Seed structure

A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some
stored food. The formation of the seed completes the process of reproduction in seed plants.

A seed develops from an ovule after fertilization. It consists of a tough coat or testa enclosing
an embryo which is made up of a plumule, a radicle and one or two cotyledons. In favourable
conditions the seed can grow and become a fully independent plant, bearing flowers and seeds
during its life cycle. In the embryo of the seed are all the potentialities of development and
growth to a mature plant resembling other members of its species in almost every detail of leaf
shape, cell distribution and flower colour and structure.

• External (outer part)

– Seed coat (testa)
– Hilum
• Embryo (inner part)
– Cotyledon
– Epicotyl / Hypocotyl
– Pumule
– Radical

The embryo is what forms the new plant once the opportune conditions are present.

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The main components of embryo are are:
 The cotyledons:: Endosperm, a temporary food supp supply,
ly, is packed around the embryo in
the form of special leaves called cotyledons or seed leaves. These generally are the first
parts visible when the seed germinates.
The cotyledon leaves, attached to the embryonic axis. There may be one ((Monocotyledons
Monocotyledons), or
two (Dicotyledons). ). The cotyledons are also the source of nutrients in the non-endospermic
non
dicotyledons, in which case they replace the endosperm, and are thick and leathery. In
endospermic seeds the cotyledons are thin and papery.
 The epicotyls: the embryonic
mbryonic axis above the point of attachment of the cotyledon(s).
 The plumule: the tip of the epicotyl, and has a feathery appearance due to the presence
of young leaf primordia at the apex, and will become the shoot upon germination.
 The hypocotyls: the embryonic
bryonic axis below the point of attachment of the cotyledon(s),
connecting the epicotyle and the radicle, being the stemroot transition zone.
 The radicle: the basal tip of the hypocotyl, grows into the primary root.

Seed coat:

The seed coat forms from the he two integuments or outer layers of cells of the ovule, which
derive from tissue from the mother plant, the inner integument forms the tegmen and the
outer forms the testa. Seed coats can be thin and soft as in beans or thick and hard as in locust
or coconut seeds.

Hilum: Scar from the seed being

ng attached to the parent plant.

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Cotyledon:

• The cotyledon is the first leaf that germinates.

• It is filled with stored food that the plant uses before it begins photosynthesis.
• Some plants have 1 cotyledon (mon (monocot)
ocot) and some have 2 cotyledons (dicot).

Dicot vs. monocot

• Dicot has two cotyledons (like bean) food is kept in the cotyledons.
• Monocot has one cotyledon which absorbs the endosperm tissue during germination
(corn).
Epicotyl vs. hypocotyls: The basis ffor
or the plant’s stem. It is known as the epicotyl above the
cotyledon and a hypocotyl below the cotyledon. These grow upward in response to light.

Plumule: The shoot tip with a pair of miniature leaves.

Radicle: The part of the seed where the root dev
develops.

Some other structures:

Perisperm : Diploid maternal food storage tissue originates from the nucellus. Reported only in
some species. e.g. Beta vulgaris, Piper nigrum, Coffea arabica, many Caryophyllales.
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Endospermic seeds: The endosperm is present in the mature seed and serves as food storage
organ. Testa and endosperm are the two covering layers of the embryo. The amount of
endosperm in mature seeds is highly species dependent and varies from an abundant
endosperm layer (Nicotiana
Nicotiana tabaccum) to a si
single layer (Arabidopsis thaliana).
Non-endospermic seeds: The cotyledons serve as sole food storage organs as in the case of pea
(Pisum sativum). During embryo development the cotyledons absorb the reserves from the
endosperm. The endosperm is almost degradegraded
ded in the mature seed and the embryo is enclosed
by the testa. Examples:: rapeseed ((Brassica napus), and the legume family including pea (Pisum
(
sativum), garden or French bean (Phaseolus vulgaris), soybean (Glycine max).
Special structural features:
Hilum and funiculus: Funicular scar on seed coat that marks the point at which the seed
was attached via the funiculus to the ovary tissue.
Micropyle: The Micropyle is a canal or hole in the coverings (seed coat) of the nucellus
through which the pollen tube us usually
ually passes during fertilization. Later, when the
matures and starts to germinate, the micropyle serves as a minute pore through which
water enters.
Chalaza: Non-micropylar
micropylar end of the seed. The base of an ovule, bearing an embryo sac
surrounded by integuments.
ments.
Raphe: Ridge on seed coat formed from adnate funiculus.
Operculum: A little seed lid. It refers to a dehiscent cap of a seed or a fruit that opens
during germination.
Carunculate: Seed with an excrescent outgrowth from integuments near the hilum, as in
Euphorbia.
Pea seeds: The embryo of mature seeds of Pisum sativum consists of the embryonic axis and
the cotyledons. The fleshy storage cotyledons make up most of the seed's volume and weight.
The pea embryo is enclosed by the testa and the endosperm is obliterated during seed
development, when it's nutrients are taken up by the embryo.

