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UNIVERSITI PUTRA MALAYSIA
STUDIES ON GERMINATION AND SEEDLING GROWTH OF NEOBALANOCARPUS
HEIMII (KING) ASHTON
SITI RUBIAH ZAINUDIN
FH 1990 10
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STUDIES ON GERMINATION
AND SEEDLING GROWTH OF NEOBALANOCARPUS
HEIMI I (KING) ASHTON
By
SITI RUBIAH ZAINUDIN
Thesis Submitted in Ful f ilment o f the
Requirements for the Degree of Master of Sc ience in the Faculty of Forestry
Univers it i Pertanian Malaysia
October 1990
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ACKNOWLEDGEMENTS
I wish to expres s my most sincere and deepest thanks
to my supervisor, Dr . Lim Meng Tsai for his invaluable guidance ,
advice , encouragement , constructive criticisms and suggestions
throughout the pro j ect . To Associate Professor Mohd . Basri bin
Hamzah, my co-supervisor , I also wish to record my appreciation
for the guidance rendered .
Grateful acknowledgement is due to the Dean , Faculty o f
Forestry , Associate Professor Dr . Kamis Awang for providing the
f acilities . Sincere thanks are also due to the sta f f o f the
Faculty of Forestry , especially to Encik Haidzir bin Abdul
Wahab and Encik Yusri bin Abu for their invaluable assistance .
I a l so wish to express my special thanks to Cik Norizah
for her patience in typing this manuscript and to a l l who have
helped in one way or another in the completion of this pro j ect .
Fina l l y , I would like to dedicate this work to my family ,
husband and son Syed Hafiz , without whose love and devotion
this study would not have completed .
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TABLE OF CONTENTS
Page
ACKNOWLEDGEMENTS ii
LIST OF TABLES vi
LIST OF FIGURES viii
LIST OF PLATES x
LIST OF ABBREVIATIONS xi
ABSTRACT xii
ABSTRAK xiv
CHAPTER
ONE GENERAL INTRODUCTION 1
Status of Forestry in Malaysia 1
Obj ective o f Study 3
Literature Review 3
Chengaf Distribution and Regeneration 3
Seed Germination 4
Seedling Growth 9
Mycorrhiza and Soil 1 0
Light Requirement 13
Photosynthetic Characteristics of Tropical Forest Trees 1 5
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TWO SEED GERMINATI ON
Introduction
Material and Methods
Page
18
19
Results and Discussion • • • • • • • • • • • • . • • . • • . . • • • • 2 2
THREE: GROWTH
I ntroduction
Material and Methods • • • • • • . • • • • • • . • • • • • • • • . • . •
30
31
Results and Discussion • • • • • • • . . . • • • • • • • • • • • • • • 32
FOUR: PHOTOSYNTHESIS
Introduction 4 5
Material and Methods • • . • • • • • • • • • • . • • • • • • • • • • . • • 46
Resu lts and Discussion • • . • • • • • • • • • • • • • • • • • • • • • • 4 7
FIVE: MYCORRHI ZA AND SOIL
I ntroduct ion 53
5 5
S I X:
Material and Methods
Results and Discussion • • • • • . • • • • • • • . • • • • • • • • • • • 56
FIELD OBSERVATIONS
I ntroduction 62
study S ites • . • . • . . . • . . • • • • . . . . . • . . . . . • . • • • • • • • • 63
Material and Methods • • • • • • • . . • • • • • • • • • • • • • • • • • • 64
Results and Discussion • • • • • • • • • • • • • • • • • • • • • • • • • 65
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Page
SEVEN: GENERAL DISCUSSION , CONCLUSIONS AND RECOMMENDATIONS 73
REFERENCES 7 9
VITA 91
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Table
1
2
3
4
5
6
7
8
LIST OF TABLES
Photosynthetic Rates of Tropical Species Measured at Light S aturating Intensities
at Temperatures between 2 20C and 300C • • • • • • •
E f fects of Light Treatment on Chengal seedlings ( Relative Growth Rates : Height ,
Page
16
Number of Leaves and Nodes ) . . ............ . . . 33
Correlation Coef fic ients of Seed Weight with Height , Number of Leaves , Number o f Nodes and Collar Diameter with Different Light Treatment • • • . • • • • • • • • • • • • . • •
E ffects of Light Treatment on Relative
Growth Rate for Collar Diameter , Leaf Area , Dry Weight of Lea f , Stem and Root of Chengal Seedlings • • . . • • • • • • • • • • • • • • • • . • •
Maximum Photosynthetic Rates of Chengal
Seedlings Grown under Different Light
33
35
Regimes • • • • • • • • • • • . • • • • • • • • • • • • . • • . . • . . • • • • 49
Maximum Photosynthet ic Rates of Some
Dipterocarp Species • • • . • • • • • . . • • • • • • • . . . • •
Mechanical Analysi s , Percentage Organic
Carbon and pH of Soil From the Ampang
Forest Reserve , Pasoh Forest Reserve and
5 1
UPM Nursery . • • • • . • . • • . • • • • • • . • • . . • • • • • • . . • 5 9
Distribut ion of Chengal Seedlings and
Saplings in Pasoh Forest Reserve ( no. per sq . m ) Around the Parent Tree . • . • • • • . • . • • • •
vi
68
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9 Distribut ion of Chengal Seedl ings in Ampang Forest Reserve ( no . per sq . m ) Around the Parent Tree • • • . . • • • • • . • . . . . . . . .
