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BOTANICAL RESEARCH AND PRACTICES

SUNFLOWERS

GROWTH AND DEVELOPMENT,

ENVIRONMENTAL INFLUENCES

AND PESTS/DISEASES

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SUNFLOWERS

GROWTH AND DEVELOPMENT,

ENVIRONMENTAL INFLUENCES

AND PESTS/DISEASES

JUAN IGNACIO ARRIBAS

EDITOR

New York


Copyright 2014 by Nova Science Publishers, Inc.

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Library of Congress Cataloging-in-Publication Data

Sunflowers : growth and development, environmental influences and pests/diseases / editor: Juan

Ignacio Arribas (Electrical Engineering Department, Univ. Valladolid, Spain).

pages cm

Includes index.

1. Sunflowers. I. Arribas, Juan Ignacio.

QK495.C74S87 2014

583'.99--dc23

2014003599

Published by Nova Science Publishers, Inc. New York

ISBN: (eBook)


CONTENTS

Preface vii

Chapter 1 An Introduction to the Sunflower Crop 1 Fabin Fernndez-Luqueo, Fernando Lpez-Valdez,

Mariana Miranda-Armbula, Minerva Rosas-Morales, Nicolaza Pariona and Roberto Espinoza-Zapata

Chapter 2 Floral Biology of Sunflowers: A Histological

and Physiological Analysis 19 Basudha Sharma, Rashmi Shakya and Satish C. Bhatla

Chapter 3 Development of Female Reproductive Structures and Apomixis

in Sunflowers 43 Olga N. Voronova

Chapter 4 Genetics and Genomics Applied to Sunflower Breeding 61 Carla Filippi, Jeremas Zubrzycki, Vernica La,

Ruth A. Heinz, Norma B. Paniego and H. Esteban Hopp

Chapter 5 Sunflower Genetic Resources: Interspecific Hybridization

and Cytogenetics in Prebreeding 95 Jovanka Atlagi and Sreten Terzi

Chapter 6 Functional Genomics and Transgenesis Applied to

Sunflower Breeding 131 Sebastian Moschen, Laura M. Radonic, Guillermo F. Ehrenbolger, Paula Fernndez, Vernica La, Norma B. Paniego, Marisa Lpez Bilbao,

Ruth A. Heinz and H. Esteban Hopp

Chapter 7 Disease Management in Sunflowers 165 Regina M. V. B. C. Leite

Chapter 8 Recent Advances for Developing Resistance against

Plasmopara halstedii in Sunflowers 187 Osman Radwan


Contents vi

Chapter 9 Effects of Crop Management on the Incidence and Severity

of Fungal Diseases in Sunflowers 201 P. Debaeke, E. Mestries, M. Desanlis and C. Seassau

Chapter 10 Insect Pests of Sunflowers in Africa 227 Hannalene du Plessis

Chapter 11 Soil Amendments and Their Effects on Sunflower Growth 239 Fernando Lpez-Valdez, Fabin Fernndez-Luqueo, Perla Xchitl Hernndez-Rodrguez, Minerva Rosas-Morales

and Silvia Luna-Surez

Chapter 12 Nutrition and Fertilization of Sunflowers in Brazilian Cerrado 257 C. de Castro, F. A. Oliveira, A. Oliveira Junior

and N. P. Ramos

Chapter 13 Environmental Issues in the Sunflower Crop of Midwestern Brazil:

Diversification and Complementarities in the Biodiesel Chain 281 N. P. Ramos, A. M. M. Pires, C. C. A. Buschinelli,

H. B. Vieira, C. de Castro and G. S. Rodrigues

Chapter 14 Micro and Macro-Morphological Variation of Cosmos bipinnatus

and Cosmos bipinnatus var. Albiflorus in Sympatric Zones in

Central Mexico 297 M. Paniagua-Ibaez, A. Zepeda-Rodrguez, P. Mussali-Galante,

R. Ramrez-Rodrguez and E. Tovar-Snchez

Index 309


To Juan Ignacio, Jr., Elena, Jr. and Elena


PREFACE

We are all well aware that the importance of the sunflower (Helianthus Annuus) as a crop

has increased significantly in recent years, not only in the food industry but also as a natural

energy resource in oil production. I am, thus, very pleased to be able to present this

comprehensive monograph on a wide range of important issues regarding sunflowers, with an

emphasis on environmental influences, pests and diseases in order to maximise production

whilst minimising costs.

Contributors where selected based on their proven experience in the field of sunflowers.

Contributors submitted an extended abstract that was assessed for relevance. They were then

invited to contribute draft chapters. Each chapter underwent a stringent and thorough peer

review process by other experts in the field, with final approval by the editor who, thus, was

able to balance the topics from all contributors.

The book contains important original results. Each chapter deals with a different topic,

and draws, where appropriate, from studies and results previously published by the authors.

Authors were encouraged to complement their writing with original and high quality graphs,

charts, tables, figures, pictures and photographs.

Its my honour and pleasure to acknowledge the rigorous work carried out by all authors

in this book, and at the same time I am very grateful to them for trusting me in leading this

project in the role of the editor of their work. My thanks also go to the anonymous reviewers

who contributed their time so generously to this book, and without whom it would not exist.

I am also very grateful to Nova Science Publishers for inviting me to lead this book, and

thank them for the help and coverage provided during the whole time that this project lasted.

I really do hope that you find this book of interest and wish you enjoy its reading as much

as I have done through the whole editing process and as much I am sure all authors have done

while writing it.

The book is structured as follows: Chapter 1 introduces sunflowers. Chapters 2 and 3

detail the biology of a sunflower. Chapters 4, 5 and 6 explore the important topics of

sunflower production: genetics, genomics and the breeding of sunflowers. Chapters 7, 8, 9

and 10 address with other important aspects of sunflower production, pests and diseases.

Chapters 11 and 12 deal with sunflower nutrition and growth. Finally, Chapter 13 presents

environmental sunflower crop issues and Chapter 14 studies sunflower morphological

Cosmos variations.


Juan Ignacio Arribas x

In Chapter 1, entitled An Introduction to the Sunflower Crop, Dr. Luqueno and colleagues

from the Natural Resources and Energy Group, Cinvestav-Saltillo, Mexico, present an

enjoyable historical perspective on sunflowers. Sunflower (Helianthus annuus L.) belongs to

the family Asteraceae. The sunflower plant originated in eastern North America. It is thought

to have been domesticated around 3000 B.C. by Native Americans. In the late 1800s the

sunflower was introduced in the Russian Federation where it became a food crop and Russian

farmers made significant improvements in the way that the sunflower was cultivated. Since

3000 B.C. a wide range of uses of sunflower have been reported throughout the world.

Sunflower is well known by its phytoremediation potential and by its seed oil content.

