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Page 1: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

Plant Cells and their Organelles

Plant Cells and their OrganellesEditEd By

William V Dashek

and

Gurbachan S Miglani

This edition first published 2017 copy 2017 by John Wiley amp Sons Ltd

Registered OfficeJohn Wiley amp Sons Ltd The Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

Editorial Offices9600 Garsington Road Oxford OX4 2DQ UKThe Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

For details of our global editorial offices for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at wwwwileycomwiley‐blackwell

The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright Designs and Patents Act 1988

All rights reserved No part of this publication may be reproduced stored in a retrieval system or transmitted in any form or by any means electronic mechanical photocopying recording or otherwise except as permitted by the UK Copyright Designs and Patents Act 1988 without the prior permission of the publisher

Designations used by companies to distinguish their products are often claimed as trademarks All brand names and product names used in this book are trade names service marks trademarks or registered trademarks of their respective owners The publisher is not associated with any product or vendor mentioned in this book

Limit of LiabilityDisclaimer of Warranty While the publisher and author(s) have used their best efforts in preparing this book they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom If professional advice or other expert assistance is required the services of a competent professional should be sought

Library of Congress Cataloging‐in‐Publication Data

Names Dashek William V editor | Miglani Gurbachan S editorTitle Plant cells and their organelles by William V Dashek Gurbachan S MiglaniDescription [Hoboken NJ] Wiley 2017 | Includes bibliographical references and indexIdentifiers LCCN 2016024724 (print) | LCCN 2016026083 (ebook) | ISBN 9780470976869 (cloth) |

ISBN 9781118924761 (pdf) | ISBN 9781118924754 (epub)Subjects LCSH Plant cells and tissuesndashTextbooksClassification LCC QK725 D36 2017 (print) | LCC QK725 (ebook) | DDC 5813ndashdc23LC record available at httpslccnlocgov2016024724

A catalogue record for this book is available from the British Library

Wiley also publishes its books in a variety of electronic formats Some content that appears in print may not be available in electronic books

Cover image copy Alan John Lander PhillipsGettyimages

Set in 9513pt Meridien by SPi Global Pondicherry India

1 2017

v

Contributors ix

Preface xi

Acknowledgments xii

1 An introduction to cells and their organelles 1

William V Dashek

Cells 1

Cell organelles ndash an introduction 6

Ion channels 10

Proton pumps 14

Water channels 14

Carriers 15

Cell death 17

References 18

Further reading 24

2 Isolation and characterization of subcellular organelles from plant cells 25

Milee Agarwal P Desai and Harish Padh

Isolation of subcellular organelles 26

Identification and characterization of isolated organelles 33

Summary 39

References 39

Further reading 41

3 Endoplasmic reticulum 42

William V Dashek

Structure 42

Chemical composition 42

Biogenesis 45

Functions 45

Posttranslational events 49

Inhibitors 53

In vitro protein synthesis 54

Other functions 54

References 54

Further reading 60

Contents

vi Contents

4 The Golgi apparatus 61

D Davis TE Wilkop and Georgia Drakakaki

The Golgi apparatus 61

Plant Golgi introduction 66

Structure and organization 69

Golgi‐mediated vesicular trafficking 71

Plant Golgi‐dependent cellular processes 74

Imaging and visualization 76

Isolation and analysis 78

Golgi genetics and genomics 81

Significance 84

Acknowledgment 85

References 85

Further reading 87

5 Microbodies 88

Robert Donaldson

Introducing peroxisomes 88

Leaf peroxisomes 89

Peroxisomes in oil seeds and pollen 91

References 107

Further reading 109

6 Microtubules intermediate filaments and actin filaments 110

William V Dashek

Microtubules 110

Intermediate filaments 113

Actin filaments (microfilaments) 116

References 119

Further reading 124

7 The mitochondrion 125

Ray J Rose Terence W‐Y Tiew and William V Dashek

Structure and dynamics 125

The mitochondrial genome 128

Comparison of the mitochondrial genome with chloroplast and nuclear

genomes 131

The mitochondrial proteome and protein import 132

Respiratory metabolite transporters 133

The electron transport chain and oxidative phosphorylation 133

The alternative electron transfer chain in plant mitochondria 139

Plant mitochondria stress responses and programmed cell death 139

Other functions of plant mitochondria 140

References 144

Further reading 145

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

ells

Uns

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aliz

ed c

ells

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uter

cov

erin

g of

var

ious

pla

nt p

arts

varia

ble

in s

hape

but

oft

en t

abul

ar

Ever

t (2

006)

Ex

ampl

es

G

uard

cel

lsSp

ecia

lized

epi

derm

al c

ells

cre

scen

t sh

aped

con

tain

chl

orop

last

s f

orm

def

ines

sto

mat

al p

ore

Will

e an

d Lu

cas

(198

4)

Su

bsid

iary

cel

lsC

ells

whi

ch s

ubte

nd t

he s

tom

atal

gua

rd c

ells

http

an

ubis

ru

acz

aM

ain

AN

ATO

MY

gua

rdce

llsh

tml

Tr

icho

mes

An

outg

row

th o

f an

epi

derm

al c

ell

can

be u

nice

llula

r or

mul

ticel

lula

rC

allo

w (2

000)

Pare

nchy

ma

cells

Isod

iam

etric

thi

n‐w

alle

d pr

imar

y ce

ll w

all

in s

ome

inst

ance

s m

ay h

ave

seco

ndar

y w

alls

not

high

ly d

iffer

entia

ted

fun

ctio

n in

pho

tosy

nthe

sis

sec

retio

n o

rgan

ic n

utrie

nt a

nd w

ater

stor

age

reg

ener

atio

n in

wou

nd h

ealin

g

Ever

t (2

006)

and

Saj

eva

and

Mau

seth

(199

1)

Ex

ampl

es

Tr

ansf

er c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

lasm

alem

ma

grea

tly e

xpan

ded

irre

gula

r ex

tens

ions

of

cell

wal

l int

o pr

otop

lasm

tra

nsfe

r di

ssol

ved

subs

tanc

es b

etw

een

adja

cent

cel

l oc

cur

in p

ith

and

cort

ex o

f st

ems

and

root

s p

hoto

synt

hetic

tis

sues

of

leav

es f

lesh

of

succ

ulen

t fr

uits

endo

sper

m o

f se

eds

Das

hek

et a

l (1

971)

and

Off

ler

et a

l

(200

3)

Col

lenc

hym

a ce

llsLa

mel

lar

or p

late

col

lenc

hym

a w

ith t

hick

enin

gs o

n th

e ta

ngen

tial w

alls

Ang

ular

col

lenc

hym

a w

ith t

hick

enin

gs a

roun

d th

e ce

ll w

alls

Pres

ent

in a

eria

l por

tions

of

the

plan

t bo

dy

Vasc

ular

cel

lsEv

ert

(200

6)

Ph

loem

Siev

e ce

lls

Siev

e el

emen

ts

Com

pani

on c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

osse

ss n

umer

ous

plas

mod

esm

atal

con

nect

ions

Opa

rka

and

Turg

eon

(199

9)

Alb

umin

ous

cells

in

gym

nosp

erm

s

Abs

ence

of

star

ch c

ytop

lasm

ic b

ridge

s w

ith s

ieve

cel

ls d

ense

pro

topl

asm

abu

ndan

ce o

f

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ighl

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nden

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euch

rom

atin

and

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itoch

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ia

Alo

si a

nd A

lfier

i (19

72) a

nd

Saut

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(19

76)

X

ylem

Trac

heid

s

Vess

els

Long

tap

erin

g ce

ll w

ith li

gnifi

ed s

econ

dary

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l thi

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ings

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e pi

ts in

wal

ls d

evoi

d of

prot

opla

sm a

t m

atur

ity n

ot a

s sp

ecia

lized

as

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els

wid

espr

ead

Tyre

e an

d Zi

mm

erm

an (2

002)

Fuku

da (2

004)

and

Eve

rt (2

006)

(Con

tinue

d )

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Spec

ializ

ed c

ells

ndash H

ydat

hode

s

(mod

ified

par

ts o

f le

aves

and

leaf

tip

s or

mar

gins

)

Con

sist

of

term

inal

tra

chei

ds e

pith

em t

hin‐

wal

led

chlo

ropl

ast‐

defic

ient

cel

ls a

she

ath

with

wat

er p

ores

gut

tatio

n di

scha

rge

of li

quid

con

tain

ing

vario

us d

isso

lved

sol

utes

fro

m a

leaf

rsquos

inte

rior

Lers

ten

and

Cur

tis (1

996)

http

sw

ww

bio

sciu

texa

sed

u a

nd

Mae

da a

nd M

aeda

(198

8)

Latic

ifer

cells

Cel

ls o

r a

serie

s of

cel

ls w

hich

pro

duce

late

xFa

hn (1

990)

Pic

kard

(200

8) a

nd

Botw

ebu

wsp

Edu

Si

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-cel

led

C

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und

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ulat

ed

Uni

on o

f ce

lls c

ompo

und

in o

rigin

and

con

sist

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itudi

nal c

hain

s of

cel

ls w

all s

epar

atin

g

cells

rem

ain

inta

ct c

an b

ecom

e pe

rfor

ated

or

entir

ely

rem

oved

Salt

glan

dsM

odifi

ed t

richo

mes

tw

o‐ce

lled

and

posi

tione

d fla

t on

the

sur

face

in r

ows

para

llel t

o th

e le

af

surf

ace

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ur in

Poa

ceae

Ever

t (2

006)