vii
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70
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Figure
1
2
3
4
5
6
7
8
9
1 0
11
LIST OF FIGURES
Distribution of Neobalanocarpus heimii
( Chenga1 ) in Peninsular Malaysia • • • • • • • • . • . .
Progres s of Seed Germination under Different L ight Condition
in Chengal
The Rel at ionship Between Seed Length And
Percentage Germination of chengal • . • . • . . • • . . .
The Relationship Between Seed Weight And
Percentage Germinat ion of Chengal . . . . . . • • . . • .
Longitudinal Section of Chengal
( Percentage Area Stained by Iodine ) Seeds
Col l ar Diameter Growth of Chengal Seedl ings under Different Light Intensit ies . . • . . . . . . . . .
Leaf Area of Chengal Seedl ings under
Different Light Intens ities . . . . . . . . . . . . . . . . . . .
Dry Weight of under Different
Leaf of Chengal Seed l ings Light Intensities • • . • • • • • • •
Stem Growth of
Different Light
Root Growth of Different Light
Weight of Seedl ings Intensit ies
Shoot grown
After
Chengal Seedl ings under Intensit ies . . . . . . . . . . . . . . . .
Chengal Seedlings under Intensit ies . . . . . . . . . . . . . . . .
and Root of Chengal under Different Light 12 months • • • . . . . . . . . . . . .
viii
Page
5
2 3
2 5
2 6
2 8
34
36
37
38
39
40
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1 2
13
14
Photosynthetic Response curve of Chengal Seedl ings Grown under Dif ferent Light Intensities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Relationship between Density of Chengal
Seed l ings , Saplings and Distance from the Parent Tree in Pasoh Forest Reserve • . • • . . . .
Relationship between Density of Chengal
Seedl ings and Distance from the Parent Tree in Ampang Forest Reserve • • . . . • . • • • • • . . • • . . . .
ix
Page
48
6 6
69
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P l ate
1
2
3
LIST OF PLATES
Chengal Seeds
Cross Section of Chengal Root Showing the Mant le and Hart ig Net ( Magnif icat ion 44x ) • • • . . • • • • • . • • . • . . . . • • •
Chengal Roots ( Ectornycorrhiza is
Brown in Colour ) ( Magnificat ion 44x ) • • • • • • • • • • • • • • • • • • • . . •
x
Page
2 4
5 8
5 8
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LIST OF ABBREVIATIONS
a . s . l = Above Sea Level
FAA = Formalin Acetic Acid
FRIM = Forest Research Institute of Malaysia
IRGA = Infra-Red Gas Analyser
PAR = Photosynthetic Act ive Radiat ion
RGR = Relative Growth Rate
RLI = Relative Light Intensity
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Abstract o f thes is submitted to the Senate o f Universiti Pertanian Malays ia in ful f ilment of the requirements for the degree of Master of Science .
Supervisor
Faculty
STUDIES ON GERMINATION AND SEEDLING GROWTH
OF NEOBALANOCARPUS HEIMI I ( KING ) ASHTON
By
SITI RUBIAH ZAINUDIN
June 1990
Dr . Lim Meng Tsai
Forestry
Neobalanocarpus heimii ( King ) Ashton locally known as
chengal is endemic to Southern Thailand and Peninsular
Malay s ia . It is an important genera as it produces wood
that has become the standard by which many others are compared .
