Seed Development and Maturation

podocarpus and taxads, and wings in conifers develop during the later
stage of seed maturation.
Some main differences between angiosperms and gymno
sperms with respect to pollination,fertilization and seed development
are summarised in Table 1.

Table 1.Differences between angiosperms and gymnosperms

with respect to polination,fertilization and seed development
Angiosperms Gymnosperms
Male and female organs in the same Male and female
ower (hermaphrodite) or in separate organs always borne
lowers separately
Pollen produced in anthers in male or Pollen produced in
hermaphrodite flowers micro-sporngia or
micro-sporophylls
Dvules enclosed in pistils or carpels Ovules borne open
on the megasporophylls
Pollination by wind, water oranimals Pollination by wind

Pollen deposited on the stigma Pollen deposited on a

pollination droplet at the
tip of the intugements
Megasporogenesis usually prior to Megasporogenesis
pollination (meiosis and formation of
egg cell) occurs after
pollination

Fertilization occurs soon after pollination Fertilization may occur

several weeks or months
after pollination. Pollen
remains dormant for a
prolonged period inthe
pollen chamber or
archegonial chamber
prior to fertilization
 10 An Introduction to Tree Seed Technology
Double fertilization; one sperm cell
form the
unites with the egg cell to the central
zygote, the other fuses with
cellto form triploid endosperm
Endosperm made up of triploid tissue
lof both maternal and paternal genetic
lorigin
Usually short reproductive cycle
Ovaries and ovules small at the time
of pollination
Unfertilized ovules usually abort
Single fertilization, sperm
fuses with egg cell to
form zygote
Endosperm made up of
haploid maternal tissue
Often long reproductive
cycle
Seed- bearing structures
and ovules often of near
mature size at the time of
pollination
Unfertilised ovules often
remain as 'empty seeds'
 CHAPTER-5

SEED DORMANCY

Seed dormancy is the physiological state in which a viable

seed does not germinate ,even in the presence of favourable environ
mental conditions. Dormancy is adevice thatoptimize the distribution
of germination in time or space, and it has great ecological importance.
This is possible because seeds of many species show variability in
depth of dormancy, which exhibits sporadic release from dormancy
resulting in iregular germination. The temporaldispersal thus, enhances
the spread and survival of the species. Seasonal environmentalchanges
(light and temperature) ultimately regulate seed dormancy and germi
nation, controlling the seed dispersal. Four basic patterns of germina
tion have been recognized in relation to its temporal distribution.
1. Quasi-simultaneous- when germination of all the seeds Occurs
over a relatively brief period.
2. Continuous- in which seeds germinate over an extended
period of time.
3. Intermittent- irregular germination over long periods of time
(multimodal distribution)
4. Periodic- multimodal germination showing more
periodicity. regular

These patterns result from the dormancy characteristics and

from their interplay with various environmental factors
ternperature. Dormancy is also important for preventing like light and
precocious germination i.e., germination while the seed is onvivipary and
the mother
plant,which is important in certain crop
species like cereals.
Dormancy is most commonly found in species which occur in
areas of seasonally harsh climates,
Vive as a seed when it might perish and enables the organism to sur
winter or a long dry season. Even in as a seedling eg., during a cou
moist tropical forests, dormanoy
Occurs in pioneer, light
Macaranga, Trena, etc. demanding, of gap-colonizing
Seedlings these genera genera such as
are incapable of
surviving under thelow light
canopy. Dormancy is useful inconditions found
enabling the undertoundisturbed
species forest
survive as seeds a

unui agap is formed by natural or

broken when the seeds are man-made causes. Dormancy to
expOsed to the increased ig
33 An Introduction to Tree
Seed Technology
temperature in canopy gaps. In drier tropics, seed coat dornancy is
common, most probably a built-in mechanism to withstand unfavourable
environmental conditions. In nature, the dormancy or after-ripening
nhenomenon lasts till the conditions for germination become favourable
Seeds of tropical hard woods in central India for example, mature dur
ing summer months; in case these seeds did not
ripening phenomenon (post harvest dormancy) the youngpossess the after
would not have survived eg., Tectona grandis. germinants
Classifications
Seed domancy can be classified based on
i)Manner of origination (Harper, 1977)
1. Innate dormancy
2. Induced dormancy
3. Enforced dormancy
ii) Time of origination
1. Primary dormancy
2. Secondary dormancy
iii) Location of dormancy
1. Exogenous dormancy (Seed coat dormancy)
2. Endogenous dormancy (Embryo dormancy)
3. Combined dormancy
iv) Cause of dormancy
1. Physical dormancy
2. Chemicaldormancy
3. Mechanical dormancy
4. Morphological dormancy
5. Physiological dormancy
6. Morpho-physiological dormancy
7. Combinational dormancy
6 and 7 are called Double dormancy
A summary of dormancy typesaccording to the classification
is discuSsed as follows.