Tan

et

al (

2010

) O

ross

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l (1

985)

and

Tho

mso

n et

al

(198

8)

Cap

cel

l ndash la

rge

nucl

eus

and

expa

nded

cut

icle

Nai

doo

and

Nai

doo

(199

8)

Basa

l cel

l ndash n

umer

ous

and

larg

e ex

tens

ive

part

ition

ing

inva

gina

tions

of

plas

mal

emm

a

Nec

tarie

sFo

und

in n

ecta

rines

pro

duce

nec

tar

usua

lly a

t th

e ba

se o

f a

flow

erFa

hn (1

990)

Nic

olso

n an

d N

epi (

2005

)

and

Paiv

a (2

009)

Idio

blas

tsC

ryst

al‐c

onta

inin

g ce

llsLe

rste

n an

d H

orne

r (2

005)

Ex

ampl

e

Ra

phid

esPr

oduc

e ne

edle

‐sha

ped

crys

tals

Muc

ilage

cel

lO

ccur

in a

larg

e nu

mbe

r of

dic

ots

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mon

in c

erta

in c

acti

slim

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ucila

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nts

evap

orat

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of w

ater

by

bind

ing

to w

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dic

tyos

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as

in s

eed

coat

s c

ell w

alls

are

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sic

and

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w

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l

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Spec

ializ

ed c

ells

app

ear

like

larg

e pa

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an o

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ascu

lar

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l wal

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ct la

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aft

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l lay

er

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osite

d

Rode

las

et a

l (2

008)

htt

pbr

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ica

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and

Ler

sten

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al (

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)

Dru

ses

Sphe

rical

agg

rega

tes

of p

rism

atic

cry

stal

sLe

rste

n an

d H

orne

r (2

005)

Cel

ls in

non

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s

Br

yoph

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Gem

mae

One

to

man

y ce

llsht

tp

build

ingt

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ide

com

fac

ulty

pgda

viso

nbr

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gy_l

inks

htm

Hyd

roid

sW

ater

‐con

duct

ing

cells

http

w

ww

Bio

logy

‐onl

ine

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Lept

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ndash P

terid

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tes

Org

anic

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poun

d‐co

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ting

cells

spo

roge

nous

cel

ls p

rese

nt in

spo

rang

ia o

f so

ri

Tab

le 1

1 (

Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 2: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

Plant Cells and their OrganellesEditEd By

William V Dashek

and

Gurbachan S Miglani

This edition first published 2017 copy 2017 by John Wiley amp Sons Ltd

Registered OfficeJohn Wiley amp Sons Ltd The Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

Editorial Offices9600 Garsington Road Oxford OX4 2DQ UKThe Atrium Southern Gate Chichester West Sussex PO19 8SQ UK

For details of our global editorial offices for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at wwwwileycomwiley‐blackwell

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Limit of LiabilityDisclaimer of Warranty While the publisher and author(s) have used their best efforts in preparing this book they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose It is sold on the understanding that the publisher is not engaged in rendering professional services and neither the publisher nor the author shall be liable for damages arising herefrom If professional advice or other expert assistance is required the services of a competent professional should be sought

Library of Congress Cataloging‐in‐Publication Data

Names Dashek William V editor | Miglani Gurbachan S editorTitle Plant cells and their organelles by William V Dashek Gurbachan S MiglaniDescription [Hoboken NJ] Wiley 2017 | Includes bibliographical references and indexIdentifiers LCCN 2016024724 (print) | LCCN 2016026083 (ebook) | ISBN 9780470976869 (cloth) |

ISBN 9781118924761 (pdf) | ISBN 9781118924754 (epub)Subjects LCSH Plant cells and tissuesndashTextbooksClassification LCC QK725 D36 2017 (print) | LCC QK725 (ebook) | DDC 5813ndashdc23LC record available at httpslccnlocgov2016024724

A catalogue record for this book is available from the British Library

Wiley also publishes its books in a variety of electronic formats Some content that appears in print may not be available in electronic books

Cover image copy Alan John Lander PhillipsGettyimages

Set in 9513pt Meridien by SPi Global Pondicherry India

1 2017

v

Contributors ix

Preface xi

Acknowledgments xii

1 An introduction to cells and their organelles 1

William V Dashek

Cells 1

Cell organelles ndash an introduction 6

Ion channels 10

Proton pumps 14

Water channels 14

Carriers 15

Cell death 17

References 18

Further reading 24

2 Isolation and characterization of subcellular organelles from plant cells 25

Milee Agarwal P Desai and Harish Padh

Isolation of subcellular organelles 26

Identification and characterization of isolated organelles 33

Summary 39

References 39

Further reading 41

3 Endoplasmic reticulum 42

William V Dashek

Structure 42

Chemical composition 42

Biogenesis 45

Functions 45

Posttranslational events 49

Inhibitors 53

In vitro protein synthesis 54

Other functions 54

References 54

Further reading 60

Contents

vi Contents

4 The Golgi apparatus 61

D Davis TE Wilkop and Georgia Drakakaki

The Golgi apparatus 61

Plant Golgi introduction 66

Structure and organization 69

Golgi‐mediated vesicular trafficking 71

Plant Golgi‐dependent cellular processes 74

Imaging and visualization 76

Isolation and analysis 78

Golgi genetics and genomics 81

Significance 84

Acknowledgment 85

References 85

Further reading 87

5 Microbodies 88

Robert Donaldson

Introducing peroxisomes 88

Leaf peroxisomes 89

Peroxisomes in oil seeds and pollen 91

References 107

Further reading 109

6 Microtubules intermediate filaments and actin filaments 110

William V Dashek

Microtubules 110

Intermediate filaments 113

Actin filaments (microfilaments) 116

References 119

Further reading 124

7 The mitochondrion 125

Ray J Rose Terence W‐Y Tiew and William V Dashek

Structure and dynamics 125

The mitochondrial genome 128

Comparison of the mitochondrial genome with chloroplast and nuclear

genomes 131

The mitochondrial proteome and protein import 132

Respiratory metabolite transporters 133

The electron transport chain and oxidative phosphorylation 133

The alternative electron transfer chain in plant mitochondria 139

Plant mitochondria stress responses and programmed cell death 139

Other functions of plant mitochondria 140

References 144

Further reading 145

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

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ypes

Cel

l typ

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4)

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d )

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Tab

le 1

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)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 3: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

This edition first published 2017 copy 2017 by John Wiley amp Sons Ltd

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Library of Congress Cataloging‐in‐Publication Data

Names Dashek William V editor | Miglani Gurbachan S editorTitle Plant cells and their organelles by William V Dashek Gurbachan S MiglaniDescription [Hoboken NJ] Wiley 2017 | Includes bibliographical references and indexIdentifiers LCCN 2016024724 (print) | LCCN 2016026083 (ebook) | ISBN 9780470976869 (cloth) |

ISBN 9781118924761 (pdf) | ISBN 9781118924754 (epub)Subjects LCSH Plant cells and tissuesndashTextbooksClassification LCC QK725 D36 2017 (print) | LCC QK725 (ebook) | DDC 5813ndashdc23LC record available at httpslccnlocgov2016024724

A catalogue record for this book is available from the British Library

Wiley also publishes its books in a variety of electronic formats Some content that appears in print may not be available in electronic books

Cover image copy Alan John Lander PhillipsGettyimages

Set in 9513pt Meridien by SPi Global Pondicherry India

1 2017

v

Contributors ix

Preface xi

Acknowledgments xii

1 An introduction to cells and their organelles 1

William V Dashek

Cells 1

Cell organelles ndash an introduction 6

Ion channels 10

Proton pumps 14

Water channels 14

Carriers 15

Cell death 17

References 18

Further reading 24

2 Isolation and characterization of subcellular organelles from plant cells 25

Milee Agarwal P Desai and Harish Padh

Isolation of subcellular organelles 26

Identification and characterization of isolated organelles 33

Summary 39

References 39

Further reading 41

3 Endoplasmic reticulum 42

William V Dashek

Structure 42

Chemical composition 42

Biogenesis 45

Functions 45

Posttranslational events 49

Inhibitors 53

In vitro protein synthesis 54

Other functions 54

References 54

Further reading 60

Contents

vi Contents

4 The Golgi apparatus 61

D Davis TE Wilkop and Georgia Drakakaki

The Golgi apparatus 61

Plant Golgi introduction 66

Structure and organization 69

Golgi‐mediated vesicular trafficking 71

Plant Golgi‐dependent cellular processes 74

Imaging and visualization 76

Isolation and analysis 78

Golgi genetics and genomics 81

Significance 84

Acknowledgment 85

References 85

Further reading 87

5 Microbodies 88

Robert Donaldson

Introducing peroxisomes 88

Leaf peroxisomes 89

Peroxisomes in oil seeds and pollen 91

References 107

Further reading 109

6 Microtubules intermediate filaments and actin filaments 110

William V Dashek

Microtubules 110

Intermediate filaments 113

Actin filaments (microfilaments) 116

References 119

Further reading 124

7 The mitochondrion 125

Ray J Rose Terence W‐Y Tiew and William V Dashek

Structure and dynamics 125

The mitochondrial genome 128

Comparison of the mitochondrial genome with chloroplast and nuclear

genomes 131

The mitochondrial proteome and protein import 132

Respiratory metabolite transporters 133

The electron transport chain and oxidative phosphorylation 133

The alternative electron transfer chain in plant mitochondria 139

Plant mitochondria stress responses and programmed cell death 139

Other functions of plant mitochondria 140

References 144

Further reading 145

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

ells

Uns

peci

aliz

ed c

ells

one

laye

r of

cel

ls in

thi

ckne

ss o

uter

cov

erin

g of

var

ious

pla

nt p

arts

varia

ble

in s

hape

but

oft

en t

abul

ar

Ever

t (2

006)