Because the sunflower has several potential markets, it is a good choice for growers on both

small and large scales. However, it has to be remembered that scientific, technical or

agricultural projects linked with sunflower have to include side effects elsewhere in order to

shape a sustainable future.

In Chapter 2, entitled Floral Biology of Sunflower - A Histological and Physiological

Analysis, Dr. Bhatla and colleagues from the Laboratory of Plant Physiology and

Biochemistry, Department of Botany, University of Delhi, India, introduce a meticulous

approach to the development of sunflower inflorescence as considered under three phases

listed next: inflorescence initiation, floret development and anther formation. Anthesis of disc

florets is a phytochrome-mediated response and is also modulated by phytohormones, such as

auxins and gibberellic acid. Dr. Bhatla and colleagues focus on the role of various

biomolecules, like glycoproteins, calcium, nitric oxide, reactive oxygen species, and

associated scavenging enzymes in relation to stigma maturation. Specific expression of

lignoceric acid (24:0) in the pollen coat and localization of lipase in pollen and stigma are

likely to have possible roles during pollen-stigma interaction. The phenomenon of self-

incompatibility and pseudo self-compatibility in sunflower has been discussed. The initial

processes accompanying pollen-stigma interaction and their regulation, especially the

adhesion of pollen on the stigma surface, hydration, formation of an "attachment foot" and

pollen tube germination in sunflower with respect to self-and cross-pollinated situations, has

also been dealt with in detail.

In Chapter 3, entitled Development of Female Reproductive Structures and Apomixis in

Sunflowers, Dr. Voronona from the Department of Embryology and reproductive biology,

Komarov Botanical Institute of RAS, Saint-Petersburg, Russia, presents an scrupulous visual

analysis of archesporial cells which are formed and gave rise to megaspore mother cells. The

meiotic divisions produced a linear tetrad of haploid megaspores and from one chalazal

megaspore a Polygonum-type embryo sac is formed. Under natural conditions the apomixis

phenomenon was hardly observed in genus Helianthus L. In addition, author shows how on

plants of CMS-lines a number of anomalies in development of female reproductive system

were detected, including such phenomena as total absence of embryo sac, apospory and

integumentary embryony. Lack of the main embryo sac and formation of additional

aposporous embryo sacs could be observed in the same ovule. Finally, investigation of the

early stages of ovule development showed that aposporous embryo sacs originated from the

same ovule subepidermal cells as a normal embryo sac.

In Chapter 4, entitled Genetics and Genomics Applied to Sunflower Breeding, Dr. Hopp

and colleagues from the Instituto de Biotecnologia, Centro de Investigaciones Veterinarias y

Agronomicas, Instituto Nacional de Tecnologia Agropecuaria, Hurlingham, Argentina,

present a thoughtful study about new breeding strategies based on molecular markers, like


Preface xi

quantitative trait loci mapping, association mapping and genomic selection that are currently

being developed for commercial crop improvement. The need to increase efficiency and

precision, and save time, resources and efforts, has motivated the application of Marker

Assisted Selection (MAS) in sunflower breeding programs. Furthermore, nowadays, the focus

is on tolerance improvement to biotic and abiotic stresses and oil quality and yield increasing,

in order to reduce the gap between potential and actual sunflower production.

In Chapter 5, entitled Sunflower Genetic Resources Interspecific Hybridization and

Cytogenetics in Prebreeding, Drs. Atlagic and Terzic present a rigorous description of the

genus Helianthus by reviewing genetic resources, cytogenetic research and application in

breeding. Besides the review of available literature, research results of the Institute of Field

and Vegetable Crops are presented in detail since the establishment of its collection in 1980.

Experience collected during this period indicates the difficulties in collection maintenance,

interspecific crosses and isolation of desired genes. Nevertheless, genus Helianthus proved to

be a good source of material for the constant improvement of cultivated sunflower.

In Chapter 6, entitled Functional Genomics and Transgenesis Applied to Sunflower

Breeding, Dr. Hopp and colleagues from the Instituto de Biotecnologia, Centro de

Investigaciones Veterinarias y Agronmicas, Instituto Nacional de Tecnologia Agropecuaria,

Hurlingham, Argentina, introduce an interesting chapter where they analyze different

strategies which have been developed in the last decade from functional genomics and post

genomics disciplines to contribute to the elucidation of gene regulation and identification of

key metabolic pathways involved in the response to biotic and abiotic stresses in sunflower.

The state of the art of strategies for gene function, studies in silico and in planta, by stable

gene transfer or agroinfiltration in sunflower as well as in the model system for Asteraceae

species, lettuce, are discussed within the frame of their application in sunflower breeding.

In Chapter 7, entitled Disease Management in Sunflowers, Dr. Leite from Embrapa

Soybean, Brazil, presents an interesting approach regarding the most important sunflower

diseases and strategies for disease management. Sunflower can be affected by the presence of

diseases, which may, depending on climatic conditions that favor the occurrence of pathogens

and the infective process, lead to a significant reduction on yield and quality of product.

Disease management should be based on an integrated program, in order to give support for

the sustainability and competitiveness of the sunflower crop.

In Chapter 8, entitled Recent Advances for Developing Resistance against Plasmopara

Halstedii in Sunflowers, Dr. Radwan from the Department of Natural Resources and

Environmental Sciences, University of Illinois at Urbana-Champaign, Urbana, IL, USA,

presents an interesting visual approach to Downy Mildew disease, as one of the most

important diseases of sunflower, which leads to an economic yield loss. In last two decades,

different approaches of genetics and genomics have significantly contributed to better

understand sunflower-Plasmopara halstedii. This progress directed to development of

sunflower lines carrying resistance to different races of this pathogen.

In Chapter 9, entitled Effects of Crop Management on the Incidence and Severity of

Fungal Diseases in Sunflowers, Dr. Debaeke and colleagues from the Institut National de la

Recherche Agronomique (INRA), Toulouse, France, dissected the effects of crop

management on the incidence and severity of major fungal diseases in sunflower, including

downy mildew, phoma, phomopsis and sclerotinia. They deeply reviewed and discussed the

influence of sowing date, plant population, N fertilization, and irrigation on sunflower

diseases from numerous experiments conducted in France during the last twenty years. They


Juan Ignacio Arribas xii

proposed indicators of canopy development and nutritional status that could be useful when

developing crop management systems with reduced chemical applications.