I t i s f a i r l y w i d e l y d i stributed but f o un d i n s c att ered
occurrence and at low numbers . This proj ect focuses on the
germinat ion , seedl ing growth , presence of mycorrhiza and
photosynthetic rates under dif ferent l ight condit ions. Soil
analys i s and f ield observat ion of seedl ing distribut ion around
the parent tree were also conducted . Germination o f chengal
was more rapid at lower l ight intensities than at higher l ight
intensities. The germinat ion percentage is also influenced by
seed size i . e . , the bigger the seed the better the germinat ion .
Treatment means for plant height and leaves showed no
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s igni f icant difference among treatments . However , the number
of nodes was s ignif icant in all treatments . Seed weight was
pos it ively correlated to height , number of leave s , number o f
nodes and col lar diameter at all l ight intensit ies . The
b iomass and total area of leaves per plant were highest at 5 5 %
RLI . Chengal grows on sandy loam soil with very low organic
carbon content and low pH . Ectomycorrhiza was found in roots
at the age of 2 months . The highest photosynthet ic rate o f
4 . 37 micro mol m-2 s-l was attained by plant grown at 5 5 % RLI
at the saturating photon flux of 400 micro mol m-2 s-l . P l ants
g rown at h igher and l ower l ight inten s it i e s had l ower
photosynthet ic rates of 1 . 30 micro mol m-2s-l and became
saturated at 100 micro mol m-2 s-l . After a seed fal l , the
number of seeds and seedlings around the parent tree is high .
However , there is correspondingly a high mortal ity rate with
high dens ity; the number fall ing to one quarter the total
number after a year . This study shows that the germinat ion
of chengal was best at 2 5% RLI and optimum growth of seedl ings
was attained at 5 5 % RLI.
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Ab s t r a k t e s i s y ang d i kemukakan kepada Senat Un iver s i t i Pertanian Malays ia sebagai memenuhi keperluan untuk Ij azah Master Sains .
KAJIAN MENGENAI PERCAMBAHAN DAN PERTUMBUHAN ANAK BENIH NEOBALANOCARPUS HEIMII ( KING ) ASHTON
OLEH
SITI RUBIAH ZAINUDIN
June 1990
Penyelia Dr . Lim Meng Tsai
Fakulti Perhutanan
Neo b a l anocapu s h e i m i i ( K ing ) A s h t o n d i ke na l i s e b ag a i
c he ng a l a d a l ah endemik kepada Se l a t a n T h a i l and d an
Semenanj ung Malaysia . Ia merupakan salah satu kayu yang
men j adi satu piawai untuk perbandingan spes ies kayu keras l ain.
pro j ek i n i membe ntangkan k a j i a n ke a t a s per c amb a h a n ,
pertumbuhan anak benih , kehad i r a n m i ko r i z a d a n kadar
fotos intesis di bawah keadaan cahaya yang berbeza . Ana l i s is
tanah dan pemerhatian di lapangan terhadap taburan anak benih
di s eke l i l ing pokok i nduk j uga d i j a l ankan . P e r c amb ah a n
daripada chengal adalah lebih cepat pada intens it i cahaya yang
rendah iaitu 2 5 % RLI berbanding dengan intens it i cahaya yang
t ingg i . Peratus percambahan j uga dipengaruhi oleh saiz b i j i
benih iaitu bij i benih yang lebih besar memberikan percambahan
y an g l eb i h b a i k . P u r a t a r awat an untuk ket i n g g i a n
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pokok dan daun menunj ukkan tiada perbezaan bererti d i antara
rawatan . Walau bagaimanapun, bilangan nod adal ah berert i di
antara semua rawatan cahaya . Berat bij i benih menunj ukkan
perhubungan posit i f dengan ket inggian , bilangan dau n , bilangan
nod dan garispusat kolar pad a semua rawatan cahaya . B iomas
dan j umlah keluasan daun per pokok adalah pal ing t inggi pada
5 5 % RLI . Chengal t umbuh d i t anah l iat berp a s ir dengan
kandungan organik karbon dan pH yang rendah . Ektomikoriza di
dapati didalam akar pada umur 2 bu lan . Kadar fotosintesis yang
pal ing t inggi , 4 . 37 micro mol m-2 s- l dicapai oleh tumbuhan
pada 5 5 % RLI pada keamatan ketepuan foton f luk 400 micro mol
m-2 s- l . Tumbuhan pad a intensit i cahaya yang lebih t inggi dan
rendah mempunyai kadar fotosintesis yang rendah iaitu 1 . 30
micro mol m-2 s-l dan menj adi tepu pada 100 micro mol m-2 s- l .