Ex

ampl

es

G

uard

cel

lsSp

ecia

lized

epi

derm

al c

ells

cre

scen

t sh

aped

con

tain

chl

orop

last

s f

orm

def

ines

sto

mat

al p

ore

Will

e an

d Lu

cas

(198

4)

Su

bsid

iary

cel

lsC

ells

whi

ch s

ubte

nd t

he s

tom

atal

gua

rd c

ells

http

an

ubis

ru

acz

aM

ain

AN

ATO

MY

gua

rdce

llsh

tml

Tr

icho

mes

An

outg

row

th o

f an

epi

derm

al c

ell

can

be u

nice

llula

r or

mul

ticel

lula

rC

allo

w (2

000)

Pare

nchy

ma

cells

Isod

iam

etric

thi

n‐w

alle

d pr

imar

y ce

ll w

all

in s

ome

inst

ance

s m

ay h

ave

seco

ndar

y w

alls

not

high

ly d

iffer

entia

ted

fun

ctio

n in

pho

tosy

nthe

sis

sec

retio

n o

rgan

ic n

utrie

nt a

nd w

ater

stor

age

reg

ener

atio

n in

wou

nd h

ealin

g

Ever

t (2

006)

and

Saj

eva

and

Mau

seth

(199

1)

Ex

ampl

es

Tr

ansf

er c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

lasm

alem

ma

grea

tly e

xpan

ded

irre

gula

r ex

tens

ions

of

cell

wal

l int

o pr

otop

lasm

tra

nsfe

r di

ssol

ved

subs

tanc

es b

etw

een

adja

cent

cel

l oc

cur

in p

ith

and

cort

ex o

f st

ems

and

root

s p

hoto

synt

hetic

tis

sues

of

leav

es f

lesh

of

succ

ulen

t fr

uits

endo

sper

m o

f se

eds

Das

hek

et a

l (1

971)

and

Off

ler

et a

l

(200

3)

Col

lenc

hym

a ce

llsLa

mel

lar

or p

late

col

lenc

hym

a w

ith t

hick

enin

gs o

n th

e ta

ngen

tial w

alls

Ang

ular

col

lenc

hym

a w

ith t

hick

enin

gs a

roun

d th

e ce

ll w

alls

Pres

ent

in a

eria

l por

tions

of

the

plan

t bo

dy

Vasc

ular

cel

lsEv

ert

(200

6)

Ph

loem

Siev

e ce

lls

Siev

e el

emen

ts

Com

pani

on c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

osse

ss n

umer

ous

plas

mod

esm

atal

con

nect

ions

Opa

rka

and

Turg

eon

(199

9)

Alb

umin

ous

cells

in

gym

nosp

erm

s

Abs

ence

of

star

ch c

ytop

lasm

ic b

ridge

s w

ith s

ieve

cel

ls d

ense

pro

topl

asm

abu

ndan

ce o

f

poly

som

es h

ighl

y co

nden

sed

euch

rom

atin

and

abu

ndan

t m

itoch

ondr

ia

Alo

si a

nd A

lfier

i (19

72) a

nd

Saut

er e

t al

(19

76)

X

ylem

Trac

heid

s

Vess

els

Long

tap

erin

g ce

ll w

ith li

gnifi

ed s

econ

dary

wal

l thi

cken

ings

can

hav

e pi

ts in

wal

ls d

evoi

d of

prot

opla

sm a

t m

atur

ity n

ot a

s sp

ecia

lized

as

vess

els

wid

espr

ead

Tyre

e an

d Zi

mm

erm

an (2

002)

Fuku

da (2

004)

and

Eve

rt (2

006)

(Con

tinue

d )

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Spec

ializ

ed c

ells

ndash H

ydat

hode

s

(mod

ified

par

ts o

f le

aves

and

leaf

tip

s or

mar

gins

)

Con

sist

of

term

inal

tra

chei

ds e

pith

em t

hin‐

wal

led

chlo

ropl

ast‐

defic

ient

cel

ls a

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 4: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

v

Contributors ix

Preface xi

Acknowledgments xii

1 An introduction to cells and their organelles 1

William V Dashek

Cells 1

Cell organelles ndash an introduction 6

Ion channels 10

Proton pumps 14

Water channels 14

Carriers 15

Cell death 17

References 18

Further reading 24

2 Isolation and characterization of subcellular organelles from plant cells 25

Milee Agarwal P Desai and Harish Padh

Isolation of subcellular organelles 26

Identification and characterization of isolated organelles 33

Summary 39

References 39

Further reading 41

3 Endoplasmic reticulum 42

William V Dashek

Structure 42

Chemical composition 42

Biogenesis 45

Functions 45

Posttranslational events 49

Inhibitors 53

In vitro protein synthesis 54

Other functions 54

References 54

Further reading 60

Contents

vi Contents

4 The Golgi apparatus 61

D Davis TE Wilkop and Georgia Drakakaki

The Golgi apparatus 61

Plant Golgi introduction 66

Structure and organization 69

Golgi‐mediated vesicular trafficking 71

Plant Golgi‐dependent cellular processes 74

Imaging and visualization 76

Isolation and analysis 78

Golgi genetics and genomics 81

Significance 84

Acknowledgment 85

References 85

Further reading 87

5 Microbodies 88

Robert Donaldson

Introducing peroxisomes 88

Leaf peroxisomes 89

Peroxisomes in oil seeds and pollen 91

References 107

Further reading 109

6 Microtubules intermediate filaments and actin filaments 110

William V Dashek

Microtubules 110

Intermediate filaments 113

Actin filaments (microfilaments) 116

References 119

Further reading 124

7 The mitochondrion 125

Ray J Rose Terence W‐Y Tiew and William V Dashek

Structure and dynamics 125

The mitochondrial genome 128

Comparison of the mitochondrial genome with chloroplast and nuclear

genomes 131

The mitochondrial proteome and protein import 132

Respiratory metabolite transporters 133

The electron transport chain and oxidative phosphorylation 133

The alternative electron transfer chain in plant mitochondria 139

Plant mitochondria stress responses and programmed cell death 139

Other functions of plant mitochondria 140

References 144

Further reading 145

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

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erm

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ells

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tain

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orm

def

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ore

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d Lu

cas

(198

4)

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bsid

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ch s

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nd t

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etric

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late

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a w

ith t

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n th

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l por

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loem

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e ce

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ts

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Spec

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lls p

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ous

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mod

esm

atal

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Opa

rka

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eon

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9)

Alb

umin

ous

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erm

s

Abs

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ch c

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ith s

ieve

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and

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(19

76)

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Trac

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Vess

els

Long

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g ce

ll w

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gnifi

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d of

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ead

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e an

d Zi

mm

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an (2

002)

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da (2

004)

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rt (2

006)

(Con

tinue

d )

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Spec

ializ

ed c

ells

ndash H

ydat

hode

s

(mod

ified

par

ts o

f le

aves

and

leaf

tip

s or

mar

gins

)

Con

sist

of

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inal

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chei

ds e

pith

em t

hin‐

wal

led

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ropl

ast‐

defic

ient

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ls a

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with

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er p

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n di

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rge

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tain

ing

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us d

isso

lved

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utes

fro

m a

leaf

rsquos

inte

rior

Lers

ten

and

Cur

tis (1

996)

http

sw

ww

bio

sciu

texa

sed

u a

nd

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da a

nd M

aeda

(198

8)

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ifer

cells

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ls o

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s of

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ls w

hich

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duce

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hn (1

990)

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kard

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nd

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ngle

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ompo

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ulat

ed

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on o

f ce

lls c

ompo

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in o

rigin

and

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sist

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itudi

nal c

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ls w

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ain

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ct c

an b

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e pe

rfor

ated

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entir

ely

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oved

Salt

glan

dsM

odifi

ed t

richo

mes

tw

o‐ce

lled

and

posi

tione

d fla

t on

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face

in r

ows

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llel t

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surf

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ur in

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ceae

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t (2

006)

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et

al (

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) O

ross

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l (1

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mso

n et

al

(198

8)

Cap

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l ndash la

rge

nucl

eus

and

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nded

cut

icle

Nai

doo

and

Nai

doo

(199

8)

Basa

l cel

l ndash n

umer

ous

and

larg

e ex

tens

ive

part

ition

ing

inva

gina

tions

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emm

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und

in n

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rines

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duce

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tar

usua

lly a

t th

e ba

se o

f a

flow

erFa

hn (1

990)

Nic

olso

n an

d N

epi (

2005

)

and

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a (2

009)

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blas

tsC

ryst

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onta

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g ce

llsLe

rste

n an

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orne

r (2

005)