Tomato and pepper seeds are type members of the Solanoideae subgroup of Solanaceae:
Solanaceae
In mature tomato (Lycopersicon
Lycopersicon esculentum
esculentum) and pepper (Capsicum annuum)) seeds the embryo
is surrounded by an abundance of endosperm cells and by the testa (seed coat). The embryos

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are curved and flattened, the seeds are discoid, and a micropylar cap
cap-like
like structure consisting of
endosperm and testa covers the radicle tip. This micropyla
micropylarr cap of Solanoideae-type
Solanoideae seeds is
the place of radicle protrusion, but there is no visible distinction beween testa rupture and
endosperm rupture.

Embryo Development and Deposition of Reserves during Seed Development

Development stage of seed

During seed development seed is composed of several tissue

tissues including Embryo,
Embryo Endosperm &
Testa. Embryo comprises
ises the axis which contains the root and shoots meristem that ultimately
grows to form the vegetative plant. Endosperm is a nutritive storage tissue it i may be
reabsorbed during seed development as a food sources. Testa is derived from integument.
integument

Embryo Development

Begin once the egg cell is fertilized

fertilized. The growing pollen tube enters angiosperm embryo sac and
releases two sperm cells, one
ne sperm fert
fertilizes central cell and initiates endosperm development
and the other
ther sperm fertilizes the egg to produce a zygote
zygote. Cell division soon follows, creating
the embryo

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Phases of embryogenesis
Embryogenesis occur in three overlapping phases i.e. Differentiation of tissue,, cell
c enlargement
& Maturation. During histo-differentiation
differentiation the fertilized egg undergoes multiple divisions
division and
differentiates to form the embryonic tissue and organ. A suspensor forms and facilitates
embryo o division until mid maturation (torpedo stage). Cell enlargement accommodates
accommodate the
deposition of stored reserves d during the torpedo stage. During maturation drying
developmental processes are terminated as the embryo prepares for and is subjected to
desiccation.

Octant stage: Four of eight

ight cells (darkly stained) in two tiers are visible. Cells of the upper and
lower tier of the octant will give rise to specific parts of the seedling
seedling.

Dermatogen stage: A tangential division of each of the eight ‘octant’ cells produces inner cells
and epidermis
pidermis (protoderm) cells
cells.

Early globular stage: The

he divisions of the inner cells immediately after the dermatogen stage
are oriented in the apical-basal
basal dimension, endowing the embryo with a morphologically
recognizable axis.

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Procambial stage: Differences in the orientation of cell divisions in the center and periphery of
the newly generated narrow procambial cells. At this stage cells remain isodiametric.

Triangular stage: Here a polarized pattern of major elements is recognizable cells have
generated two symmetrically positioned cotyledon primordia and cells a radially patterned
cylinder (comprising epidermis, ground tissue and vascular tissue). Additional divisions
distinguish the ‘hypophyseal cell’ from other suspensor cells. Its descendants will ultimately
form the quiescent center of the primary root meristem and the columella initials.