Selepas bij i benih gugur I terdapat banyak bij i dan benih
benih yang tumbuh mengelil ingi pokok induk . Bilangan
anak benih menurun dengan cepat dengan kenaikan j arak dari
pokok induk . Terdapat kadar kemat ian yang t inggi bilamana
ketumpatan ad a l ah s angat t i ngg i . O l e h i t u b i l angan
berkurangan kepada satu perempat selepas setahun . Kaj ian ini
menunj ukkan percambahan bij i benih chengal adalah pal ing baik
pad a 2 5% RLI dan anak benih chengal memerlukan keamatan cahaya
5 5 % RLI untuk pertumbuhan yang opt imum .
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CHAPTER ONE
GENERAL INTRODUCTION
status of Forestry in Malaysia
For the past three decades , tropical forests dominated by
dipterocarps have been exploited on an increasing scale due to
popular world demand for hardwood . Malaysia has 1 7 . 37 mil l ion
hectares of dipterocarp forests which contributed
s igni f icantly to industrial development , earning of foreign
exchange and soc io-economic changes ( Thang , 1986 ) . In view o f
the rapid depletion of the forest resources , ef forts to meet
future demand for t imber has forced the Forestry Department o f
Peninsular Malaysia to embark o n a Compensatory Plantation
Programme in which fast growing soft hardwood species such as
Acacia mangium and Gmel ina arborea have been successfully
planted .
I n a n att empt t o pursue a s u s t a i n a b l e y i e l d po l ic y ,
dipterocarp forests have to be regenerated after logging with
dipterocarps , either naturally or art i f ically by enrichment
planting . Natural regenerat ion of hill dipterocarp forest i s
general ly poor ( Wyatt-Smit h , 19 63; Burgess , 1976 ) . This i s due
to the variabil ity of terrain and stocking of economic spec ies .
Efforts to improve the regenerat ion status of these forests by
extending the natural distribution range of the more product ive
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species through planting have not been very succes s ful ( Tang
and Wadley , 1 9 7 6 ) . T h i s c o u l d b e due t o t he l im it e d
understanding of the ecophysiology of the species.
Neobalanocarpus heimii ( King ) Ashton is l isted as one o f
the ten endangered species i n Malaysia ( Kiew et al . , 1985 ) . Due
t o it s s t re ngth and durab i l it y it i s o f g r e at e c o n o m i c
importance and demand for chengal local ly is high ( Maskayu , Vol
I, Jan , 1989 ) . It produces a strong and naturally durable wood
used for power-l ine posts , building boat s , carriage-bodies for
lorries and buses and frames for doors and windows .
Growth measurements of 319 planted chengal trees at Forest
Research Inst itute Malays ia at 22 years old has a mean girth o f
3 7 c m and a mean annual increment of 1 . 70 cm . Measurement o f
5370 trees i n Negeri Sembilan and Pahang gave annual increment
of 1 . 0 cm , so that in it s natural state , it probably takes
about 80 years to reach a girth of 153 cm, and 1 2 0 years to
reach a girth measurement of 230 cm ( Edwards , 1930 ) .
In the natural environment big trees of chengal are
usual ly found with a marked lack of suitable representat ion o f
sma l l and intermediate s izes ( Symington , 1943 ) . To ensure
s uc ce s s f u l regenerat ion o f cheng a l , t here is a need f o r
information o n its distribution and requirements for growt h .
These would help i n the management and manipulation o f the
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forest to favour the regenerat ion of chengal in accordance to
its tolerance .
Objective of study
The unusual distribution of chengal suggest that the
ecological requirements for growth and regenerat ion may be very
specific . As a result of this , these studies were conducted to
explain how chengal estab l ishes itsel f . Parameters inc lude the
r e s po n s e of c heng a l s e ed l i ng s t o l ight , me a s urement o f
photosynthetic rates , presence o f mycorhizza and soil analys i s .