Ex

ampl

e

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phid

esPr

oduc

e ne

edle

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ped

crys

tals

Muc

ilage

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lO

ccur

in a

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e nu

mbe

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nts

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omes

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ell w

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d

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l

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ovsk

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ear

like

larg

e pa

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a ce

lls c

an o

ccur

in v

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lar

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nd t

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l wal

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ct la

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r (2

005)

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ide

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roid

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ater

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w

ww

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logy

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ine

org

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ndash P

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tes

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anic

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roge

nous

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ls p

rese

nt in

spo

rang

ia o

f so

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Tab

le 1

1 (

Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 5: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

vi Contents

4 The Golgi apparatus 61

D Davis TE Wilkop and Georgia Drakakaki

The Golgi apparatus 61

Plant Golgi introduction 66

Structure and organization 69

Golgi‐mediated vesicular trafficking 71

Plant Golgi‐dependent cellular processes 74

Imaging and visualization 76

Isolation and analysis 78

Golgi genetics and genomics 81

Significance 84

Acknowledgment 85

References 85

Further reading 87

5 Microbodies 88

Robert Donaldson

Introducing peroxisomes 88

Leaf peroxisomes 89

Peroxisomes in oil seeds and pollen 91

References 107

Further reading 109

6 Microtubules intermediate filaments and actin filaments 110

William V Dashek

Microtubules 110

Intermediate filaments 113

Actin filaments (microfilaments) 116

References 119

Further reading 124

7 The mitochondrion 125

Ray J Rose Terence W‐Y Tiew and William V Dashek

Structure and dynamics 125

The mitochondrial genome 128

Comparison of the mitochondrial genome with chloroplast and nuclear

genomes 131

The mitochondrial proteome and protein import 132

Respiratory metabolite transporters 133

The electron transport chain and oxidative phosphorylation 133

The alternative electron transfer chain in plant mitochondria 139

Plant mitochondria stress responses and programmed cell death 139

Other functions of plant mitochondria 140

References 144

Further reading 145

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

ells

Uns

peci

aliz

ed c

ells

one

laye

r of

cel

ls in

thi

ckne

ss o

uter

cov

erin

g of

var

ious

pla

nt p

arts

varia

ble

in s

hape

but

oft

en t

abul

ar

Ever

t (2

006)

Ex

ampl

es

G

uard

cel

lsSp

ecia

lized

epi

derm

al c

ells

cre

scen

t sh

aped

con

tain

chl

orop

last

s f

orm

def

ines

sto

mat

al p

ore

Will

e an

d Lu

cas

(198

4)

Su

bsid

iary

cel

lsC

ells

whi

ch s

ubte

nd t

he s

tom

atal

gua

rd c

ells

http

an

ubis

ru

acz

aM

ain

AN

ATO

MY

gua

rdce

llsh

tml

Tr

icho

mes

An

outg

row

th o

f an

epi

derm

al c

ell

can

be u

nice

llula

r or

mul

ticel

lula

rC

allo

w (2

000)

Pare

nchy

ma

cells

Isod

iam

etric

thi

n‐w

alle

d pr

imar

y ce

ll w

all

in s

ome

inst

ance

s m

ay h

ave

seco

ndar

y w

alls

not

high

ly d

iffer

entia

ted

fun

ctio

n in

pho

tosy

nthe

sis

sec

retio

n o

rgan

ic n

utrie

nt a

nd w

ater

stor

age

reg

ener

atio

n in

wou

nd h

ealin

g

Ever

t (2

006)

and

Saj

eva

and

Mau

seth

(199

1)

Ex

ampl

es

Tr

ansf

er c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

lasm

alem

ma

grea

tly e

xpan

ded

irre

gula

r ex

tens

ions

of

cell

wal

l int

o pr

otop

lasm

tra

nsfe

r di

ssol

ved

subs

tanc

es b

etw

een

adja

cent

cel

l oc

cur

in p

ith

and

cort

ex o

f st

ems

and

root

s p

hoto

synt

hetic

tis

sues

of

leav

es f

lesh

of

succ

ulen

t fr

uits

endo

sper

m o

f se

eds

Das

hek

et a

l (1

971)

and

Off

ler

et a

l

(200

3)

Col

lenc

hym

a ce

llsLa

mel

lar

or p

late

col

lenc

hym

a w

ith t

hick

enin

gs o

n th

e ta

ngen

tial w

alls

Ang

ular

col

lenc

hym

a w

ith t

hick

enin

gs a

roun

d th

e ce

ll w

alls

Pres

ent

in a

eria

l por

tions

of

the

plan

t bo

dy

Vasc

ular

cel

lsEv

ert

(200

6)

Ph

loem

Siev

e ce

lls

Siev

e el

emen

ts

Com

pani

on c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

osse

ss n

umer

ous

plas

mod

esm

atal

con

nect

ions

Opa

rka

and

Turg

eon

(199

9)

Alb

umin

ous

cells

in

gym

nosp

erm

s

Abs

ence

of

star

ch c

ytop

lasm

ic b

ridge

s w

ith s

ieve

cel

ls d

ense

pro

topl

asm

abu

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 6: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

Contents vii

8 Nucleus 146

Yogesh Vikal and Dasmeet Kaur

Structural organization of the NE 147

Nuclear pores 152

The nucleolus 157

Chromatin and chromosomes 165

DNA structure 170

DNA replication 173

RNA structure function and synthesis 176

Nucleocytoplasmic transport nuclear import and nuclear export 183

The dynamics of NE biogenesis during mitosis 188

The dynamics of nuclear pore complex biogenesis 196

Cell cycle control 200

Summary 205

References 206

Further reading 207

9 Plant cell walls 209

James E Bidlack and William V Dashek

Introduction 209

Structure 209

Biosynthesis 216

Chemical composition 217

Biogenesis 222

Function 225

References 231

Further reading 238

10 Plastid structure and genomics 239

Gurbachan S Miglani

Plastid structure 239

Different forms of plastids 240

Plastid stromules 248

Chlorophyll biosynthesis 248

Plastid genomics 250

Sequenced plastomes 253

Promiscuous DNA 258

Plastid genome organization 260

Plastid gene organization expression and regulation 265

Systems biology approach in understanding chloroplast development 269

Chloroplast genetic engineering 284

Recent trends in chloroplast research 289

Summary 293

References 294

Further reading 299

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

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Uns

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ecia

lized

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al c

ells

cre

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t sh

aped

con

tain

chl

orop

last

s f

orm

def

ines

sto

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al p

ore

Will

e an

d Lu

cas

(198

4)

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bsid

iary

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lsC

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ch s

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nd t

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atal

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rd c

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http

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etric

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ome

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lar

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late

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lenc

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a w

ith t

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enin

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n th

e ta

ngen

tial w

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l por

tions

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t bo

dy

Vasc

ular

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6)

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loem

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e ce

lls

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emen

ts

Com

pani

on c

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Spec

ializ

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lls p

osse

ss n

umer

ous

plas

mod

esm

atal

con

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ions

Opa

rka

and

Turg

eon

(199

9)

Alb

umin

ous

cells

in

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nosp

erm

s

Abs

ence

of

star

ch c

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s w

ith s

ieve

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ls d

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ighl

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and

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Alo

si a

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i (19

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nd

Saut

er e

t al

(19

76)

X

ylem

Trac

heid

s

Vess

els

Long

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erin

g ce

ll w

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gnifi

ed s

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dary

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l thi

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ings

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e pi

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wal

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d of

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s sp

ecia

lized

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espr

ead

Tyre

e an

d Zi

mm

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an (2

002)

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da (2

004)

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rt (2

006)

(Con

tinue

d )

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Spec

ializ

ed c

ells

ndash H

ydat

hode

s

(mod

ified

par

ts o

f le

aves

and

leaf

tip

s or

mar

gins

)

Con

sist

of

term

inal

tra

chei

ds e

pith

em t

hin‐

wal

led

chlo

ropl

ast‐

defic

ient

cel

ls a

she

ath

with

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er p

ores

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tatio

n di

scha

rge

of li

quid

con

tain

ing

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us d

isso

lved

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utes

fro

m a

leaf

rsquos

inte

rior

Lers

ten

and

Cur

tis (1

996)

http

sw

ww

bio

sciu

texa

sed

u a

nd

Mae

da a

nd M

aeda

(198

8)

Latic

ifer

cells

Cel

ls o

r a

serie

s of

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ls w

hich

pro

duce

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xFa

hn (1

990)

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kard

(200

8) a

nd

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ebu

wsp

Edu

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ngle

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led

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ompo

und

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ulat

ed

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on o

f ce

lls c

ompo

und

in o

rigin

and

con

sist

of

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itudi

nal c

hain

s of

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ls w

all s

epar

atin

g

cells

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ain

inta

ct c

an b

ecom

e pe

rfor

ated

or

entir

ely

rem

oved

Salt

glan

dsM

odifi

ed t

richo

mes

tw

o‐ce

lled

and

posi

tione

d fla

t on

the

sur

face

in r

ows

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llel t

o th

e le

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surf

ace

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ur in

Poa

ceae

Ever

t (2

006)

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et

al (

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) O

ross

et a

l (1

985)

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Tho

mso

n et

al

(198

8)

Cap

cel

l ndash la

rge

nucl

eus

and

expa

nded

cut

icle

Nai

doo

and

Nai

doo

(199

8)