Heart stage: Cotyledon outgrowth. Subsequently, cells not contributing to cotyledon formation
initiate the primary shoot meristem.

Mid-torpedo stage: Enlargement of cotyledons and hypocotyl and further elaboration of the
radial pattern. Vascular differentiation in the cotyledons is visible.

Bent cotyledon stage embryo with elaborated radial pattern in different organs.
• Many genes are involved in the regulation of development.
• Leafy cotyledon(LEC) and FUSCA (FUS) genes control late embryogenesis,
• The heart stage and encode proteins that might function as critical components in the
transduction of environmental and cellular signals during late development.

Embryo development in a dicot

Stages of embryogenesis:

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 • Proembryo
• Globular
• Cotyledon
• Mature embryo
pro-embryo development
• Following fusion of the egg and sperm nuclei, a proembryo is formed by a transverse cell
division to form an apical and basal cell.
• The basal cell divides again and will form the suspensor.
• The apical cell forms the embryo.
• The suspensor in dicots is usually a column of single or multiple cells.
• The suspensor functions to push the pro-embryo into the embryo sac cavity and to
absorb and transmit nutrients to the proembryo.

Globular and cotyledon stage

• Basal cell derivatives in the globular embryo form the hypophysis that goes on to
develop into the radicle.
• Tissue differentiation becomes evident in the 16-celled globular embryo where
peripheral cell division begin to form a protoderm.
• Cell division continues as the embryo enters the cotyledon stage.
• Two recognizable stages are the heart and torpedo stages based on the degree of
elongation seen in the developing cotyledons.
• In the cotyledon stage, the embryo has organized to form an apical meristem, radicle,
cotyledons and hypocotyl.

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Mature embryo stage
In mature embryo stage, the embryo is fully formed and separate cotyledons can be easily seen
as well as a distinct shoot and root meristem. At this time the suspensor and the basal cell
begin to disappear.

Root and shoot formation

Established during globular stage of development formation of each controlled independently
Morphogenesis
Globular stage gives rise to heart-shaped embryo with cotyledons produced by embryonic cells

Endosperm development follows different pathway: Two major types of endosperm

development
• Nuclear endosperm development
• Cellular endosperm development

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Deposition of stored reserves during seed development
There is no direct vascular connection between the parent plant and the embryo or endosperm.
Assimilates are transported from the parental seed coat cells to the extracellular space
(apoplast) by a passive facilated membrane transport process and are subsequently taken up by
the filial tissues of the endosperm or embryo. During early development, sucrose is genereted
by the parent plant released from the pholem into the apoplast and hydrolyzed by invertase in
the cell wall. The hexoses products are loaded into the fillial storage tissue.
Invertase also influences the development of cereals seeds, miniature-1 a maize mutant lackng
invertase activity demonstrated aberrant development of both the endosperm and the area of
phloem unloading.
During the early stages of development sucrose may be resynthesized from the hexose
imported into the storage tissues by the sucrose-phosphate synthatase pathway. The major
forms of nitrogen translocated into the seed in the pholem are asparagine and glutamine.
Although in some legumes about 10 to 15% nitrogen translocated into seeds is used, the
composition of the amino acid reaching the developing storage tissue often differs from that
entering the pholem of the parental tissue, so enzyme that interconvert amino acid must be
present in the seed coat.
The products of seed coat asparaginase and aspartate aminotransferase are imported into the
embryo or endosperm and serve as substrates for synthesis and inter-converson of amino acid.
Later during development when reserve protein synthesized is high asparaginases activity in the
seed coat is very low and asparagine is taken up directly into the cotyledons, where it is
deamineated by a cotyledon localized isoenzyme of asparaginase the activity of which increase
during cotyledon development. Glutamine synthatese and glutamate synthatase which act

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together to assimilate the NH4+released by asperginases also become more active in cotyledons
as development progress. No mutants for the Nitrogen translocattranslocation
ion and biosynthetic
pathways.

Stages of seed development and germination

Dormancy block during seed development and germination

Seed germination

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Germination in plants is the process by which a dormant seed begins to sprout and grow into a
seedling under the right growing conditions.