In Chapter 10, entitled Insect Pests of Sunflowers in Africa, Dr. du Plessis from the Unit

for Environmental Sciences and Management, North-West University, Potchefstroom, South

Africa, present a complete study regarding a number of insect species which have adapted to

cultivated sunflower as a source of food and have consequently become economically

important pests. However, although many insect species is associated with sunflower in

Africa, only few are considered to be of potential economic importance. Insects most

commonly reported as injurious to this crop, occur sporadically, but usually in high numbers.

The families Noctuidae, Tenebrionidae, Curculionidae, Pentatomidae and Orsillidae are the

most important. Various types of damage are caused to sunflower seedlings, but the damage

symptoms are specific to a particular pest species. Dr. du PLessis argues that these seedling

pests mainly constitute dusty surface beetles (Gonocephalum simplex), greater false wire

worms (Somaticus spp.), cutworms (Agrotis spp.) and ground weevils (Protostrophus spp.).

Total defoliation can be incurred by the Plusia looper, Trichoplusia orichalcea. Hemipterans

and the African bollworm, Helicoverpa armigera are the most important insect pests of

sunflower during the heading stages of crop development. Intensive feeding by hemipterans

during this development stage results in deformed heads, which delay flower opening. The

occurrence of the false chinch bug, Nysius natalensis on sunflower during the heading stage

onwards in South Africa, is similar to that of N. stali in Nigeria. Since high summer

temperatures prevail throughout the sunflower production area of South Africa and the most

favourable temperature range for N. natalensis development is between 26C and 38 C, the

potential for rapid population build-up by this pest during the sunflower production season is

good. It is likely that N. natalensis can become important in sunflower production in other

African countries with similar weather conditions too. The insect is polyphagous and a variety

of wild host plants, mainly weed species as well as crops such as grain sorghum play an

important role in sustaining its populations. Sunflower is not the preferred host but N.

natalensis lays its eggs on sunflower when its preferred host plants are removed or dead. This

behaviour explains the insects injuriousness to late-planted sunflower because weed species

have already senesced before the sunflower. This period often coincides with seed fill of late-

planted sunflower, providing an alternative for the insect for moisture, as well as seeds that

are necessary for reproduction of the pest. Weeding in and around sunflower during seed fill

of the crop, therefore results in destruction of the preferred host plants of N. natalensis, and

they consequently move to sunflower where they feed and cause damage. When considering

application of insecticides for control of this pest, it should take into consideration that N.

natalensis is highly mobile and continuous re-infestations could occur. Timing of insecticide

application is therefore important. African bollworm (H. armigera) is frequently present

during the reproductive stage of cultivated sunflower in Africa. The attractiveness of

sunflower to this pest is demonstrated by the traps use as a trap crop in and around organic

cotton fields in Tanzania. Larvae occur from the budding stage onwards. Levels of infestation

vary between localities and seasons, sporadically reaching epidemic proportions. Sunflower

has, however, the ability to compensate for head damage and along with the fact that

preferential feeding sites of H. armigera are not the achenes, a significant number of larvae

could be tolerated without any significant effect on yield. Actual damage is, therefore, the

only criterion that could be used in the determination of economic injury levels for control of

African bollworm on sunflower crop.


Preface xiii

In Chapter 11, entitled Soil Amendments and Their Effects on Sunflower Growth, Dr.

Lopez and colleagues from the Centro de Investigacion en Biotecnologia Aplicada, Instituto

Politecnico Nacional, Tlaxcala, Mexico, present a novel approach to several forms of amend

or fertilize sunflowers as alternative to improves this important cultivar. In particular authors

are interested in the study of organic materials that could be applied to soil in order to

improve their properties and plant growth, keeping in mind that we must recycle or reuse this

kind of materials. Finally, the organic amendments could be a beneficial disposal approach

that must be considered.

In Chapter 12, entitled Nutrition and Fertilization of Sunflowers in Brazilian Cerrado,

Dr. Castro and colleagues present an interesting chapter centered in a particular region in

Brazil, which authors argue that is recognized as a major global food producer and despite the

fact that its agriculture occupies only less than 5 % of the national territory, the estimated

grain production for 2013/2014 growing season is 195 million tons. The Brazilian Cerrado is

the main agriculture expansion region in the country, driven by soybean cultivation in an area

of 13 million ha. In this tropical agricultural region, sunflower has great potential for

expansion and consolidation as an important component of sustainable crop rotation

production systems. This chapter addresses the major limiting soil fertility factors which

hinder the crop development and discusses fertilization management practices related to the

main limiting nutrients, like the macronutrients nitrogen, phosphorus and potassium and

micronutrients such as boron and molybdenum. Adequate management of soil acidity and

fertilization has been proved as a powerful tool to improve the natural conditions of acidic

and chemically poor arable tropical soils. In addition to the soil fertility assessment, leaf

analysis is essential for the proper interpretation of plant nutritional status, thereby enabling

better refinement of the crop nutritional management.

In Chapter 13, entitled Environmental Issues in the Sunflower Crop of Midwestern Brazil

Diversification And Complementarities in the Biodiesel Chain, Dr. Ramos and colleagues

from Embrapa Environment, Brazil, present an interesting study regarding the increase in

global demand for renewable energy, the production of oilseeds, including sunflower, as

feedstock for biodiesel. The increase in global demand for renewable energy has encouraged,

both directly and indirectly, the production of oilseeds, including sunflower, as feedstock for

biodiesel. In this scenario, authors argue that Brazil stands out for its excellent agronomic and

climatic conditions for growing these crops throughout its territory. Sunflower is considered

an interesting option and your production in the mid-western region of Brazil has done

valuable contributions as a second, and especially the production practices adopted by the

reference farmers of the country, rendering complementarities and diversification to both the

food and the bioenergy sectors.

In Chapter 14, entitled Micro and Macro-Morphological Variation of Cosmos Bipinnatus

and Cosmos Bipinnatus Var. Albiflorus in Sympatric Zones in Central Mexico, Dr. Paniagua

and colleagues from the Centro de Investigacion en Biodiversidad y Conservacion, Univ.