Basa

l cel

l ndash n

umer

ous

and

larg

e ex

tens

ive

part

ition

ing

inva

gina

tions

of

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mal

emm

a

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tarie

sFo

und

in n

ecta

rines

pro

duce

nec

tar

usua

lly a

t th

e ba

se o

f a

flow

erFa

hn (1

990)

Nic

olso

n an

d N

epi (

2005

)

and

Paiv

a (2

009)

Idio

blas

tsC

ryst

al‐c

onta

inin

g ce

llsLe

rste

n an

d H

orne

r (2

005)

Ex

ampl

e

Ra

phid

esPr

oduc

e ne

edle

‐sha

ped

crys

tals

Muc

ilage

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lO

ccur

in a

larg

e nu

mbe

r of

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ots

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mon

in c

erta

in c

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slim

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ucila

ge p

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nts

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orat

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of w

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by

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ing

to w

ater

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ll w

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omes

pro

duce

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in s

eed

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s c

ell w

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are

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lulo

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d

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uetl

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ry9

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l

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) and

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ovsk

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l (2

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ed c

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app

ear

like

larg

e pa

renc

hym

a ce

lls c

an o

ccur

in v

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lar

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nd t

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l wal

l has

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ee d

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ct la

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l lay

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osite

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et a

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ica

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tes

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atic

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rste

n an

d H

orne

r (2

005)

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ls in

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erm

s

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ytes

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mae

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y ce

llsht

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ide

com

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ulty

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nbr

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gy_l

inks

htm

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roid

sW

ater

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duct

ing

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http

w

ww

Bio

logy

‐onl

ine

org

Lept

oids

ndash P

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tes

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anic

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poun

d‐co

nduc

ting

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spo

roge

nous

cel

ls p

rese

nt in

spo

rang

ia o

f so

ri

Tab

le 1

1 (

Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 7: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

viii Contents

11 Photosynthesis 300

J Kenneth Hoober

Introduction 300

Evolution of photosynthesis 301

Development of the chloroplast 310

Absorption of light energy 317

Generation of end products 324

Distribution of the photosystems in thylakoid membranes 329

Photoinhibition damage and repair of the PS II reaction center 332

Protection of PS II by carotenoids 332

Incorporation of carbon as CO2 into carbohydrate 334

End products of carbon assimilation 346

Conclusions for the reactions of photosynthesis 348

References 348

Further reading 350

12 Vacuoles and protein bodies 351

William V Dashek and Amy M Clore

Vacuoles 351

PBs and other protein storage compartments 359

References 365

Further reading 370

13 Systems biology in plant cells and their organelles 371

Rajdeep Kaur Grewal Saptarshi Sinha and Soumen Roy

Systems biologymdashldquoomicsrdquo 371

Genomics 373

Lipidomics 378

Metabolomics 380

Proteomics 382

Transcriptomics 384

Synthetic biology 386

Acknowledgments 388

References 389

Further reading 391

Appendix A 392

Appendix B 400

Appendix C 403

Index 407

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

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Uns

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aliz

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ble

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Ever

t (2

006)

Ex

ampl

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G

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ecia

lized

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derm

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ells

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scen

t sh

aped

con

tain

chl

orop

last

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orm

def

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mat

al p

ore

Will

e an

d Lu

cas

(198

4)

Su

bsid

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lsC

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ch s

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nd t

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atal

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an

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be u

nice

llula

r or

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ticel

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allo

w (2

000)

Pare

nchy

ma

cells

Isod

iam

etric

thi

n‐w

alle

d pr

imar

y ce

ll w

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in s

ome

inst

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ay h

ave

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Tr

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Spec

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aren

chym

a ce

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lasm

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grea

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r ex

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of

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wal

l int

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otop

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nsfe

r di

ssol

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subs

tanc

es b

etw

een

adja

cent

cel

l oc

cur

in p

ith

and

cort

ex o

f st

ems

and

root

s p

hoto

synt

hetic

tis

sues

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leav

es f

lesh

of

succ

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uits

endo

sper

m o

f se

eds

Das

hek

et a

l (1

971)

and

Off

ler

et a

l

(200

3)

Col

lenc

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mel

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or p

late

col

lenc

hym

a w

ith t

hick

enin

gs o

n th

e ta

ngen

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alls

Ang

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Pres

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Vasc

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loem

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ts

Com

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Opa

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Alb

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Abs

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and

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(19

76)

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Trac

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els

Long

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espr

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da (2

004)

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rt (2

006)

(Con

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l typ

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Ref

eren

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ed c

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(mod

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and

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gins

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Con

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ls a

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er p

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n di

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us d

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utes

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rsquos

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rior

Lers

ten

and

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tis (1

996)

http

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bio

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texa

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u a

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terid

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tes

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anic

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ls p

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nt in

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ia o

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Tab

le 1

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Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 8: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

ix

Milee AgarwalScientist

Pharmacology and Toxicology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

James E BidlackProfessor of Biology and CURE‐STEM

Scholar

Department of Biology

University of Central Oklahoma

Edmond OK USA

Amy M CloreProfessor of Biology

Division of Natural Sciences

New College of Florida

Sarasota FL USA

William V DashekRetired Faculty

Adult Degree Program

Mary Baldwin College

Staunton VA USA

D DavisGraduate Student

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

P DesaiScientist

Cellular and Molecular Biology Department

BV Patel PERD Centre

Ahmedabad Gujarat India

Robert DonaldsonProfessor

Department of Biological Sciences

George Washington University

Washington DC USA

Georgia DrakakakiAssociate Professor

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

Rajdeep Kaur GrewalSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

J Kenneth HooberProfessor Emeritus

School of Life Sciences

Center for Photosynthesis

Arizona State University

Tempe AZ USA

Dasmeet KaurResearch Assistant

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Gurbachan S MiglaniVisiting Professor

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

Contributors

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

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Tab

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 9: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

x Contributors

Harish PadhVice-Chancellor

Sardar Patel University

Vallabh Vidyanagar Anand

Gujarat India

Ray J RoseEmeritus Professor

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Soumen RoyAssociate Professor

Department of Physics

Bose Institute

Kolkata India

Saptarshi SinhaSenior Research Fellow

Department of Physics

Bose Institute

Kolkata India

Terence W‐Y TiewGraduate Student

Center of Excellence for Integrative

Legume Research

School of Environmental and Life Sciences

The University of Newcastle

Callaghan New South Wales Australia

Yogesh VikalSenior Geneticist

School of Agricultural Biotechnology

Punjab Agricultural University

Ludhiana India

TE WilkopSenior Project Scientist

Hellman Fellow Plant Sciences

Department of Plant Sciences

University of California Davis

Davis CA USA

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

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ypes

Cel

l typ

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acte

rist

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Ref

eren

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Epid

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Uns

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cas

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4)

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Vasc

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lls p

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ous

plas

mod

esm

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Opa

rka

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eon

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Alb

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Abs

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Alo

si a

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lfier

i (19

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er e

t al

(19

76)

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Trac

heid

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Vess

els

Long

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g ce

ll w

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gnifi

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ead

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002)

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004)

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rt (2

006)

(Con

tinue

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Cel

l typ

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Ref

eren

ces

Spec

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ed c

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ndash H

ydat

hode

s

(mod

ified

par

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f le

aves

and

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gins

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Con

sist

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us d

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rior

Lers

ten

and

Cur

tis (1

996)

http

sw

ww

bio

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texa

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u a

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8)

Latic

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990)

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e pe

rfor

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ely

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oved

Salt

glan

dsM

odifi

ed t

richo

mes

tw

o‐ce

lled

and

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tione

d fla

t on

the

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face

in r

ows

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llel t

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ur in

Poa

ceae

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t (2

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et

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ross

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8)

Cap

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nded

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icle

Nai

doo

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(199

8)

Basa

l cel

l ndash n

umer

ous

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larg

e ex

tens

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gina

tions

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e ba

se o

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005)

Ex

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e

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phid

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crys

tals

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lO

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e nu

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renc

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a ce

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an o

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005)

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ls p

rese

nt in

spo

rang

ia o

f so

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Tab

le 1

1 (

Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 10: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

xi

Plant Cells and their Organelles is an advanced textbook to enhance the plant

biology studentrsquos knowledge of the structure and function of plant cells and their

organelles The book assumes that the student has had introductory courses in

plant science and chemistry The book emphasizes the research literature in

plant cell biology concerning cell and organellar structure However the litera-

ture from plant physiology molecular genetics and biochemistry has been

utilized to augment the discussions of cell and organellar function

Preface

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

ells

Uns

peci

aliz

ed c

ells

one

laye

r of

cel

ls in

thi

ckne

ss o

uter

cov

erin

g of

var

ious

pla

nt p

arts

varia

ble

in s

hape

but

oft

en t

abul

ar

Ever

t (2

006)

Ex

ampl

es

G

uard

cel

lsSp

ecia

lized

epi

derm

al c

ells

cre

scen

t sh

aped

con

tain

chl

orop

last

s f

orm

def

ines

sto

mat

al p

ore

Will

e an

d Lu

cas

(198

4)

Su

bsid

iary

cel

lsC

ells

whi

ch s

ubte

nd t

he s

tom

atal

gua

rd c

ells

http

an

ubis

ru

acz

aM

ain

AN

ATO

MY

gua

rdce

llsh

tml

Tr

icho

mes

An

outg

row

th o

f an

epi

derm

al c

ell

can

be u

nice

llula

r or

mul

ticel

lula

rC

allo

w (2

000)