Types of germination

(i) Epigeal germination

– In this, the cotyledons are raised out of the soil and generally become green and
photosynthetic. In dicots, they are pushed up by rapid extension of hypocotyl
before growth of the epicotyl.

– Epigeal germination occurs in bean, caster, mustard, tamarind, sunflower etc.

(ii) Hypogeal germination

– In this type of germination, the cotyledons remain underground. Hypocotyl

growth is restricted. The epicotyl grows to raise the first leaves out of the soil.

– Hypogeal germination occurs in dicotyledenous seeds of gram, pea, mango,

ground nut etc and in monocotyledons like rice, maize, wheat etc.

Special type of germination

(iii) Viviparous germination

– This is a special type of germination occurring in mangrove plants. These plants

generally grow in salty lakes, sea coasts and deltas. Here, the seed germinates while still
attached to the parent plant.

– The embryo emerges out of the fruit with a massive radicle pointing downwards. Due to
increased weight, the seedling separates from the parent plant and establishes itself in
the muddy soil below.

– Example: Rhizophora

Mechanism of germinating seed

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Process behind germination

STAGE EVENTS

PREGERMINATION (a) Rehydration – imbibition of water.

(b) RNA & protein synthesis stimulated.
(c) Increased metabolism – increased respiration.
(d) Hydrolysis (digestion) of food reserves by enzymes.
(e) Changes in cell ultrastructure.
(f) Induction of cell division & cell growth.

GERMINATION (a) Rupture of seed coat.

(b) Emergence of seedling,
ling, usually radicle first.

POST GERMINATION (a) Controlled growth of root and shoot axis.

(b) Controlled transport of materials from food stores to
growing axis.
(c) Senescence (aging) of food storage tissues.

There are several factors affecting seed germina

germination.
tion. First and foremost, seed must be viable meaning it
must be capable of germination.

1. Abiotic Factors:
1) Light
2) Temperature
3) Aeration (Oxygen)
4) Soil type and depth of sowing
2. Biotic factors:
1) Viability of seed
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 2) Dormancy period
3. Other factors:

Factors that affect seed germination are divided in two categories, internal and external factors. The
internal factors include seed vitality, genotype, seed maturation and seed dormancy. The external
factors are: water, temperature, oxygen, light and smoke.

Light
• Generally seeds require darkness to germinate. However, lettuce, tobacco, tomato and
many grasses need to light exposure to germinate.
• These seeds require the red portion of the light spectrum, while far red light inhibits
germination.
• Many small seeds with low amounts of storage reserves (such as lettuce) show such a
red light requirement.
Phytochrome is a plant pigment found in cytoplasm that senses the presence of red light. The
red-absorbing form of phytochrome changes to the far-red absorbing form when it absorbs red
light (660 nm) and back again when it absorb far-red light (730 nm). Thus the presence of the
far-red form of phytochrome ends the inhibition of germination in these seeds.

Temperature
• As with most reactions germination generally occurs faster when at warmer
temperatures. However there is sometimes a need for cool temperatures to break
dormancy.
• Stratification is one strategy that is employed in woody species in particular. It requires
a moist, cool period that degrades growth inhibitors that prevent germination. Once the
inhibitors are degraded and all other conditions are met then germination will occur.

Oxygen
• Another requirement for germination is aeration (oxygen).
• Respiration rates for germinating seeds are very high, therefore adequate oxygen is
necessary. The germination percent of most seeds will be retarded if the oxygen percent
goes below 20 percent. (Normal air is 20 percent oxygen.)
• B. Seedbeds that are over-watered or poorly drained will cause the oxygen supply to
become limited, so the germination percent will diminish.
Soil type
• Soil type is widely affected on seed germination, there are so many parameters of soil
affecting on seed germination.