Innate dormancy
This type of dormancy is present when the seeds are ready for
dispersal t prevents germination during development and maturation
of the seed on the mother plant and also for sometime after shedding
and harvesting. Seeds with impermeable coats and those requiring
prechilling treatments represent innate dormancy. This is also referred
as primary dormancy.
Seed Dormancy 34

Induced dormancy
Aseed may become dormant if, during germination, it is ex
posed to unfavourable conditions when it is predisposed to germinate
viz.. excessive moisture or desiccation, extremes of climates etc. Thie
is also referred to as secondary dormancy.

Enforced dormancy
When a light requiring seed is buried deep in soil and fails to
geminate due to non-availability of light, this is called as enforced dor
mancy.

Exogenous dormancy
Dormancy caused due to factors relating to seed coat or peri
carp..Analogously, mechanical resistance, physical impermeability,in
hibitors or light sensitivity are associated with the seed coat.

Endogenous dormancy
Dormancy caused due to factors relating to embryo eg., imma
ture development or chemical inhibitors located in the embryo.
Combined dormancy
Where twOor more types occur in the same seeds it is called
combined dormancy. Dormancy may be caused due to factors relating
to seed coat as well as embryo.

Acomprehensive classification
Asimplified classification and terminology is presented here
under which basically follows one used by Nikolaeva (1977) and Baskin
and Baskin(1998)with few modifications.
A. Exogenous dormancy
i) Physical: Impermeability of seed coat or pericarp to water
i) Chemical : Inhibitors in pericarp or seed coat
ii) Mechanical : Mechanical resistance of pericarp on seed coat
to embryo growth.
B. Endogenous dormancy
i)
ii Morphological Under developed embryo
Physiological
a)Shallow Physiological inhibiting mechanism
Weak inhibition
b) Intermediate Intermediate inhibition
 35 An Introduction to Tree Seed
Technology
c) Deep Strong inhibition.
Thermodormacy
Temperature sensitive embryo
Light sensitive embryo
Photodormancy
c. Combined morpho-physiological dormancy
Combination of under-developed embryo with strong physiologi
cal inhibiting mechanism of epicotyl growth

D. Combined exogenous / endogenous dormancy (A+B)

A.Exogenous dormancy
1. Physical dormancy
This is caused due to seed coats impermeable to water. The
impermeability is due to a layer of thick walled, cutinised paliside like
macroscleroid cells. If the coat of such seeds is ruptured or softened,
so as to permit the ingress of water, the state of dormancy may be over
come eg., Acacia nilotica and Albizzia lebbeck.
2. Chemical dormancy
Seed coats may contain germination inhibitors. These seeds
lack physiological dormancy and thus the inhibitor chemical can be
leached out of the seed or some how deactivated. Chemical dormancy
is broken by removal of pericarp or leaching of the fruits. Light sensi
tive inhibitors are present in intact seeds of Betula pubescens and
Betula pendula.
3. Mechanical dormancy
It is due to the presence of ahard woody fruit wall usually en
docarp sometime merocarp. Itisproposed that the endocarp acts as
mechanical obstance to a germinating embryo.
In general, once the seed absorbs water and if the embryo is
not dormant, the force of expansion of the germinating embryo rup
ures the seed coat and breaks apart the coverings. But, the coats or
many seeds and indehiscent fruit eg., nuts, are hard and have very
tough tissues; although they allowentry of water, yet offer resistance to
ne growth of germinating embryo. This is because the internal pres
Sure developed is insufficient to burst the seed, a process that must be
effected before swelling can be completed and germination com
menced. eg., Eucalyptus delegatensis and E. pauciflora.
 Seed Dormancy 36

Long periods of coldstratification are required for loss of dor.

mancy. Once the dormancy of embryo is broken however, it has the
arowth potential to push through the endocarp. Thus it appears that
mechanical dormancy inthese species should be reviewed as an as
pect or manifestation of physiological dormancy.
B. Endogenous (embryo) dormancy
1. Morphological dormancy
In some species, at the timeof dispersal the embryo is not fully
grownto get differentiated into radical and cotyledons, there fore ger.
mination does not take place until both differentiation and qrowth oc
cur. Fraxinus spp, requires about 4months to complete development:
In case of Ginkgo biloba even fertilization of the ovule is said to take
place after the seed falls. Requirements for growth are moist subsrate,
suitabie tenmperature and specific dark: light condition. Gibbrelic acid
also increased germination of certain species at super optimum em
perature.