Pare

nchy

ma

cells

Isod

iam

etric

thi

n‐w

alle

d pr

imar

y ce

ll w

all

in s

ome

inst

ance

s m

ay h

ave

seco

ndar

y w

alls

not

high

ly d

iffer

entia

ted

fun

ctio

n in

pho

tosy

nthe

sis

sec

retio

n o

rgan

ic n

utrie

nt a

nd w

ater

stor

age

reg

ener

atio

n in

wou

nd h

ealin

g

Ever

t (2

006)

and

Saj

eva

and

Mau

seth

(199

1)

Ex

ampl

es

Tr

ansf

er c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

lasm

alem

ma

grea

tly e

xpan

ded

irre

gula

r ex

tens

ions

of

cell

wal

l int

o pr

otop

lasm

tra

nsfe

r di

ssol

ved

subs

tanc

es b

etw

een

adja

cent

cel

l oc

cur

in p

ith

and

cort

ex o

f st

ems

and

root

s p

hoto

synt

hetic

tis

sues

of

leav

es f

lesh

of

succ

ulen

t fr

uits

endo

sper

m o

f se

eds

Das

hek

et a

l (1

971)

and

Off

ler

et a

l

(200

3)

Col

lenc

hym

a ce

llsLa

mel

lar

or p

late

col

lenc

hym

a w

ith t

hick

enin

gs o

n th

e ta

ngen

tial w

alls

Ang

ular

col

lenc

hym

a w

ith t

hick

enin

gs a

roun

d th

e ce

ll w

alls

Pres

ent

in a

eria

l por

tions

of

the

plan

t bo

dy

Vasc

ular

cel

lsEv

ert

(200

6)

Ph

loem

Siev

e ce

lls

Siev

e el

emen

ts

Com

pani

on c

ells

Spec

ializ

ed p

aren

chym

a ce

lls p

osse

ss n

umer

ous

plas

mod

esm

atal

con

nect

ions

Opa

rka

and

Turg

eon

(199

9)

Alb

umin

ous

cells

in

gym

nosp

erm

s

Abs

ence

of

star

ch c

ytop

lasm

ic b

ridge

s w

ith s

ieve

cel

ls d

ense

pro

topl

asm

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 11: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

xii

Dashek is grateful to Drs WG Rosen WF Millington and DTA Lamport for

training enabling a career in teaching and research in plant biology Dashek

appreciates the grant support of the USArsquos NIH NSF DOE and USDA Forest

Service Dashek thanks Ms Katherine Mumford Ms Retha Howard and

Ms Abigail M Johnson for technical assistance in the preparation of the

manuscript

Miglani wishes to record his appreciation for Dr Darshan S Brar Honorary

Adjunct Professor School of Agricultural Biotechnology Punjab Agricultural

University Ludhiana India for his valuable technical suggestions Miglani

thanks Dr (Mrs) Parveen Chhuneja Director School of Agricultural

Biotechnology Punjab Agricultural University Ludhiana India for motivat-

ing me to prepare this volume and the Punjab Agricultural University for

providing facilities for this work

We thank the Wiley editorial staff members for their attention to detail

Acknowledgments

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

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ypes

Cel

l typ

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4)

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d )

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Tab

le 1

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)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 12: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

1

Plant Cells and their Organelles First Edition Edited by William V Dashek and Gurbachan S Miglani

copy 2017 John Wiley amp Sons Ltd Published 2017 by John Wiley amp Sons Ltd

Cells

Parenchyma chlorenchyma collenchyma and sclerenchyma are the four main

plant cell types (Figure 11 Evert 2006) Meristematic cells which occur in

shoot and root meristems are parenchyma cells Chlorenchyma cells contain

chloroplasts and lack the cell wall thickening layers of collenchyma and scleren-

chyma Certain epidermal cells can be specialized as stomata that are important

in gas exchange (Bergmann and Sack 2007) The diverse cell types (Zhang et al

2001 Yang and Liu 2007) are shown in Table 11 Photomicrographs of certain of

these cell types can be found in Evert (2006) Fahn (1990) Beck (2005) Rudall

(2007) Gunning (2009) MacAdam (2009) Wayne (2009) Beck (2009) Assmann

and Liu (2014) and Noguchi et al (2014)

How do cells ariseCells arise by cell divisions (see Chapter 8 for mitosis and meiosis) in shoot and

root (Figures 12 and 13) meristems (Table 12 Lyndon 1998 McManus and Veit

2001 Murray 2012) The shoot apex is characterized by a tunicandashcorpus organiza-

tion (Steeves and Sussex 1989) The tunica gives rise to the protoderm and its

derivative the epidermis In contrast the corpus provides the procambium which

yields the primary xylem and phloem In addition the ground tissue derives from

the corpus originating the pith and cortex Following divisions cells can differenti-

ate into tissues (Table 13) and organs of the mature plant body (Leyser and Day

2003 Sachs 2005 Dashek and Harrison 2006) The leaf primodium arises on the

apex (Micol and Hake 2003) The mature angiosperm leaf consists of palisade cells

and spongy mesophyll cells sandwiched between the upper and the lower epider-

mis (Figure 14) The epidermis possesses guard cells with associated stomata that

function in gas exchange KNOX genes affect meristem maintenance and suitable

patterning of organ formation (Hake et al 2004) In dissected leaves KNOX genes

are expressed in leaf primordia (Hake et al 2004) Hake et al (2004) suggest that

An introduction to cells and their organellesWilliam V DashekRetired Faculty Adult Degree Program Mary Baldwin College Staunton VA USA

CHapter 1

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

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ypes

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l typ

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e an

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mm

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004)

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rt (2

006)

(Con

tinue

d )

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Spec

ializ

ed c

ells

ndash H

ydat

hode

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(mod

ified

par

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f le

aves

and

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gins

)

Con

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tis (1

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l

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e pa

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Tab

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)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 13: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

2 Plant cells and their organelles

KNOX genes may be important in the diversity of leaf form Extensive discussions

of leaf development occur in Sinha (1999) Micol and Hake (2003) and Efroni

et al (2010) Under appropriate stimuli the vegetative apex can be converted to a

floral apex (Figure 15) Photoperiod (Mazumdar 2013) such as short days and

long days and combinations of the two is one such stimulus (Glover 2007

Kinmonth‐Schultz et al 2013) This induction results in the production of florigen

(Turck et al 2008) the flowering hormone (Zeevaart 2006) While early reports

suggest that florigen is an mRNA species (Huang et al 2005) a more recent inves-

tigation indicates that florigen is a protein complex (Yang et al 2007 Taoka et al

2013) Taoka et al state that florigen protein is encoded by the gene Flowering

Locus T in Arabidopsis species (Shresth et al 2014) It is believed that florigen is

induced in leaves and that it moves through the phloem to the shoot apex Plant

hormones (see Appendix A) can influence floral development (Howell 1998)

Gibberellins (Blaacutezquez et al 1998) auxins and jasmonic acid can affect petal

development In contrast auxin can influence gynoecium development The ABC

model has been proposed for regulating the development of floral parts (Soltis

et al 2006) The A gene expression is responsible for sepals while the petals are

the result of co‐expression of A and B genes The B and C genes are responsible for

stamen development and carpels require C genes In certain plants vernalization

(low temperature) can induce flowering in certain plants (Kemi et al 2013)

A diagram of the mature angiosperm plant body is presented in Figure 16 Plant

Primary phloem bres

CO

par

A

Epidermis

Cortex

Phloem

Vascularcambium

Secondaryxylem

Figure 11 Plant cell types Left parenchyma (par) and collenchyma (co) Right

sclerenchyma Source Evert (2006) Reproduced with permission of John Wiley amp Sons

Tab

le 1

1 P

lan

t ce

ll t

ypes

Cel

l typ

esC

har

acte

rist

ics

Ref

eren

ces

Epid

erm

al c

ells

Uns

peci

aliz

ed c

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r of

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ls in

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ble

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Ever

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cre

scen

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tain

chl

orop

last

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orm

def

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ore

Will

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cas

(198

4)

Su

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lsC

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whi

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nd t

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w (2

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cells

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ome

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Pres

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Abs

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Ref

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 14: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

Tab

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1 P

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An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 15: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

Cel

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esC

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acte

rist

ics

Ref

eren

ces

Spec

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ed c

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ndash H

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(mod

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and

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of

term

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of li

quid

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vario

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rsquos

inte

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Salt

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ls p

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nt in

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rang

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Tab

le 1

1 (

Con

tin

ued

)

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 16: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 5

development is discussed in Fosket (1999) Moore and Clark (1995) Greenland

(2003) Leyser and Day (2003) and Rudall (2007)

What is the composition of cellsCertain plant components exhibit polar growth for example the tip growth of

pollen tubes (Hepler et al 2001) The tubes elongate via the fusion of Golgi‐

derived vesicles with the plasmalemma and subsequent deposition of the vesi-

clesrsquo contents into the cell wall (Taylor and Hepler 1997 Parton et al 2001 and

others as reviewed in Malho (2006a 2006b)) In 2007 Dalgic and Dane (2005)

published a diagram depicting the now known tube‐tip structural elements and

physiological processes that facilitate tube elongation The diagram represents a

Figure 12 Angiosperm shoot meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

Figure 13 Angiosperm root meristem section Source Alison Roberts Reproduced with

permission of University of Rhode Island

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 17: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