Literature Review
Chengal Distribution and Regeneration
Chengal is found throughout Peninsular Malaysia except
for Perlis and Malacca ( Symington , 1943 ) ( Figure 1 ) . It i s
found i n the lowland forests below 600m a . s . 1 and seems to
prefer sandy soil with l ittle laterite . It is also found in
low densities , Le less than 5 trees/ha ( symington , 1943 ) .
According to Symington ( 19 43 ) , numerous seedl ings and sapl ings
can somet imes be found beneath the parent tree . Chengal is a
l arge tree that can attain a diameter of up to 4 metres and
height of 42 metres to the f irst branch . Reported to be s low
growing , it does not achieve a maximum rate of growth unt i l it
is about 75 cm in girt h . However , measurements o f certain
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plants have indicated that under favourable conditions growth
may be greater than mentioned above ( Symington , 1 9 4 3 ) .
According to the Forest Research Inst itute of Malaysia
Phenology Reports ( 1989 ) chengal flowers and fruit s every year .
Flowering does not seem to occur in mast with f l owering o f
other species of Dipterocarps ( Ng , Pers . comm . , 1 9 89 ) . I t
begins to bear flowers and fruits while sti l l young , and
planted trees have been known to bear fruits at less than ten
years old ( Symington , 1943 ) .
Chengal seeds are heavy and wingless and hence dispersal
i s restricted. Seeds are dispersed by animal s and by rol l ing
down h i l l s lopes. Because of its l imited dispersal mechanism
seedl ings are usually found in high densities beneath the
parent trees ( Symington , 1943 ) . According to Symington ( 19 43 ) ,
although these seedl ings are able to survive for long periods
under dense shade , light is still required for better growth .
Seed Germination
Seeds of many forest trees of the humid tropic s germinate
soon after dispersal . Germination of dipterocarps seeds is
rapid and Ng ( 19 78 ) concludes that rapid germination gives the
plants the advantage of escaping seed predation. For example ,
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04°
2
1 QOO[
, '.
.- ..... " .
. . · /1···········
.. ·
;' .. _' . .
'O" . .� i ; .' � ..... ' . ."
•
�"'" . , . ,
• . .... ,�
Ke'; • = chenga i
o ,
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t
Figure 1 Distribution of Neobalanocarous he�mii (Chengali in P�ntnsular M�laysia
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6
Dryobalanops aromat ica germinates within 1-2 weeks while Hopea
odorata , Shorea material i s , Shorea
lucida within 1-3 weeks .
maxima and Parashorea
The viab i l ity of seeds is af fected by various factor s .
Amongst them are the method of collection , stage of seed
maturation , infestation by pests, moisture content of seeds and
storage condit ions . Yap ( 1981 ) compared seeds o f Shorea
assamica col lected from the ground with those from the tree and
found l ittle dif ference in germinat ion percentage . Tang ( 19 71 )
has a l so indicated that selected good seeds of Shorea curt i s i i
from ground col lection are j ust as viable as those from
trees . However , ground col lected seeds of unknown
durat ion can show very low viabl ity ( Yap , 198 1 ) .
The viabil ity of seeds is also affected by the dif ferent
stages of maturation . Barnard ( 19 5 0 ) found that seeds with
green seed coats and brown wings germinate better than seeds
w i t h brown seed c o at s . A s l ight l y h i gher germi nat i o n
percentage was observed in the fruits o f Shorea materialis with
brown wings over those with part ially brown wings ( Yap , 1 9 8 1 ) .
In Shorea sumatrana , fruit s with green pericarps germinated
better than those with brown pericarps .
The e s t ab l i s hment o f c ert a i n d ipterocarp spe c ie s i s
critical where the product ion o f viable seeds i s very low . The
m a j or pe s t s o f seeds wh i l e s t i l l on t re e s are weevi l s
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7
( Al c idodes spp . and Nanophytes spp . ) while Scolit id beetles
( Poec i i l ips spp . ) infest fal len seeds ( Dalj eet-s ingh , 19 74 ) .
Their infestat ion on Shore a curtisii reduces viable seeds to as
low as 1 . 6% of the total seeds produced ( Burgess , 1972 ) .