2. Epicotyl dormancy
In seeds of some plants, only a part of the embryonic axis may
be dormant. In Viburnum spp, and Asarum canedense, the radicle
growth proceeds normally while the embryo and embryonic axis re
mains dormant at room temperature. The hypocotyl or root of such
seeds elongate and develop until the stored food is consumed, after
which it withers and dies. This is called epicotyl dormancy, which may
be overcome by chillingor other dormancy breaking
sure of such treatments. Expo
germinated seeds to low temperatures (33 - 50° F), may
induce the tops to resume growth.
3. Physiological dormancy
Non Deep Physiological dormancy
Freshly matured seeds with non deep physiological dormancy
either cannot germinate at any temprature (or) they
Over avery narrow range of temperatures germinate only
Fagus, Pinus and Eucalyptus. Such dormacny (Thermo dormancy) eg.
broken by relatively short periods of cold in some species s
stratification ranging from
days in Triticum sp to 60-90 days in /mpatiens
at room tenperatures come out of biflora. Seeds stored dry
dormancy after ripen. However, the
time requiredfor seeds to after ripen is usually much longer than
required for dormancy loss during cold uia
may ocCur at high stratification. Dormancy loss
temperatures regardless of whether seeds ale
 37 An
imbibed. Non
Introduction to Tree Seed Technology
deepphysiological
including potassium nitrate, thiourea,dormancy can be broken by
A
light requirement for
kinetines, ethylene andchemcials,
gibbrellin.
non deep physicological germination is another
and some dormacy (Photo
Eucalyptus. Seeds of some speciesdormancy) manifestation
eg., Spathodea
of
for germination as they come out of lose their
fcation or high summer dormancy in response torequirement
cold strati-
requirement temperatures, but some species have
for germination after they have a light
ing treatments. recieved dormancy break
Physiological dormancy is caused by embryo and by interaction be
tween the embryo and its covering strutures
Seed coat may restrict movement of
inhibitors (eg., phenolics in seed coat oxygen
may fix
to the embryo or
dation, thus making it unavailable to embryo). Oxygen by oxi
Seed coat may prevent leaching of inhibitors, retard the
of the oxygen which can inactivate or prevent entrance
production of inhibi
torsor themselves contain growth inhibitors. After breaking of dor
mancy,embryo becomes insensitive to inhibitors, even though it is
present.
The embryo covers i.e. the seed coat may mechanically restrict
embryo growth. If seeds are placed under appropriate dormancy
breaking and /or germination conditions, the growth potential of
the embryo increases andgermination occurs. Depending on the
species,cold stratification, gibbrellin incubation temperatures, light
dark may increase the growih potential of the embryo.
IV Endosperm rather than seed coats are the mainforce restricting
embryogrowth and thus germination in some species. Resistance
has been shown tovary depending on the temperature and dark :
ight conditions under which seeds were imbibed and was also
influenced bygibbrellin.
in many species interaction between embryo andcovering struc
ures can be the best explanation for cause of dormancy. Embryo
may produce signal to stimulate production of hydrolases in
edasperm. Embryo may remove or absorb products of enzyme
nydrolysis that inhibit enzyme activity. Embryo may also regulate
inhibitors in the endosperm.
 Seed Dormancy 38

Intermediate Physiological Dormancy

Embryos isolated from the seeds with intermediate PD will grow

the resulting seedlings are normal. The dornancy of intact seeds
or dispersal units is broken by cold stratication but upto 6 months of
this treatment may be rquired.

Deep Physiological Dormancy

Embryos isolated from seeds with deep PD either do not grow
orthey produce abnormal seedlings. The only treatment that overcomes
the dormancy of intact seeds (or dispersal units) is a relatively long
period of cold stratification.

After-ripening
In this type, no morphological changes can be seen from the
time the seed falls from the tree, until it is fully developed and is ready
for normal germination. However, the physiological process that inter
venes is called "after-ripening". This proceeds most actively at alow
temperature (6-10°C).
Chemical changes accompanying after- ripening are listed as follows:
Softening of the seed coat
2. Increase in amount of water absorbed
Increase intotal acidity. Usually,the hypocotyl is at first alkaline but
becomes acidic when the after-ripening is complete, the H-ion
concentration andenzyme activity also increase.
3. Treating with dilute acid (HCL, acetic acid, butyric acid) may shorten
the period required for after - ripening. The increase in acidity is
often correlated with greater water absorbing power of colloids
4 increase of
reducing sugars, amino nitrogen and amino acids
5. Increased respiration
6. Appearance of HCN in cotyledons
7. Increase in catalase, oxidase and
peroxidase activity
8. Short chain fatty acids such as volatile fatty acids may induce seed
dormancy and their loss during after ripening may trigger dormancy
termination.
LFreshly coliected seeds of Tectona grandis would not germi
nate, no matter howelaborate weathering the seed has been
subjected
39 An Introduction to Tree Seed Technology