Autonoma Estado de Morelos, Morelos, Mexico, present a concise but at the same time

precise analysis of the morphological variations in various sunflower species in central

Mexico area. Mexico is considered one of the centers of diversification of the Asteraceae

family, which contains the greatest richness of flowering plants. Particularly, the Trans-

Mexican Volcanic Belt (TMVB) is a heterogeneous mountain belt, located in the central part

of the country in an eastwest direction, which has been considered a diversification site for

many genera due to the vast number of species that it contains, as well as its high degree of


Juan Ignacio Arribas xiv

endemism. Cosmos bipinnatus is present in various types of vegetation along the TMVB

which is favored by disturbances. Taxonomic studies have documented that this sunflower

species presents white to lilac ligules. However, horticulturists have considered the white

variety as C. bipinnatus var. albiflorus. Still, there is no scientific evidence to support their

observations. Therefore, the goal of this study was to compare the micro and macro

morphology characters between C. bipinnatus individuals with white and lilac ligules to

determine a possible morphological differentiation between both phenotypes in sympatric

zones in the central Mexico region. Principal Component Analysis and Non-Metric

Multidimensional Scaling showed a clear morphological differentiation between both groups;

this pattern was consistent even when the ligules color was not considered for the statistical

analyses. Dr. Paniagua and colleagues results sign a possible speciation between these

phenotypes and support a taxonomic shift for the Mexican sunflower with white ligules to C.

bipinnatus var. albiflorus.

Juan Ignacio Arribas, PhD

Associate Professor of Electrical Engineering

University Valladoild, Spain

Valladolid, November 2013

Tel: +34 983423000

Fax: +34 983423667

[email protected]


In: Sunflowers ISBN: 978-1-63117-347-9

Editor: Juan Ignacio Arribas 2014 Nova Science Publishers, Inc.

Chapter 1

AN INTRODUCTION TO THE SUNFLOWER CROP

Fabin Fernndez-Luqueo1,

, Fernando Lpez-Valdez2,

Mariana Miranda-Armbula2, Minerva Rosas-Morales

2,

Nicolaza Pariona1 and Roberto Espinoza-Zapata

3

1Natural Resources and Energy Group, Cinvestav-Saltillo, Coahuila, Mxico

2CIBA - Instituto Politcnico Nacional, Tepetitla de Lardizbal, Tlaxcala, Mxico

3Crop Breeding Department, UAAAN, Saltillo, Coahuila, Mxico

ABSTRACT

Sunflower (Helianthus annuus L.) belongs to the family Asteraceae. The Helianthus

genus contains 65 different species of which 14 are annual plants. The sunflower plant

originated in eastern North America. It is thought to have been domesticated around 3000

B.C. by Native Americans. In the late 1800s the sunflower was introduced in the Russian

Federation where it became a food crop and Russian farmers made significant

improvements in the way that the sunflower was cultivated. Since 3000 B.C. a wide

range of uses of sunflower have been reported throughout the world such as ornamental

plant, medicinal, alimentary, feedstock, fodder, dyes for textile industry, body painting,

decorations, and so on. Sunflower species are allelopathic in nature and this crop appears

to have a bright future, especially if the scientists can translate the cutting-edge research

into technologies that will reduce the reliance on synthetic herbicides, pesticides, and

crop protection chemicals. On the one hand sunflower is well known by its

phytoremediation potential, thus it can be speculated that the good tolerance of sunflower

towards pollutants coupled with an increased accumulation/degradation capacity might

contribute to an efficient removal of pollutants from soil and water; on the other hand

sunflower possesses the potential to develop bioenergy systems that allow for synergies

between food and energy production. Because the sunflower has several potential

markets, it is a good choice for growers on both small and large scales. However, it has to

be remembered that scientific, technical or agricultural projects linked with sunflower

have to include side effects elsewhere in order to shape a sustainable future.

* Corresponding author: F. Fernndez-Luqueo, Natural Resources and Energy Group, Cinvestav-Saltillo, Coahuila.

C. P. 25900, Mxico Tel.: +52 844 4389625; Fax: +52 844 4389610. E-mail address: cinves.cp.cha.luqueno@

gmail.com.


F. Fernndez-Luqueo, F. Lpez-Valdez, M. Miranda-Armbula et al. 2

Keywords: Allelopathy, biodiesel, phytoremediation, renewable energy, sustainable

development, symbiosis

1. INTRODUCTION

Sunflower (Helianthus annuus L.) belongs to the family Asteraceae. Helianthus genus

contains 65 different species (Andrew et al., 2013). The name Helianthus, being derived from

helios (the sun) and anthos (a flower), has the same meaning as the English name Sunflower,

which has been given these flowers from a supposition that they follow the sun by day,

always turning towards its direct rays. The sunflower that most people refer to is H. annuus,

an annual sunflower. In general, it is an annual plant which possesses a large inflorescence

(flowering head), and its name is derived from the flower's shape and image, which is often

used to depict the sun. The plant has a rough, hairy stem, broad, coarsely toothed, rough

leaves and circular heads of flowers (Khaleghizadeh, 2011). The heads consist of many

individual flowers which mature into seeds on a receptacle base (Seghatoleslami et al., 2012).

Sunflower is the worlds fourth largest oil-seed crop and its seeds are used as food and its

dried stalk as fuel. It is already been used as ornamental plant and was used in ancient

ceremonies (Harter et al., 2004; Muller et al., 2011). Additionally, medical uses for

pulmonary afflictions have been reported. In addition, parts of this plant are used in making

dyes for the textile industry, body painting, and other decorations. Sunflower oil is used in

salad dressings, for cooking and in the manufacturing of margarine and shortening

(Kunduraci et al., 2010). Sunflower is used in industry for making paints and cosmetics. A

coffee type could be made with the roasted seeds. In some countries the seed cake that is left

after the oil extraction is used as livestock feed. In the Soviet Union the hulls are used for

manufacturing ethyl alcohol, in lining for plywood and growing yeast. The dried stems have

also been used for fuel. The stems contain phosphorous and potassium which can be

composted and returned to soil as fertilizer. Sunflower meal is a potential source of protein

for human consumption due to its high nutritional value and lack of anti-nutritional factors

(Fozia et al., 2008).

Sunflower was a common crop among American Indian tribes throughout North

America. Evidence suggests that the plant was cultivated by natives in present-day Arizona

and New Mexico about 3000 B.C. Some archaeologists suggest that sunflower may have been

domesticated before corn (NSA, 2013). Although the scientific consensus had long been that

sunflower was domesticated once in eastern North America, the discovery of pre-Columbian

sunflower remains at archaeological sites in Mexico led to the proposal of a second

domestication center in southern Mexico. However, evidences from multiple evolutionary

important loci and from neutral markets support a single domestication event for extant

cultivated sunflower in eastern North America (Blackman et al., 2011).