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 • In that factors, soil salinity, acidity, salt concentrate, EC of soil, soil porosity, water
holding capacity, texture of soil also affect on seed germination. Most problematic
factor is water holding capacity of soil water holding capacity.
Depth of sowing
• Proper planting depth is a direct correlation to seed size.
• The general rule of thumb is larger seeds can be planted more deeply than small seeds.
This is due to the energy needed to emerge. Larger seeds have greater food reserves
from which to draw energy for respiration and growth. They are able to emerge from
greater depths.
• Soil types also affect the planting depths. The surface of sandy soils tends to dry out
quickly, so seeds planted in these soils should be planted deeper, than in loam soils.
Stages of germination: Phase of activation followed by phase of digestion and translocation.
Process of seed germination: Imbibition, GA formation, Transcription, transportation and
translation of amylase, amylase accelerates starch hydrolysis, movement of hydrated starch to
cotyledon and radical development.

Germination stimulator and inhibitors

Although hormones have been shown to promote germination, it is by no means clear-cut.
Endogenous and exogenous levels of hormones were shown to affect the germination of seeds.
The role of endogenous levels or exogenously applied hormones e.g. indole acetic acid (IAA)
gibberellic acid (GA3) and kinetin on germination has been widely investigated.

Auxins: The relative concentrations of auxins, their precursors and other hormonal regulators
of growth are known to control the development of different plant parts. There was a rapid and
sharp increase in the endogenous IAA content of lupin, bean, maize, wheat and pine seeds in
response to water imbibition, especially during the time of radicle emergence.

Gibberellins: Gibberellic acid (GA3) stimulates the germination of seeds. Furthermore, some of
the known gibberellins occur in bound forms in fruits and seeds; these being converted to free
gibberellins during germination of seeds.

Cytokinins: Kinetin promotes the germination of seeds. Zeatin, its riboside and ribotide are
natural cytokinins and were shown to stimulate germination, with lower activity in case of the
derivatives than zeatin itself. The cytokinins are actively metabolized in germinating seeds.

Abscisic acid: Abscisic acid (ABA) was detected just prior to germination in some crop seeds. It soon
disappeared after the emergence of radicle. In Zea mays root tip extracts, ABA and xanthoxin were
confirmed to the root cap.

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Ethylene: When exogenously applied it stimulates germination. Germinating seeds are able to produce
ethylene. Some ethylene effects are gene activation and formation of some mRNA and membrane
integrity, changes in the level of hydrolytic enzymes, endogenous auxins.

Effect of coumrin and thio-urea: Seed treatment with coumarin resulted in reduction in % germination,
while thiourea stimulated % germination. Coumarins interfere with various metabolic process during
germination. Coumarin prevents the rise of lipases in fatty seeds. Nitrogen metabolism was also affected
by coumarin where it prevented any rise in soluble nitrogen during germination. It inhibits breakdown of
proteins due to inhibition of proteases.

Chemical composition of seed

About 70% of all food for human being comes from seed and remaining 30% from animals
indirectly. In addition to the normal chemical constituents found in plant tissues seed contains
extra substances, food reserves to support early seedling growth. The major food reserves
1. Carbohydrates
2. Proteins
3. Fats and oils
The minor food reserves: Phytin, Raffinose and oligosaccharides, Proteinase inhibitors, Lectins
and Alkaloids.
Chemical composition of seed is determined by: Genetic factors, Agronomic practices and
Environment
Average % composition: (Bewley and Black. 1994)

Crops Protein Oil Carbohydrates Major storage

organs

Barley 12 3 76 Endosperm

Maize 10 5 80 Endosperm

Oats 13 8 66 Endosperm

Rye 12 2 76 Endosperm

Wheat 12 2 75 Endosperm

Broad bean 23 1 56 Cotyledons

Garden pea 25 6 52 Cotyledons

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 Peanut 31 48 12 Cotyledons

Soybean 37 17 26 Cotyledons

Castor bean 18 64 Negligible Endosperm

Oilpalm 9 49 28 Endosperm

Rapeseed 21 48 19 Cotyledons

Carbohydrates: Major storage substances in seed. Cereal and grasses are rich in carbohydrates
and low in proteins and fats. Major forms of carbohydrate storage in seed are
A. starch
B. Hemicellulose
• Amyloids and raffinose series oligosaccharides are present as minor carbohydrate
reserves.
• Sometimes non storage forms like cellulose, pectins and mucilages occur.