to: Other examples are Cassia fistula (12 months), Samania saman
(13months) and Lagerstroemia flosreginae (7 months). Dalbergia
latifolia, Pterocarpus marsupium,Adina cordifolia, Albizzia lebbek,
Cassia siamea, Terminalia chebula, Tectona grandis, Zizyphus
jujuba, etc., also require some period of after ripening.
C. Double dormancy (or)combined dormancy
In some species,seed dormancy is caused by the combined
effect of factors associated with the embryo and seed coat. Seed of
Fraxinusexcelsior exhibited deep dormancy, which was caused by
combined effect of restriction of oxygen supply to ermbryo and a re
quirement for chiling. Therefore,more than one treatment may be nec
essary in order to break the double dormancy of such seeds.
Treatments to overcome dormancy
Pretreatment for dormant seeds is a must to enchance rapid
and uniform germination of seed sown in the nursery, field or during
seed testing.
i)Mechanical scarification
Large quantities of hard coated seeds can be mechanically
Scarified using concrete mixer with sharp gravel or sand; special drum
lined with abrasive material such as sand paper, cement or crushed
glass or fitted with abrasive disks may be made. Care should be taken
to avoid over treatment eg., Acacia catechu, A. nilotica, Albizzia
falcataria, A. lebbek, Cassiafistula, C. javanica, C. nodosa, Delonix
regia, Santalum album, Terminalia arjuna, T. tomentosa.
i) Water soaking
This treatment is intended both to soften hard seeds and to
leach out chemical inhibitors. Soaking in cold water for a period rang
ing from a few hours to several days, is asafe and simple treatment,
which can be tried on alI, but the smallest seed. Leguminous seeds
often respond wellto immersion in boiling water; the seed to be treated
being placed in a quantity of boiling water more or less equal to its own
Volune and left tili the water cools. eg., seeds of oaks and chestnuts
are reported to respond well to a 1/2 minute immersion in boiling wa
ter. Soaking in water for periods ranging from 2 to 48 hours according
to species viz., Acacia mearnsii, A. nilotica, Adenanthera
microsperma, Albizzia amara, Grevillea robusta, etc.Prescriptions
for hot water treatment must be applied meticulously to remove seed
Coat dormancy without kiling the seeds through successive heating.
Seed Dormancy 40

ii) Acid scarification

Soaking in concentrated sulphuric acid is the most common
method of treatment to soften the seed coat. For this, the materials
needed include commercial grade sulphuric acid (95%,36N), acid re
sistant containers, wire containers, Screens and an abundant supply of
water for rinsing the seeds after treatment. To scarify the seeds. pour
the undiluted acid until all the seeds are wetted. Stir them for few min
utes to one hour depending on the species and immediately wash them
thoroughly in cool runningwater for 5-10 minutes to remove all traces
of acid. Shade dry he seeds unless wef sowing is preferred. In seed
coat of Prosopis juliflora, an oily deposit in the coat is responsible for
delayed germination,seeds germinated when this is dissolved in ether.
iv) Dry heat treatment
Spread the light inflammabBe litter over the fruits and then ig
nite it. However, adjusting the heat of the fire to achieve the maximum
effect on the pericarp without damaging the seed embryo requires ex
perience. The exposure of seeds of Acacia mangium to dry heat at
100°C was nearly as effective as immersion in water at 100°C in over
coming dormancy. In Tectona grandis, the drupes exposed to 45°C -
50°C showed 50 per cent germination.
v) Warm stratification
This has been recommended both to overcome mechanical
and morphoiogical dormancy. Soak the seeds in several times their
volume of coid water at approximately 3-5°C for 48 h. Drain off the
water and mix the seeds with two to four times their volume of a moist
ened, water retained medium such as sand, sandpeat mixture or ver
miculite. Store at a warmn temperature. A constant temperature of 20
25°Cor alternat ingtemperature of,20°C and 30°C is suitable for many
species. Open the containers weekly, mix seeds and if surfaces show
signs of drying out, remoist with water spray.
ir
vi) Cold stratification
This is recommended to remove physiological dormancy. Seeds
shoutd be soaked in severat times their volume of water before
prechilling, a 48 h soak at 3-5°C. After soaking, the water is drained o ro
and moist seeds are stored in 3-5°Cfor the period appropriate to eaci
species. Storage may be without any mediumi.e. seeds as such or the
seeds may be mixed with 2-4 times its volume of a medium such as ha
 41 AnIntroduction to Tree Seed
Technology
moist sand, peat or a mjxture of the two. Containers shal! be opened
and mixed if surfaces dry off eg., Abeis spp,Eucalyptus
etc. delegatensis,
The combination of high moisture and low
to trigger off changes in cellenergy relationships temperature appears
leading to biochemi
cal activity which transforms complex food substances into simpler forms
utilized by the embryowhen it renews growth at germination. Low tern
erature not only favours these biochemical changes, but also reduces
microorganisms' activity and the risk of overheating and prermature
germination of the after-ripened seeds.
vii) Biological methods
innature, animals and microorganisms are an important factor
in the break down of seedcoat impermeability. It is difficult tomake use
of these organisms as a controlled pretreatment of seed, but in few
cases successfu! results have been obtained. There is no doubi that
certain seeds germinate more freely after they have passed through
the bodies of animais. The combination of moisture, warmth and chemi
cal action of the digestive juices doubtlessly soltens and renders the
hard seed coat permeable. This method has been experimented with
Santalum album, Acacia nilotica and Macaranga denticuiata, but
effective control is very difficult.
Hormonal regulation of Seed Dormancy
Abscisic acid inhibits germination of non dormant seeds. It is
the main chemical factorfor induction of dormancy frequently accumu
lating in maturing seeds.
Gibberelin stimuiates seed germination of many species. Dor
mancies with chilling and light requirements are often overcome by
treating with GA.
Cytokinin is more effective than GA in overcoming the effect of
inhibitors like ABA of various GA sensitive processes.
pa