The objective of this chapter is to present and discuss a summary about the huge amount

of information in which the sunflower is the main subject. The chapter aims to assist people

involved in all aspects of sunflower management, including conservation, agriculture, mining,

energy, food production, health and other industries, to obtain a broad knowledge of

sunflower and of its ecosystem services.


An Introduction to the Sunflower Crop 3

2. BOTANICAL AND MORPHOLOGICAL DESCRIPTION

Sunflowers are botanically classified as Helianthus annuus L. (Table 1). They are large

plant and are grown throughout the world because of their relatively short growing season.

Sunflower is an annual herb, with a rough, hairy stem, 3 to 12 feet high, broad, coarsely

toothed, rough leaves, 3 to 12 inches long and circular heads of flowers, 3 to 6 inches wide in

wild specimens and often a foot or more in cultivation. The flower-heads are composed of

many small tubular flowers arranged compactly on a flattish disk: those in the outer row have

long strap-shaped corollas, forming the rays of the composite flower. Each sunflower head, or

inflorescence, is actually composed of two types of flowers. What appears to be yellow petals

around the edge of the head are actually individual ray flowers. The face of the head is

comprised of hundreds of disk flowers, which each form into a seed (achene).

The basic chromosome number for the Helianthus genus is 17. Diploid, tetraploid and

hexaploid species are known. There are only 14 annual species of Helianthus. Plant breeders

have made interspecific crosses within the genus and have transferred such useful characters

as higher oil percentage, cytoplasmic male sterility for use in production of hybrids, and

disease and insect resistance to commercial sunflower.

Table 1. Scientific classification of H. annuus L.; this genus counts 65 different species

Taxa

Kingdom Plantae

Subkingdom Viridaeplantae

Infrakingdom Streptophyta

Division Tracheophyta

Subdivision Spermatophytina

Infradivision Angiospermae

Class Magnoliopsida

Superorder Asteranae

Order Asterales

Family Asteraceae

Subfamily Helianthoideae

Tribe Heliantheae

Genus Helianthus

Specie annuus

The taxonomic classification has been in place since 1753.

3. PRODUCTION

In recent years, the sunflower cultivated area has been steadily increasing due to the

breeding of dwarf high yielding hybrids that also facilitate mechanization and the emphasis

given to polyunsaturated acids for human consumption. Global production grew steadily in

last 25 years (PSD-USDA, 2011), and FAO expect a total world output close to 60 million

tons towards 2050. The four largest producers (Russia, Ukraine, European Union and

Argentina) account for 70% of global volume, with an exponential growth of production in



the last ten years in the Black Sea region, with increased acreage an higher yields achieved by

the replacing old varieties by hybrid seeds.

According to data from FAOSTAT (FAOSTAT, 2011) Russia Federation ranked first

producing ca. 9.7 millions of tons of sunflower seeds or 26% of the world total. Ukraine and

Argentina ranked second and third place with 8.6 and 3.6 tons of sunflower seeds,

respectively. France, Romania, China, Bulgaria, Hungary, Turkey, and Spain produced

between 1.0 and 1.9 millions of tons of sunflower seeds (Table 2). The United States

produced ca. 1.0 millions of tons of sunflower seeds, or 5% of the worlds total production.

That is enough to make the United States rank eleventh in that category. South Africa ranked

twelfth producing ca. 0.9 millions of tons of sunflower seeds.

Table 2. The highest twelve sunflower seed producing countries

in the world during 2011

Place Countries Production (tons)

1 Russia Federation 9,696,450

2 Ukraine 8,670,500

3 Argentina 3,671,750

4 France 1,882,450

5 Romania 1,789,330

6 China 1,700,000

7 Bulgaria 1,439,700

8 Hungary 1,374,780

9 Turkey 1,335,000

10 Spain 1,084,300

11 United States of America 924,550

12 South Africa 860,000

Russia followed by Ukraine are harvesting almost half of the world sunflower seed production. The

total sunflower seed production is reaching ca. 35 millions of tons

Data source: data obtained from FAOSTAT (2011).

According to FAO (FAO, 2010), there are some key production parameters which have

to be known by farmers throughout the world:

Sunflowers are grown in warm to moderate semi-arid climatic regions of the world from Argentina to Canada and from central Africa to the Commonwealth of

Independent States (Esmaeli et al., 2012; Onemli, 2012).

Frost will damage sunflowers at all stages of growth. The plant grows well within a temperature range of 20-25C; temperatures above 25C reduce yields and oil

content of the seeds (Thomaz et al., 2012).

Plants are drought-resistant, but yield and oil content are reduced if they are exposed to drought stress during the main growing and flowering periods. Sunflowers will

produce moderate yields with as little as 300 mm of rain per year, while 500-750 mm

are required for better yields (Gholamhoseini et al., 2013; Ghaffari et al., 2012).

Sunflowers adapt to a wide variety of soil, but perform best on good soils suitable for maize or wheat production (Radanielson et al., 2012).



Sunflower plant density of 5-8 plants per m2 is required to form the optimum leaf area for plant photosynthesis. Kernel weight (40-80 g per 1000 kernels) and the

average number of kernels in a sunflower head (1200-1500) are the other most

important yield component (Seassau et al., 2012; Emami-Bistghani et al., 2012).

Sunflower growth depends more on nitrogen than any other nutrient. Due to its deep rooting system, sunflower is able to use nitrogen from soil layers that are inaccessible

to wheat, corn or other field crops. The plant requires a maximum of 150 kg of

nitrogen per hectare to produce a three tons ha-1

yield. Over fertilization may lead to

sunflower lodging. Phosphorous, potassium, boron, magnesium and molybdenum are

also needed to achieve the best yields (Jabeen and Ahmad, 2012; Babaeian et al.,

2011).

The average fatty acid composition of oil from temperate sunflower crops is 55-75% linoleic acid and 15-25% oleic acid. Protein content is 15-20% (Aznar-Moreno et al.,

2013; Ali and Ullah, 2012).

Planting in the Western Balkan countries, Eastern Europe and countries of the Former Soviet Union takes place during March and April (Zheljazkov et al., 2012;

Saleem et al., 2008).

Sunflower has one of the shortest growing seasons of the major economically important crops of the world. Early maturing varieties are ready for harvesting 90 to

120 days after planting, and late maturing varieties 120 to 160 days after planting.