6 Plant cells and their organelles

significant advance over the early studies of pollen tubes as it assigns function to

ultrastructural components for example signalling molecules the Rho family of

GTPases and phosphatidylinositol 45 bisphosphate appear to be localized in the

apical plasma membrane Besides pollen tubes root hairs exhibit polar growth

Cell organelles ndash an introduction

Organelles are required for plant growth development and function (Sadava

1993 Gillham 1994 Herrmann 1994 Agrawal 2011) These organelles

(Figure 17) are the loci for a myriad of physiological and biochemical processes

(Tobin 1992 Daniell and Chase 2004 ndash see individual chapters)

There are many diagrams of a generalized plant cell Some of these are available

at wwwexplorebiologycom httpwwwdaviddarlinginfoimagesplant_celljpg

Table 12 Meristems and their derivatives

Meristems Derivatives

Primary

Protoderm Epidermis

From tunica (Evert 2006)

Procambium (provascular) Primary xylem and phloem

From corpus (Evert 2006) Vascular cambium

Ground Ground tissue pith and cortex

Lateral

Vascular cambium

Fusiform initials Secondary xylem

Secondary phloem

Ray initials (Evert 2006) Ray cells

Cork cambium

Phellogen Replaces the epidermis when cork cambium initiates stem girth

increase composed of lsquoboxlikersquo cork cells which are dead at

maturity protoplasm secretes suberin some cork cells that are

loosely packed give rise to lenticels which function in gas exchange

between the air and the stemrsquos interior httpwwwBiology‐online

org Evert (2006) httpwwwvebrioSceincevunlenvirut

Periderm (Evert 2006)

Phelloderm Parenchyma cells produced on the inside by the cork cambium

Meristems are discussed by Steeves and Sussex (1989)

Table 13 Plant tissues

Tissue system

Meristematic Ground Vascular Dermal

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 18: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 7

and httpmicromagnetfsuedu The organelle contents of plant and animal

cells in common and those unique to plant cells are depicted in Table 14 The

dimensions of plant organelles are presented in Table 15 A plant organelle data-

base (PODB) has been reviewed by Mano et al (2008)

To enter a plant cell molecules must traverse both the cell wall and the fluid

mosaic plasmalemma (Singer and Nicolson 1972 Leshem et al 1991 Larsson and

Miller 1990) In contrast to the fluid mosaic model (Figure 18) of the plasmalemma

Figure 14 SEM of a pecan leaf Diagram of a leafrsquos interior is available at httppics4learning

com Source Reproduced with permission of Asaf Gal

Both day length and temperature regulate owering

Flowering is regulated by a protein hormone named origen

Leaf primordia

Includes the shoot meristem and the owers Inorescence meristem

Floral meristems generate oral organs ie sepals petals stamens and carpels

Genes occur in oral m

eristemsH

omeo

tic g

enes

enc

ode

prot

eins

Figure 15 Schematic of the floral meristem

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 19: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

H2O

CO2H2O

Sugars

Sugars

Sugarsstarch

starch or sugarstorage organ

Light energy

vapour

Starch or sugarstorage organPhotosynthesis

respiration andphotorespiration

Respiration nophotorespiration

O2

CO2

O2 H2O andmineralsenter throughroot hairs

C6H12O6

Figure 16 Diagram of angiosperm plant body Source From httpwwwmsueducourse

te8021science08plantsfoodshtml

Figure 17 Electron micrograph of a plant cell and its organelles Source Reproduced with

permission of HJ Horner

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 20: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 9

the picketndashfence model proposes the accumulation of membrane protein anchored

in an actin network beneath the membrane (Kusumi et al 2012)

The plasmalemma is composed of water protein and lipids There are both

integral and peripheral proteins (Leshem et al 1991) The integral proteins may

be simple (classical α‐helical structure that traverses the membrane only once)

or complex (globular ndash composed of several α‐helical loops which may span the

membrane several times) Peripheral proteins can be easily isolated by altering

Table 14 Comparison of organelle contents of plant and animal cells

Organelle Animal cell Plant cell

Cell wall Absent Present

Centrioles Present Absent

Endoplasmic reticulum Present Present

Glyoxysomes Absent Present

Golgi apparatus Present Present

Microfilaments Present Present

Mitochondrion Present Present

Nucleus Present Present

Peroxisomes Present Present

Plastids Absent Present

Protein bodies Absent Present

Spindle Present Present

Vacuoles Sometimes small Present (mature

cell ndash large central)

Early discussions of plant cell organelles occur in Hongladarom et al (1964) Pridham (1968) Reid

and Leech (1980) and Tobin (1992)

Table 15 Dimensions of subcellular organelles

Organelles Dimension

Chloroplast 4ndash6 microm in diameter

Golgi apparatus Individual cisternae 09 microm

Coated vesicles 50ndash280 microm in diameter

Microbodies 01ndash20 microm in diameter

Microtubules 05ndash10 microm in diameter

Mitochondria 1ndash10 microm

Nuclear envelope pores 30ndash100 microm in diameter

Nucleus 5ndash10 microm in diameter

Peroxisome 02ndash07 microm

Plasmodesmata 2ndash40 microm in diameter

Primary wall 1ndash3 microm

Protein bodies 2ndash5 microm in diameter

Vacuoles 30ndash90 of cell volume

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 21: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

10 Plant cells and their organelles

the ionic strength or pH of the encasing medium The transport proteins are

pumps carriers or chemicals (see section on membrane transport) The lipids are

electro-negative and anionic phospholipids sphingolipids (Figure 19) chloroplast‐

specific glycerolipids and sterols (Table 16)

Lipid rafts are specialized phase domains containing sterols and sphingolipids

which may be important in signal transitions (Gray 2004 Furt et al 2007

Grennan 2007 Mongrand et al 2004) Caveolae which give rise to clathrin‐

coated vesicles (Brodsky et al 2001) are anchored multifunctional platforms in

lipids (Van Deurs et al 2003 Patel and Insel 2009)

The organization of the caveolae (Bastani and Parton 2010) in the plasma-

lemma and clathrin‐coated vesicles (Samaj et al 2005) is presented in Figure 110

The current discussion focuses on membrane transport mechanism Plants can

internalize certain molecules by endocytosis via invaginations of the plasmalemma

yielding clathrin‐coated vesicles (Figure 111 Holstein 2003) which become the

endosome (Low and Chandra 1994 Battey et al 1999 Šamaj et al 2006)

Proteins involved in clathrin‐dependent endocytosis appear to be clathrin adaptor

proteins and two adaptins (Pearse and Robinson 1990 Šamaj et al 2006) Plant

endocytosis and endosomes (Contento and Bassham 2012) seem to be significant

in auxin‐mediated cellndashcell communication gravity responses stomatal move-

ments cytokinesis and cell wall morphogenesis (Šamaj et al 2006)

Ion channels

Plasma membranes contain potassium (K+) calcium (Ca++) and anion channels

(Roberts 2006) Voltage‐gated ion channels are transmembrane ion channels acti-

vated by changes in electrical potential Gating is the precise control of ion channel

opening (Krol and Trebacz 2000) An example of an ion channel is the K+ the

Fluid mosaic model of the plasmalemma

Consists of a lipid bilayer in which globular proteins are embedded There are two types of proteins integraland peripheral Oliogsaccharides (2ndash20 monosaccharides) can be attached to the integralproteins Phospholipids from the bilayer with a polar head on the outside and non-polar tails on theinside

Fence model of the plasmalemma

There is a membrane skeleton with skeleton-anchored proteins and transmembrane proteins projectedoutwards into the cytoplasm Cytoplasmic domains of proteins collide with the actin skeleton yieldingtemporary con nement of the transmembrane proteins The membrane can contain lipid rafts andrelated caveolae invaginations The rafts are combinations of proteins and the lipids which may function insignalling sphingolipids are prevalent in the rafts

Picket model of the plasmalemma

Phospholipids can also be con ned by the membrane skeleton Some investigators combine the fenceand picket models

Figure 18 Top Fluid mosaic model of the plasmalemma Middle Fence model of the

plasmalemma Bottom picket model of the membrane

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 22: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 11

inwardly potassium channel This type of channel possesses a positive charge in

the cell Stomatal pore movements are mediated by a rise in intracellular K+ and

anion contents of guard cells (Schroeder and Hagiwara 1989) Another example

is the adenosine triphosphate (ATP) binding cassette transporter or ABC trans-

porter These transport toxic substances from the cell or into the vacuole These

Figure 19 Structures of (a) phospholipids and (b) sphingolipids

O

Phosphatidic acid Phosphatidylethonolamine Leeithin

Phosphatidylserine Phosphatidylinositol 1-Lysoleeithin

Glycerol-3 phosphate

Phosphatidate

Phosphatidylcholine(leeithin)

OH

OO

O

O

O

O

OH

OH

R1

R2

P

P

OO

O

O O

O

R1

R2 O

OH

P NH2

CH3

OO

OO

O

OO

R1

R2

OH

PN

CH3

CH3CH3

+

OOO

O

OOO O

OOO O

OO

+

+

+

+

+O

OOO

O

P

R1

R2 OHOH

OH

OH

OH

P

OO

OO

O

OO

HO

PN

+

OO

O

OO CH3

CH3

CH3OH

HO

H

HOCH

CHCH

CH3

CH2CH2

H3C

P

CCCC

N

CH2

CH2

CH2

H2CH2C

H2C

OO

O

O O

O

O O

R1

R2

PNH2OH

OH

(a)