Auxin (|AA)may be associatedwith the later stages of embryo

expansion during seed germination,but may not play any significant
role in development or termination of seed dormancy in forest seeds.
Ethylene breaks the dormancy of seeds of many species and
has been implicated in the regulation of seed dormancy.
 Seed Dormancy 42

Regulationof dormancy by relative contents of promoters and

inhibitors, is depicted as under Fig. 10.
Fig10. The essential role of gibberelins and the enabling role of
cytokinin inseed germination and dormancy. (From Ref 55).
Without Gibberellin With Gibberellin

+Cytokinin Cytokinin
- Inhibitor No GerminationGermination
germination Inhibitor No Germination
germination

Chemical regulation of seed dormancy

Potassium nitrate
it can completely counteract light inhibition of seed germina
tion in certain trees seeds.

Hydrogen peroxide
Hydrogen peroxidereleases the seed coat restraint by causing
cracks in hilum along the ridge of the seed coat resulting in increased
uptake of water. This chemical is known to act as respiration stimulant,
which accelerates the breakdown of food substances, thus providing
rapid supply of energy and materiats for synthesis in growing points.
Thiourea
This is abie to substitute light and temperature requirements of
germination.
 CHAPTER-12
SEED COLLECTION
The practice of colecting tree
tion of forests and seeds is very old. The exploita
seeding and plantingrecognition
of the need for their
gave rise to the activity of seed replacerment by
1own. Success of any planting collection as now
The quality of seeds in tun programme depends on quality seeds.
niques and handling. It costs depends
on the source, collection tech
just as
seed as it does from seed of highestmuch to establish tree frorn poor
ence in material and returns could be genetic potential, but the differ
quality seeds improved the survival bygreat. The benefit of usingbetter
two folds and gave greater eco
nomic returns.

Seeds or fruits should usually be collected just as they

reached full ripeness and before natural dispersal begins. The timinghave
of seed collection is important, failingwhich it may leadto a
totalloss of
seed of the season i.e., until the next seasonyear no seeds can
be
collected. Seeds may be lost due to dispersal eg., Hardwickia binnata,
Bombax malabaricum, etc.,or due to feeding by birds which find fruits
of Simarouba glauca, Azadirachta indica and Santalum album
highly attractive. In most broad leaved species, seed collection does
not pose much problem. in leguminous species eg., Pongamia pinnata,
Cassia fistula, Dalbergia sissoo, Albizzia lebbek, etc.. pods remain
hanging on the trees for a periodof time facilitating collection. How
ever, collection at seed maturity stage is envisaged to avoid pest and
disease attack, which will eventually reduce the quantity and quality of
the seed.