Delayed harvesting causes unwelcome changes in oil quality, with an increase in free

fatty acid content. The seeds are ready to harvest when the heads turn black or brown

and the seed moisture content reaches 10-12%. Grain combines are fairly easily

adapted for the harvesting of sunflower by the addition of a head snatcher (Borbely et

al., 2008).

Depending on climatic and cultivation conditions, yields can vary from as much as 600 to 3000 kg ha

-1; irrigation is a key factor for obtaining high yields (Chigeza et

al., 2013; Khan et al., 2013; Akhtar et al., 2012).

Table 3 shows the oil yields in gallons per acre of oil producing crops, the yields will

vary in different agroclimatic zones. Sunflower produces 98 Gal oil acre-1

. That is enough to

make the sunflower rank twenty-third in that category. Additionally, higher-yielding oil crops

like safflower, mustards and sunflower have significant rotational benefits. For example, deep

safflower and sunflower roots help break up hardpan and improve soil tilth.

4. GROWTH AND DEVELOPMENT

Sunflower is a broadleaf plant that emerges from the soil with two large cotyledons

(Rawat et al., 2010). The emergence will take four to five days when planted an inch deep in

warm soil, but will take a few days longer in cooler soils or when planted deeper. Soil

crusting can make it difficult for the large seedlings to push out of the soil. Sunflowers grow

rapidly, producing large and rough leaves. Current sunflower varieties reach an average

height of six feet, varying between five and seven feet depending on planting date and soil

conditions (Saensee et al., 2012). After reaching their full height and blooming, heads on



commercial cultivars turn downwards, designed to make it harder for birds to eat the seed.

Commercial sunflowers have flowers that are self-compatible for pollination, meaning they

do not require a pollinating insect, although some studies have shown bee pollinators

providing a slight yield boost (de Carvalho and de Toledo, 2008). Some farmers prefer

sowing their rows from north to south so that the capitula can lean into the row space, rather

than bumping against an adjacent plant, causing some seed to fall (Olowe and Adeyemo,

2009).

Table 3. Oil producing crops

Number Crop Scientific name Yield (Gal oil acre-1

)

1 Oil palm Elaeis guineensis Jacq. 610

2 Macauba palm Acrocomia aculeata Jacq. 461

3 Pequi Caryocar brasiliense Camb. 383

4 Buriti palm Mauritia flexuosa L. 335

5 Oiticia Licania rigida Benth 307

6 Coconut Cocos nucifera L. 276

7 Avocado Persea americana Mill. 270

8 Brazil nut Bertholletia excelsa Humb & Bonpl. 245

9 Macadamia nut Macadamia ternifolia F.V. Muell. 230

10 Jatrofa Jatropha curcas L. 194

11 Babassu palm Orbignya martiana Mart. 188

12 Jojoba Simmondsia chinensis Link 186

13 Pecan Carya illinoensis Wangenh. 183

14 Bacuri Platonia insignis Mart. 146

15 Castor bean Ricinus communis L. 145

16 Ghoper plant Euphorbia lathyris L. 137

17 Pissava Attalea funifera Mart. 136

18 Olive tree Olea europea L. 124

19 Rapessed Brassica napus L. 122

20 Opium poppy Papaver somniferum L. 119

21 Peanut Arachis hypogea L. 109

22 Cocoa Theobroma cacao L. 105

23 Sunflower Helianthus annuus L. 98

24 Tung oil tree Aleurites fordii Hemsl. 96

Yields of common energy crops are associated with biodiesel production. This is not related to ethanol

production, which relies on starch, sugar, and cellulose content instead of oil yields.

Experiments have been carried out to improve the growth and development of sunflower

under natural or stress conditions (Gerardo et al., 2013; Nasim et al., 2011; Da Silva et al.,

2012). Naz and Bano (2013) reported that the adverse effects of salt stress on sunflower

growth could be alleviated by foliar application of salicylic acid alone or in combination with

Azospirillum and Pseudomonas inoculations (Table 4). Gholamhoseini et al. (2013) shown

that the application of Glomus musseae and Glomus hoi could be critical in the cultivation of

sunflowers under arid and semi-arid conditions, where water is the most important factor in

determining plant growth and yield. Additionally, Akbari et al. (2011) reported that

inoculating the sunflower seeds with plant-growth promoting rhizobacteria increased the



qualitative and quantitative properties of sunflower significantly, as compared to the control

treatment.

Table 4. Recent uses of the sunflower during the last years; main or alternative uses

make evident the diversity of sunflower

Area Description References

Food

Blends of high linoleic sunflower oil with

selected cold pressed soils.

(Ramadan, 2013)

Production of florets of sunflower. (Liang et al., 2013)

Tocopherols and phytosterols for the human

food market.

(Fernndez-Cuesta et al.,

2012)

Sunflower flour as a rich source of high

quality proteins.

(Levic et al., 2012)

Protein hydrolysis using proteases. (Tavano, 2013)

Animal Feed

Sunflower products fed to finishing pigs. (Gonzlez-Vega and Stein,

2012)

Ingestive behavior and physiological

responses of goats fed with sunflower cake.

(Agy et al., 2013)

Nutritional value of sunflower meal on broiler

chickens.

(Moghaddam et al., 2012)

Potential nutritive value as source of feed for

ruminants in Kenya.

(Osuga et al., 2012)

Energy

Methane production. (Fernndez-Cegr et al., 2013;

Todorovic et al., 2013)

Biodiesel production. (Iriarte and Villalobos, 2013;

Iglesias et al., 2012)

Bioenergy: biotechnology progress and

emerging possibilities.

(Gonzlez-Rosas et al., 2013)

Anaerobic digestion of sunflower oil cake. (De la Rubia et al., 2013)

Oil production. (Spinelli et al., 2012)

Sustainability

Of sunflower cultivation within the EU

Renewable Energy Directive.

(Spugnoli et al., 2012)

Sustainable sunflower processing. (Weisz et al., 2013)

Economic sustainability of sunflower

production.

(Keskin and Dellal, 2011)

Symbiosis and Plant-Growth Promoting Rhizobacteria

Effect of arbuscular mycorrhizal inoculation

on sunflower.

(Naz and Bano, 2013; Audet

and Charest, 2013;

Gholamhoseini et al., 2013)



Table 4. (Continued)

Area Description References

Symbiosis and Plant-Growth Promoting Rhizobacteria

Bacterial inoculation speeds zinc release from

ground tire rubber.

(Khoshgoftarmanesh et al.,

2012)

A strain of Bacillus subtilis stimulates

sunflower growth.