Mature dipterocarp seeds are rich in moisture and quickly
die when dehydrated ( Jansen, 1971; Tang , 1971; Tang and Tamar i ,
19 73; Sasaki , 1980; Yap , 1981 ) . These are typical recalc itrant
seeds which die quickly through dehydrat ion and can only be
stored wet ( Robert s , 1973 ) . Moisture is essent ial for
germination and seed can become non-viable in seven days if
allowed to dry . Corbineau and Come ( 1986 ) found that seeds o f
Shorea roxburghii and Hopea odorata are typical recalcitrant
seeds . The seeds cannot tolerate desiccat ion and die when
their mean moisture content drop by 20% ( dry weight basis ) .
Shorea roxburghii and Shorea ovalis seeds lose viab i l ity
at 20% moisture content ( Sasaki , 1980 ) while the seeds o f
Dryobanalops aromat ica are damaged at a moisture content below
35% ( Tamar i , 1976 ) .
Yap ( 19 8 1 ) proved that stored fruits of dipterocarps also
lose their viabi lity rapidly, and different species responded
to different storage condit ions . It was found that Dipterocarpus
baudii fruits with intact wings stored in closed bags at 1 40C
for 1 7 days was able to retain viability o f over 60% while
fruits with wings removed germinated poorly after storage in
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8
the same condition . Dipterocarpus oblongifolius however , was
only able to retain viab i l ity of over 60% when stored in bags
with the air replaced with nitrogen for 60 days at room
temperature . Seeds of Shorea roxburghii and Hopea odorata
germinate very e a s i ly at high t emperatures w i t h opt i m a l
temperature close t o 300C o r 3SoC . According to Corbineau and
Come ( 1986 ) , germination was possible at 40oC , but this
temperature did not al low seedlings growth . Hopea odorata
seeds however , germinated perfectly down to lSoC , whereas
t h o s e o f Shorea roxburgh i i germ inat ed very poo r l y at
temperature below lSoC ( Corbineau and Come , 1986 ) .
Species found in c losed shaded habitats of moist temperate
and tropical environment are found to have l arger seeds than
those that regenerate in open , secondary habitat s ( Foster ,
1986 ) . It is general ly considered that large seeds enhance
seedl ings survival at low l ight intensities .
Large seeds may have nutrients , energy and water reserves
t o a c c o mmodat e metabo l i c requ irement s dur i n g per i o d s o f
dormancy . These reserves could allow seed survival unti l
c on d i t i o n s o f germ inat ion a n d growt h o f seed l i n g s are
favourable ( Foster, 1986 ) . Large seeds contain l arge amounts
o f secondary compounds that are used in the defence of the
seeds and seedl ings • Freeland and Janzen ( 1974 ) found that
s ome s e e d s cont a i n s e c ondary compounds l ike a m i n o a c id
analogues , alkaloids and phenolic compounds of up to 10% of
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9
it s dry mass . Seeds of the shade-tolerant neotropical nutmeg ,
Virola surinamensis contain 1 5 . 4% dry mass of soluble tannin
( Howe and Vande Kerckhove , 1981 ) .
Large seeds also provide energy and nutrients necessary
for growth . Grime and Jeffrey ( 19 6 5 ) found a strong positive
correlat ion between seed s ize and max imum height of seedl ings
in nine temperate zone tree species which germinated under
shade . S imilar results were obtained by Howe and Richter
( 1 9 8 2 ) and Howe et al . , ( 19 8 5 ) for the s h ade- t o l e r ant
neotropical nutmeg , Virola surinamensis.
Seedling Growth
S hade i s an import ant prerequ i s it e for d ipte r o c arp
estab l ishment ( Whitmore, 1984 ) . Several species such as
Shorea leprosula , Parashorea tomentella and Dipterocarpus
s t e l l atus require s ome s h ade for seed l ings e s t ab l i s hment
( Nicho l son, 1960 ) .
General ly , most species produce a large populat ion o f
s ee d l in g s a fter a mas s f ru it ing a l t hough t he r e i s h i g h
mortal ity i n the f irst few months. Liew and Wong ( 19 73 ) in
their study on Parashorea tomentel la in Sabah , found that
seedl ing survival rates decl ined from 4 1 . 9 5 % in the first year
to 1 6 . 90% in the fourth year . The survival of dipterocarp
seedl ings , however , can differ cons iderably between species