When large quantities of seed are collected, some planning

and organization are necessary. Factors to be considered are;
i) An estimate has to be done depending on the quantity of seeds
required.
i) Surveys have to be conducted in each seed zone for selection of
seed source, to make note of the flowering time at diferent loca
tions so that the exact period of seed ripening can be determined.
ii) The seed collector must select a number of plus trees or seed stands
from which seeds can be procured
IV) In species that exhibit periodicity in seeding, speculative collec
tions should be made for anticipated requirements for the future.
 82 An Introduction to Tree Seed Technology

In tree species,which is mostly indeterminate, harvest at any

given time yields seeds with varying stages of maturity and growth
potential. Consequently, the date of harvest is acompromise to give
the greatest amount of germinable, high vigour seeds from a popula
tion changing from immature seeds to those which are ready to shat
ter. Seed should generally be collected when it is fully ripe or mature.
Fully mature seeds have an advantage ofcomplete physical and physi
ological development, which is needed for maximum expression of
vigour. Mature seeds have higher germinative energy and greater lon
gevity in storage than immature seeds because of good physical and
physiological development, which, helps to retard the deterioration pro
cess. Premature seed succumbs to pest and insect attack very rap
idly. Complete seed maturity helps to produce maximum seedling vigour
even under adverse germination conditions.

Analternative method isto collectfruits before they ripen and

to store themin relatively cool, weli-ventilated conditions, whichper
mits further ripening of the seeds within the fruit. There are certain
advantages in collecting immature seeds i) to extend the colection
Season, i) to avoid damage to the seed crop by insects and other pests
eg.. Aibizzia Tebbek, A. procera and Shorea robusta and üi) to
salvage immature seed collectedinadvertentiy. Most of the changes in
ripening fruits and seeds have been viewed by various researchers as
probable indicators of maturity.
indices of maturity
i) Physical indices
The physical indices such as a) colour differences, b) increased
firmness or brittleness, c)decreased moisture content and d) changes
in physical dimensions are widely used, but many are highly subjed
tive estimates and their success largely depends on the
the collector (Table 9). experience o
Seed Collection 83

Table 9.Colour variation in maturing pods of different species

Species Phases of development
Maturity
Acacia auriculiformis Green Brown Red Black

AIbizzia procera Green Red Brown Brown

Azadiractha indica Green Greenish| Yellowish|Yellow
yellow green
Cassia glauca Green Brown Brown Black

Cassia siamea Green Brown Brown Brown

Casuarina equisetifolia Green Greenish Brownish Brown

brown green
Ceiba pentandra Green Yellow Brown Black

Prosopis juliflora Green Greenish Yellow |Yellow

yellow
Santalum album Green Greenish Reddish
red brown Black

i) Biochemical indices
This inciudes a) crude fat, b) iodine absorption number of fat,
c)reducing sugars, d) non-reducing sugars, e) starch, f) soluble nitro
gen and g)protein nitrogen.
Pseudotsuga menziesii seeds with reducing sugar level of
22 mglg were immature, while full maturity was achieved when the
level had fallen to 13 mg/g. n seed of Abies procera, the crude fat
level of 22 mglg is a good indicator that cones are ripe. However, such
indicators remain relatively impractical for small seed collection opera
tions.

ij) Laboratory methods

1.Dry weight
The most generally accepted measure is the time when the
seed has reached its maximum dry weight. This means the nutrients
are no longer flowing into the seed from the mother tree.
 84 AnIntroduction to Tree Seed Technology
2.Chemical analysis
Content of crude fat and protein nitrogen, which increase five
and four times respectively from immaturity to physiological mäturty,
are the best indices for Fraxinus pennsylvanica. But they have no
advantage over an examination of the embryo and the colour change
of the fruit. The extra trouble to perform the analysis does not appear
to be justified.
3. X- radiography
The examination of the development of the embryo and en
dosperm of sample seeds by means of x-ray radiographs is a quick
and relatively straightforward method for assessing seed maturity. The
technique has been successfully followed in Tectona grandis and a
number of tropical trees.
4. Moisture content of fruits
Water loss in the fruits occurs during development and is closely
related to maturity of seed.

iv) Field methods

1. Specificgravity
As moisture content decreases, the specific gravity decreases.
Seeds when droppedin the liquid of knownspecific gravity, they sink
or fioat based on its own specific gravity ultimateiy the matured seeds
sinks and gets separated from the floaters which are immature. The
specific gravity of kerosene, light SAE 20 motor oil and linseed oil is
0.80, 0.88 and 0.93, respectively. Flotation liquids can be prepared
arious mixtures of these liquids having adesignated specific
gravity.Specific gravity indices have proven reliable for some temper
ate conifers e.g., a specific gravity of 0.74 for Picea gluaca however,
this is not suitable for several hard woods.