(Lpez-Valdez et al., 2011)

Remediation

Biodegradation of PAHs. (Tejeda-Agredano et al.,

2013)

Plant response to lead. (Doncheva et al., 2013)

Metal accumulation on sunflower. (Mahmood et al., 2013; Hao et

al., 2012)

Fertilization, pesticides and environment

Foliar fertilization with molybdenum. (Skarpa et al., 2013)

Fertilization affects the agronomic traits of

high oleic sunflower hybrid.

(Mohammadi et al., 2013)

Gas exchange in sunflower plants. (Da Silva et al., 2013)

Effect of different nitrogen level on yield

components.

(Rafiei et al., 2012)

Biological control

Encrusting offers protection against

phytotoxic chemicals.

(Szemruch and Ferrari, 2013)

Biological control of Macrophomina

phaseolina on sunflower.

(Ullah, 2010)

Allelopathic effects

On growth of rice and subsequent wheat crop. (Bashir et al., 2012)

On seed germination and seedling growth of

Trianthema portulacastrum.

(Rawat et al., 2012)

Health

In vivo evaluation of an oral health toothpaste

with sunflower oil.

(Schafer et al., 2007)

Health benefits of the sunflower kernel. (Holliday and Phillips, 2001)

5. SUNFLOWER ALLELOPATHY

Sunflower species are allelopathic in nature; as well cultivated sunflower has great

allelopathic potential and inhibits weed-seedling growth of velvet leaf, thorn apple, morning

glory, wild mustard and other weeds (Macas et al., 1998a). Two members of the genus

Helianthus contain a great quantity of allelopathic compounds. H. annuus is well known for

its allelopathic compounds, including sesquiterpene lactones, heliespirones A, annoionones,

helibis-abonols and heliannols (Macas et al., 1998b). Heliannols A, D and E have special

relevance due to high phytotoxic activity (Macas et al., 1999).



Figure 1. Some molecular structures of allelopathic compounds presents in sunflower cultivars. A)

Annuithrin (sesquiterpene lactone) or Niveusin C, a growth inhibitor. B) Furanoheliangoline, a

biologically active molecule. C) Germacranolide, a toxic sesquiterpene lactone (a potent feeding

deterrents).

Helianthus tuberosus contains helian-gine and H. annuus contains a sesquiterpene

lactone; a heliangolide [Annuithrin or Niveusin C (Figure 1A)] (a growth inhibitor);

furanoheliangolide [(Figure 1B) a biologically active]; three additional sesquiterpene

lactones: the known compound niveusin B, a germacranolide (Figure 1C) (the tifruticin-type);

a 3-ethoxy-niveusin B; an ethoxyheliangolide (Spring et al., 1982) and coumarins (only

accumulate in healthy sunflower plants as a response to the variation in environmental

conditions that affect field-grown plants). In sunflower, it was reported that the concentrations

of scopolin exceeded those in both infected and uninfected plants (Gutirrez-Mellado et al.,

1996).

Scopoletin have been described as phytoalexins and allelopathic compounds, being

accumulated in response to fungal and parasitic plant infection, insect attack, mechanical

injury and treatment with abiotic elicitors such as sucrose and CuCl2, and plant hormones;

besides scopoletin has also been shown to have a physiological activity, including the

promotion of stomatal closure in sunflower and inhibition of bud growth in pea at very low

concentrations (Gutirrez-Mellado et al., 1996).

Annuithrin was tested using a bioassay with Avena straight growth test. The addition of a

concentration range from 50 to 180 M resulted in a linear reduction of growth between 10

and 90%. In fact, annuithrin was shown to have antibacterial qualities. However, fungi and

yeast were either less inhibited or not inhibited (minimal inhibitory concentration, MIC 45 g

mL-1

on Bacillus brevis; MIC 90 g mL-1

on Proteus vulgaris; MIC 90 g mL-1

on

Eremothecium ashbyi; Macas et al., 1996). In addition, in vivo DNA and RNA synthesis in

cells of the ascitic form of Ehrlich carcinoma was drastically reduced by annuithrin (at an

annuithrin concentration of 20 g mL-1

about 50% inhibition of DNA synthesis and about

75% inhibition of RNA synthesis) (Spring et al., 1981).

It is well known that there are examples of allelopathic cover crops being used for weed

management in other crops, as well as other cultural methods to employ allelopathy (Duke,

2010). However, there are still no cultivars of crops being sold with allelopathic properties as

a selling point (Cheema and Khaliq, 2000; Tesio and Ferrero, 2010). Enhancement or

impartation of allelopathy in crops through the use of transgenes could eventually be used to



produce such a cultivar. The study of allelopathic crops appears to have a bright future,

especially if the scientists can translate the cutting-edge research into technologies that will

reduce the reliance on synthetic herbicides, pesticides, and crop protection chemicals. Tesio

and Ferrero (2010) reported that the use of allelopathic traits from crops or cultivars with

important weed inhibition qualities, together with common weed control strategies, can play

an important role in the establishment of sustainable agriculture. It has to be noted that

allelopathy may also be another component of desired improved weed management. It will

not solve all weed problems in any field, but may help considerably to reduce the population

of weeds in the fields (Labrada, 2008).

6. PHYTOREMEDIATION WITH SUNFLOWER

Phytoremediation consists of mitigating pollutant concentrations in contaminated soils,

water, or air, with plants able to contain, degrade, or eliminate contaminants and its

derivatives (Malaviya and Singh, 2012). H. annuus is a plant with not only food and energy

values, but also with phytoremediation potential (Seth et al., 2011; Mukhtar et al., 2010). It is

one of the most widely studied plants for heavy metal phytoremediation (Kara et al., 2013).

However, it is well known that sunflower is able to contain, degrade or eliminate metals

(Chen et al., 2012; Ker and Charest, 2010; Lee and Yang, 2010), polycyclic aromatic

hydrocarbons (Tejeda-Agredano et al., 2013; Gan et al., 2009) and polychlorinated biphenyls

(Fiebig et al., 1997) from soil or water. Investigations with H. annuus have revealed that

several heavy metals, including lead, cadmium, copper, zinc and cobalt, accumulate at high

concentrations in shoots as well as in roots. Heavy metal uptake is minor in seeds than in

roots and shoots. However, few attempts have been made to use plant-growth promoting

rhizobacteria to facilitate phytoextraction and cadmium uptake in H. annuus planted in

cadmium-contaminated soil (Prapagdee et al., 2013). Sunflower is a documented metal

accumulator and its growth on contaminated soil for simultaneous remediation and further

energy production has been studied (Marques et al., 2013; Madejon et al., 2003). The good

tolerance of sunflower toward pollutants coupled with an increased accumulation/degradation

capacity might contribute to an efficient removal of pollutants from soil and water. Clearly it

is not an easy job, thus scientists of multidisciplinary areas have to work hard. Additionally,

there is a lack of knowledge concerning the pollutants accumulation and antioxidant

responses during the growth and development of sunflowers.