HHO

H H

H

H

H H

HH

CH3

CH3

H3C

H3C

CH3

CH3

Alpha-spinasterol

29 sterol (C29H48O)

isolated from `Phytolaccaacute

httpwww3dmetdnaaffecgojpbin2show_dataeacc-B02450

Brassicasterol

28 sterol (C28H46O)

synthesized by oilseed rape

and several unicellular algae

httpenwikipediaorgwikiBrassicasterol

HO

HO(b)

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 23: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

12 Plant cells and their organelles

transporters are composed of four core domains two cytosolic nucleotide‐binding

proteins and two transmembrane domains (Malmstrom 2006)

Besides cation channels there are anion channels regulated by voltage but

their activity is also influenced by Ca++ ATP phosphorylation or membrane

stretching (Tyerman 1992) Anion plasma membrane channels function as

efflux channels when they are open

Table 16 Composition of certain cellular membranes

Chemical composition

Fatty acyl groups in membrane lipids

160 161 t‐161 163 180 181 182

α183 δ183 184 220 221 240 241

Electroneutral phospholipids Phosphatidylcholine phosphatidylethanol

phosphatidylethanolamine

Anionic phospholipids Phosphatidylserine phosphatidylglycerol

phosphatidylinositides

Lyo‐phospholipids Cerebrosides

Sphingolipids Galactolipids sulpholipids

Chloroplast‐specific glycerolipids Diphosphatidylglycerol and monophosphatidylglycerol

Mitochondrial phospholipids

Sterols Sitosterol

Campesterol

Stigmasterol

Unusual sterols

Cycloartenol

Cholesterol minute quantities

Sterol glycosides

Lanosterol Pathogenic fungal membranes

Water

Extramembrane water Membrane is a bilayer sandwiched between two layers

of water

Water located within the bilayer which is attached to

or in approximate contact with the expanses of

membrane constituents

Proteins May cross the membrane once or several times and are

linked either electrostatically or by means of biophysical

lipophilicity to the inner domains of the bilayer

Integral proteins

Simple integral proteins Classic α‐helical structure that traverses the membrane

only once

Complex integral proteins Globular ndash comprised of several α‐helical loops that

may span the membrane several times

Peripheral proteins Associated with only leafletndasheasily isolated by altering

ionic strength or pH of the encasing medium

Transport proteins Pumps carrier and channel

Source From Leshem et al (1991)

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 24: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 13

Figure 110 Depictions of a (a) lipid raft (b) caveolae and a clathrin‐coated vesicle Source

Reproduced with permission of Caveolae and Clathrin Vesicle

CytosolCholesterol

Protein with longertransmembrane domain

Normal trans-Golginetwork membrane

Protein with shorttransmembranedomain cannotenter lipid raft

GPI-achoredprotein

Lectins

Glycolipids

Lumen

(a)

Integral proteins excludedfrom transport vesicles

Exoplasmic face

Cytosolic face

Assemblyparticle

Fibrousclathrincoat

GTP

GDP

Dynamin

Clathrin-coatedvesicle

(b)

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 25: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

14 Plant cells and their organelles

proton pumps

The transport of a substance against its electro channel gradient requires energy

generated by ATP‐proton pumps (Briskin and Hanson 1992 Evert 2006) One

such pump is the V‐ATPase found in both the plasmalemma and the tonoplast

(Barkla and Pantoja 1996 Vinay et al 2009) The H+‐ATPase in the plasma-

lemma is the P‐ATPase which forms electrochemical gradients (Elmore and

Coaker 2011) Mitochondria and chloroplast membranes possess F‐ATPases

Water channels

Aquaporins are channel proteins which exist in the plasmalemma in intracellular

spaces (Maurel et al 2008) These proteins permit water to move freely but

exclude ions and metabolites (Chrispeels and Maruel 1994 Muller et al 2007)

Intermediate endosome

δ-TIP

Late endosomePVC

Vacuole

ARA6

ARA7RHA1

AtVAMP727

AtSKD1

PM ATPase PM receptor

AtSYP21AtSYP22

VSR

AtALEU

GNOMBFA

PIN1+NPAndashbinding protein

TGNAtSYP61

Golgi

ERCPCR

SVP ndash a syntaxinGNOM ndash Plant-specic protein that participates in ADP-ribosylationESCRT ndash protein endosomal sorting complexRHA ndash a member of the Rab GTPases function in trafcking pathwaysARA6 ndash a member of the Rab GTPasesSYP ndash a SNARE component of the late endosomeVSR ndash vacuolar sorting receptorSKD ndash vacuolar protein suppressorUbiquitylation ndash signal that regulates the cell surface expression

Figure 111 Diagram of plant endocytosis Source Reproduced with permission of M Otegui

University of Wisconsin

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 26: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

An introduction to cells and their organelles 15

providing for buffering osmotic fluctuations in the cytosol Aquaporins are major

intrinsic membrane proteins which are composed of four subunits each of

which comprises six transmembrane‐spanning helices Aquaporins are encoded

by multiple gene families (Johansson et al 1998)

Carriers

Carriers are unitransporters and co‐transporters (Evert 2006) Unitransporters

transport only one solute from one side of the membrane to the other On the

contrary co‐transporters transfer one solute with the simultaneous or sequential

transfer of another solute A thorough discussion of membrane transport

processes occurs in Malmstrom (2006)

Organelle structure and function can be influenced by a variety of environ-

mental parameters which affect plant growth A discussion of parameters is pre-

sented because of the increasing pollution of the earthrsquos atmosphere and

ecosystem In addition global climate change is a current issue of urgent con-

cern (Dashek and McMillin 2009)

Both major and minor elements are required for growth and development

(Table 17) Metals and metalloids at elevated levels can result from mining

(Dashek and McMillin 2009) What effects do these levels have on the structure

and function of cellular organelles (See Lepp 1981 Medioini et al 2008 Yusuf

et al 2011 see also Table 18)

Elevated levels of SO2 CO

2 NO

2 and O

3 (Treshow and Anderson 1989) can

occur in the atmosphere as a result of industrial and contemporary activities

Table 19 presents the effects of certain gases (Bell and Treshow 2002) on the

structure and function of organelles Of special interests are the increasing levels

Table 17 Major and minor elements required for plant growth and development

Element mgkg Minor or major

Nitrogen N 15 000 Major

Potassium K 10 000 Major

Calcium Ca 5 000 Major

Magnesium Mg 2 000 Major

Phosphorus P 2 000 Major

Sulfur S 1 000 Major

Chlorine Cl 100 Minor

Iron Fe 100 Minor

Boron B 20 Minor

Manganese Mn 50 Minor

Zinc Zn 20 Minor

Copper Cu 6 Minor

Molybdenum Mo 01 Minor

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)

Page 27: Plant Cells and their Organelles · Plant Golgi introduction, 66 Structure and organization, 69 Golgi‐mediated vesicular trafficking, 71 Plant Golgi‐dependent cellular processes,

16 Plant cells and their organelles

of CO2 in the atmosphere which many scientists believe causes global warming

(Dashek and McMillin 2009) Table 110 offers the effects of sublethal and lethal

temperatures on organelles Franklin and Wigge (2014) discuss the effects of

temperature on plant development Other environmental parameters which can

Table 18 Toxic metals and metalloids

Metal or metalloid

Toxic level effects References

Aluminium Affects root cells of plasmalemma Mossor‐Pietraszewska (2001)

Arsenic Pale green to yellow lesions on leaves and

necrosis of leaves

Treshow and Anderson (1989)

Defoliation

Impaired nitrogen metabolism

Needle abscission

Cadmium General chlorosis Treshow and Anderson (1989)

Saadati et al (2012) and Khateeb

(2014)

Reduced photosynthesis

Reduced transpiration toxic effects ndash changes

in proline levels changes in lipid peroxidation

and seed germination

Copper Interference with normal metabolic reactions Treshow and Anderson (1989)

and Shah et al (2001)Blocks specific enzymatic reactions

Chromium Contamination Treshow and Anderson (1989)

and Antonovics et al (1971)Can promote white dead patches on leaves

Lead Condensation of nuclear chromatin decrease

in germination of two Brassica cultivars

Rout and Das (2003) and

Hosseini et al (2007)

Nickel Dilution of nuclear membrane Seregin and Kozhernikova (2006)

Zinc Disruption of cortical cell Rout and Das (2009)

Table 19 Effects of environmental pollutants on organelles

Elevated CO2

Stomatal openings reduce as CO2 increases Woodward et al (1991)

Affects both primary and secondary meristems

of shoots and roots alternation of leaf size and

anatomy increased branching and stem

diameter

Pritchard et al (1999)

Increase in the number of mitochondria and

amount of chloroplast stroma thylakoid

membranes

Griffin et al (2001)

Stomatal densities decrease in two species of

Spartia

Lammertsmaa et al (2011)

Acid rain Leaching of nutrients on tree needles damages

surfaces of needles and leaves and reduces a

treersquos ability to withstand cold

Godbold and Huumlttermann

(1994) Schulze et al (2000)

and White and Terninko (2003)

Nitric oxide Necrotic lesions marginal chlorosis Lamattina and Polacco (2007)

Ozone and its

derivatives

Changes in metabolism Roshchina and Roshchina (2003)