2. Examination of seed contents

Examination of seed contents by cutting open fruits can be a
reliable and simple method of assessing seed ripeness. Mature seed
has afirm white endosperm and a fully developed firm embryo.
3. Colour of fruits
Ghanges in the fruit colour provide a simple and reliable crite
rion for judging seed maturity, but the operator must be experience0
the characteristics of the species concerned. Colour changes are usu
 Seed Collection
85
ally from the green to various
'e
shades of
Since the seed normally matures before yellow, brown grey,grey, bla
black etc.
the fruit, it is
species to plan the collection an earlier at
at
stage advisable
of
in some
Az2dirachta indica, among the maturity stages, colour change.
the
drupes registered yellowish green
and green drupes. In maximum viability and vigour compared to yellow
ar
Casuarina equisetífolia, seeds from brown Cones
were superior to greenish brown
bum, when compared with green,and green cones. In Santalum al
black shriveled fruits, seeds collectedgreenish red, reddish brown and
from black fruits recorded higher
germination and seedling vigour besides maintainíng better viability
andvigour in storage (Table 10).
Table 10.Influence of fruit colour on
of sandal
germination and vigour
Fruit colour Germination Root length Dry
k%) Vigour
(cm) matter index
production
(mgl
seedling1)
Green 0.0 0.0 0.0 0.0
Greenish red 29.0 5.9 14 390
Reddish black 68.2 6.9 19 1152
Black 73.2 7.1 21 1267
CD (P=0.05) 0.72 0.10 0.5 326

Athorough knowledge about the optimum stage of maturity for

collecting the seeds of each tree species is important to ensure collec
tion of maximum quality seeds. The first seeds or fruits, which fall natu
rally, are often of poor quality in which case it is advisable to reject
them and to postpone collection until the peak and the latter half of the
season. InTh¡iland, drupes of Tectonagrandis start to shed in March
but observations have shown that most viable drupes are shed only
later, so collections starting only during April is recommended.
Types of seed collection
) Provenance seed collection
Following are the guidelines with regard to provenance seed
collection.
a) Collect from normal trees rather than plus trees. Collections if made
from superior phenotypes, it should be kept separately.
 AnIntroduction to Tree Seed
86 Technology
h) Collect from a minimum of 10trees preferably 25 to 50 in a stand
If the stand is very variatble, the number can be increased The
number of trees and the approximate
recorded.
percentage that they consti.
tute in a stand should be
c) To reduce the risk of collecting from half sib parents, seed trees
should be at least separated by seed fall distance eg., 100 mfor
Pseudotsugaand 50 m for Tectona grandis
d) Individual seed trees should be marked.
e) Collect equal number of cones, fruits or seeds per tree.
f) In first stage provenance collections, seeds from individual trees
may be mixed together. For special studies on individual geno
types,seeds from each tree should be kept separately.
i) Single tree collections
This is done in the following conditions; a) Tostudy the prov
enance differences under local conditions incases where exotics are
to be introduced into countries, b)To investigate variation between in
dividuals in the best provenance by means of progeny trials in order to
identify the best adapted provenance and c) If progeny trials are the
object of seed collection, it is essential to keep seed from individual
trees separately at all stages of seed handling.
i) Single clone collection
During seed collections in clonal seed orchards, it is important
to keep the clone ramet separately so as to maintain their identity. This
helps to make composite seedlots with equal clonal representation,
avoiding the dominating effects that some clones frequently have in a
bulked seed lot.

iv) Large scale collections

Here, emphasis is on collecting as much seed as possIDie
quickly and cheaply rather than on very careful selection of the parent
trees. The guide linesto be followed are a) collect from healthy vigor-
ous trees, b) avoid young and over mature trees, c) avoidisolated
to be
trees of naturally cross pollinating species, since these are likely
self pollinated and d) Avoid stands containing numerous poorlyfoformed
abnormal or diseased trees.
Methods of collection
i) Collection from
ground
Seeds/ fruits, which have fallen after natural ripening
can be
Seed Collection 87

collected from the ground. This is recommended for species which

have large heavy seeds, or bulky fruits which fall unbroken below the
parent trees eg., Dipterocarpus spp., Shorea robusta, Azadirachta
indica, Syzygium cumini, Tectona grandis, etc.
Advantages
1. Lesser cost is involved.
2. Does not require much of the skilled labour.

Disadvantages
1. Insects, fungi, birds and animals may attack seeds not collected
îmmediately after shedding from trees.
2. Seeds having short viabilityloose germinability quickly if collection
is delayed eg.,Syzygium cumini.
3. If single tree collection is intended, ground collected seeds pose
the danger of being shed from the tree near by tree.
4. Seeds may be collectedfrom over ripe fruits.
5. Collection of empty, damaged. illfilled,immature and unsound seed.
6. Cleaning process is difficult due to admixtures.
Whenever seeds are gathered from the ground, great care
should be taken tosee that only freshly fallen seeds are taken, and to
ensure this, it is advisable to clear and sweep the ground below the
selected seed trees, and daity coliections shall be made thereafter. A
careful watch must be kept to see that the seeds gathered are of
suitable quality, free from insect damage and fully ripe.