7. SUNFLOWER AS A RENEWABLE ENERGY SOURCE

Thousands of years ago, people in many regions throughout the world began to process

vegetable oils, utilizing whatever food stuffs they had on hand to obtain oils for a variety of

cooking purposes. The Chinese and Japanese produced soy bean oil as early as 2000 B.C.,

while southern Europeans had begun to produce olive oil by 3000 B.C. In Mexico and North

America, sunflower seeds were roasted and beaten into a paste before being boiled in water;

the oil that rose to the surface was skimmed off (FAO, 2010). During the last decade, an



increased attention would be observed being paid on the use of sunflower as renewable

energy source.

Oilseed sunflower is quickly gaining popularity as a feedstock crop for biodiesel because

it shares several positive agronomic features with other common oil crops such as canola and

soybean; yields well in a variety of conditions, and can be grown easily and profitably at both

small farm and large field scales. It is well known that a number of crops can be used for both

food and bioenergy production such as sunflower (Kibazohi et al., 2012). Under some

circumstances, the potential exist to develop bioenergy systems that allow for synergies

between food and energy production. Integrated food and energy systems could produce food

crops while simultaneously addressing energy needs (Bogdanski et al., 2010).

There is a trend world-wide to grow crops in short rotation or in monoculture (such as

sunflower), particularly in conventional agriculture (Bennett et al., 2012). This practice is

becoming more prevalent due to a range of factors including economic market trends,

technological advances, government incentives, and retailer and consumer demands. Land-

use intensity will have to increase further in future in order to meet the demands of growing

crops for both bioenergy and food production, and long rotations may not be considered

viable or practical. Notwithstanding, evidence indicates that crops grown in short rotations or

monoculture often suffer from yield decline compared to those grown in longer rotations or

for the first time (Zambrano-Navea et al., 2012). Numerous factors have been hypothesized as

contributing to yield decline, including biotic factors such as plant pathogens, deleterious

rhizosphere microorganisms, mycorrhizas acting as pathogens, and allelopathy or autotoxicity

of the crop, as well as abiotic factors such as land management practices and nutrient

availability (Sun et al., 2011). This section identifies gaps in our understanding about the

energy production of biomass and the interaction of the ecosystems. Additionally, it has to be

remembered that each bioenergy development projects have to include side effects elsewhere

in order to shape a sustainable future.

CONCLUSION

Sunflower was domesticated in eastern North America and since 3000 B.C. this crop was

bred by natives. Thenceforth a wide range of uses of sunflower have been reported

throughout the world. Sunflowers are a permanent source of food, oilseed and biofuels

because they are well adapted to a variety of conditions and often require fewer agricultural

inputs than other more common crops, while under some circumstances, the potential exist to

develop bioenergy systems that allow for synergies between food and energy production.

Because the sunflower has several potential markets, it is a good choice for growers in both

small and large scales. However, scientific, technical or agricultural projects linked with

sunflower have to include environmental side effects such as pollution, greenhouse gases

emissions, salinization, or energy consumption elsewhere in order to shape a sustainable

future.



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In: Sunflowers ISBN: 978-1-63117-347-9

Editor: Juan Ignacio Arribas 2014 Nova Science Publishers, Inc.

Chapter 2

FLORAL BIOLOGY OF SUNFLOWERS:

A HISTOLOGICAL AND PHYSIOLOGICAL ANALYSIS

Basudha Sharma, Rashmi Shakya and Satish C. Bhatla* Laboratory of Plant Physiology and Biochemistry, Department of Botany,

University of Delhi, Delhi, India

ABSTRACT

The development of sunflower inflorescence can be considered under three phases,

namely inflorescence initiation, floret development and anther formation. Floret

primordia appear at the rim of the receptacle where ray or disc florets are generated. Disc

florets are arranged in Fibonacci series whereby a spiral pattern emerges as new florets

arise in rows of bumps consisting of a bract and a floret. Floral morphogenesis in

sunflower occurs according to the ABC model, whereby genes of the MADS box are

activated. Anthesis of disc florets is a phytochrome-mediated response and is also

modulated by plant hormones, such as auxins. The disc florets are hermaphrodite and

protandrous in nature, whereas the ray florets are sterile, incomplete and have an

attractive, fused and flag-like corolla. Stigma in sunflower is semi-dry in nature,

producing lipid rich exudates in the crevices of the adjacent papillae. Stigma undergoes

physiological maturity with the passage of development from bud, staminate and, finally

to the pistillate stage. The production of extracellular lipid rich secretions is initiated at

the staminate stage of stigma development and increases at the receptive stage through

the availability of elaioplasts and endoplasmic reticulum network in the basal regions of

the papillae. Transfer cells, earlier identified only in the wet type of stigma, are also

present in the transmitting tissue of sunflower stigma. Neutral esters and triacylglycerols

(TAGs) are the major lipidic constituents in pollen grains and stigma, respectively.

Lignoceric acid (24:0) and cis-11-eicosenoic acid (20:1) are specifically expressed only

in the pollen coat. Similar long-chain fatty acids have earlier been demonstrated to play a

significant role during the initial signalling mechanism leading to hydration of pollen

grains on the stigma surface. Lipase activity is expressed both in the pollen grains and

stigma papillae. Stigma exhibits a better expression of acyl-ester hydrolase activity the

pollen grains. Specific expression of lignoceric acid (24:0) in the pollen coat and

* Corresponding author: Professor S.C.Bhatla; E mail: [email protected].


Basudha Sharma, Rashmi Shakya and Satish C. Bhatla 20

localization of lipase in pollen and stigma are likely to have possible roles during pollen-

stigma interaction. During the course of stigma development in sunflower, a correlation is

evident in the accumulation of reactive oxygen species (ROS), nitric oxide (NO) and the

activities of ROS scavenging enzymes [superoxide dismutase (SOD) and peroxidase

(POD)]. Mn-SOD (mitochondria localized) and Cu/Zn-SOD (cytoplasmic) exhibit

differential expression during the staminate stage of stigma development. An increase in

total SOD activity at the staminate stage is followed by a

sunflowers 2014

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