2. Collection directly from felled or standing trees

This is recommended in the following situations.
i) Fruits too small to be economically picked from the ground after
felling eg., Terminalia myriocarpa.
ii) Fruits or seeds that are widely dispersed by wind. eg., Ailanthus
excelsa, Casuarina equisetifolia.
ii) Fruits which suffer damage by insects or animals between the time
ca, Terminalia
of seed ripening and fruit fall eg., Michelia champaca,
tomentosa.

Thecheapest and easiest method is to gather seed from freshly

Telled trees eg., Casuarinaequisetifolia, Eucalyptus spp. If it is not
possible to fellthe entire tree, climbers may be sent up to pluck the fruit
eg.,Tamarindus indica and Ceiba pentandra. Good bearing branches
 88 An introduction to Tree Seed Technology

can also be cut downtocollect fruits eg., Albizzia lebbek. Manual or

mechanical shaking of trees can also be done to collect seeds from
Tamarindus indica, Emblica officinalis etc. Long handled cutters are
useful to collect pods from Acacia and Cassia species.
Advantages
1) Single tree collections are successfully maintained.
2) Wastage of seeds by insects and diseases can be avoided.
3) Large quantity of seeds can be obtained at the right stage of
maturity.

Disadvantages
1. While collecting from feled branches or treesunripe fruits may be
collected along with ripe fruits.
2. High risk is involved while climbing up the clean bole of superior
trees.
3. Cost involved is higher.
3. Bird droppings
Fruits highiy attractive eg., Azadirachta indica, Syzygium
cumini, Ficus religiosa, F. benghalensis, F. carica, Santulum
album, etc., are eaten by birds and after digestion and seeds are ex
pelled out. Seed viability and vigour are however, not affected due to
their passage through the digestive tract. However, in seeds that have
hard seededness or physical dormancy, the digestive enzymes may
help in scarifying the seed coat, thereby improving the germinability.
Harvesting appliances
A) Tools for seedcollection from felled or standing trees with ac
cess from ground
i. Secauteur
Operated by hand, secauteur makes it quicker to cut thewhole
bunches of fruits without permittingindividualfruits to fallon the ground.
it can easily be kept in the pocket since it is small and light.
ii. Sickle with ahandle
It can be used for cutting the twigs bearing the fruits but lt
not check the fruits that fall off on the ground, unless its edge is sharp:
This tool is often operated by hand and makes the cutting easier as
comparedto that done by fingers.
 Seed Collection
89
B) Tools for seed collectionfrom standing trees
) Ropes and spars
If trees are very high
tree with the help of
(25-30 m) it is Compulsory to clinb on the
ropes and spurs. After
ter or sickle is used to detach the fruits. climbing, long handled cut-
i) Long handied cutter
Aluminium pipe or 2 m pieces of straight bamboos are joined
together with metallic caps having compatible inner and outer
Thus the length of the handle may be adjusted threads.
but upto a maximum of 8-10 m. according the need,
to
The cutter has a knife of 15 to 30 crm
length,with sharp edges on both the sides. Aluminium pipes are better
because they are light in weight.
Seed bearing years
Seed bearing of many forest trees is rather irregular fromyear
to year. One year with a heavy crop, maybe followed by one or severa!
years with a poor seed crop or none at al. The year with heavy seed
crop is termed as seed year or mast year.

Two kinds of periodicity are recognized i) inner or spontaneous

periodicity; it is more mysterious and has long been intrigued. One
hypothesis states that seed bearing depended chiefly upon the accu
mulation of reserves. Reduction in nitrogen, magnesium and calcium
have been reported after a seed year, that insufficient food then re
mained for reproduction and i) induced periodicity; it depends upon
the periodic occurrence of the external condition favourable to fruiting.
Water shortage iseffective in retarding vegetative growth and drought
have occasionally been correlated with seed years. Insect defoliation
reduces the capacity of the tree to store reserves for the coming winter,
and may reduce the size and vigour of seed being formed at the time.
Of courseé the impetus to seed formation came from the previous years.
Tectona grandis has generally good flowering each year, although
exceptionally good seed years are observed in some localities on a
3-4 year cycle.
Advantages
1. The cost of collection will be lower,
2. Ihe seeds will usually be of higher germination capacity and will
retain their viability longer than those collected in a poor seed year.
 90 An Introduction to Tree Seed Technology

3. Aheavy seed crop usually reflects a previous heavy production

of pollen, to which all or most trees in the stand have contributed.
Collection in a good seed year therefore, conserves a higher pro
portion of the genetic diversity