Carbohydrate storage in seed: Starch Polysaccharide (…-glucose-glucose-glucose-…).

Two forms: 1. Amylose and 2. Amylopectin.

Starch grains in starchy endosperm cells of rye: 50-75% amylopectin & 20-25% amylase.

• Sl: large starch grain, Ss: small starch grain, p: protein matrix, w: cell wall

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Hemicellulose: Found in cell wall of plants and seed as reserves. It is the major storage
carbohydrate in endospermic legumes. Many hemicelluloses are mannans with β(1-4) linkage
between mannose units. Galactomannans have a β(1-6) linkage at galactose side chains.
Xyloglucans are known as amyloids.
Chemical structure of galactomannan;

Mucilages: are complex carbohydreates consisting of polyuronides and galacto uronides that
chemically resemble pectic compounds and hemicelluloses.
Pectic compounds: Found in primary cell wall and middle lamella. Occurs as pectic acid pectin
and propectin. During ripening of fruits propectin is converted into pectins.

Lipids –(Triacylglycerides/fats/oil): Simple lipids- fats, fatty oils and waxes-major

• Compound lipids-phospholipids, glycolipids

• Derived lipids- cholesterol
• The predominant FA in seeds are unsaturated FA (UFA)
• Lipids are stored in oil storage bodies called spherosomes which range in size 0.2 to 6.0
microns diameter.
• Enzymes for lipid synthesis & hydrolysis are present in spherosomes
• During germination lipids disappear and sucrose content in increases
• Lipase activity increases and hydrolysis tryglycerides to di and monoglycerides
• High lipid content is associated with low protein content

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Proteins
Osborne classified proteins into 2 categories
Metabolically inactive
• Prolamine
• Glutelin
Metabolically active
• Albumin and globulins
Albumins and globulins are not deficient in specific amino acids so they are good sources of
dietary protein. Storage proteins are oligomeric. Most storage proteins are not single but
bounded together by intermolecular disulphide groups, hydrogen bonding ionic bonding
and hydrophobic bonding. In legumes the major storage protein is globulins, which account
for 70% of total seed nitrogen. In addition to these proteins glycoproteins like lectins are
present in seeds. Seed storage proteins are deposited in protein bodies. Some proteins are
part of defence mechanism against pests and predators;
• Arcelin- conifers resistance against bruchid beetles
• Chitinase increases resistance to fungal attack

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Fatty acids: Building blocks for triglycerides and phospholipids . A chain of carbon and hydrogen
atoms with a carboxyl
arboxyl group at the alpha end and a methyl group at the omega end. end

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Saturated and unsaturated fatty acids:

Conclusion:
Sucrose entering the developing oil seed is used mainly for synthesis of storage triacylglycerols
and proteins. The triacylglycerol
triacylglycerols synthesis can be considered in three parts: The production of
the glycerol back bone, The formation of fatty acids
acids, The esterification of glycerols with fatty
acids components to give triacylglycerols. Sucrose is translocated into the developing seed andan
converted to hexose phosphates and triose phosphates by the reaction of glycolysis. Fatty acid
synthesis occurs in the plastids utilizing acetyl
acetyl-coA,
coA, which can be generated by glycolytic
reactions in the organelle.

Fruit and seed development

The ovary
ry of the flower contains the ovules. As fertilized ovules develop into seeds, the ovary
wall develops into the fruit. In science, the term “fruit” refers to a mature ovary that contains
seeds. In flowering plants – Fruit is a mature, ripened ovary th that contains
ntains the seeds pericarp
the ovary wall.

Fruits are formed by seed plants to aid in dispersing seeds

seeds. A seed contains the developing
plant embryo in a protective coat (testa)
(testa). Seeds form from ovules fertilized in the ovary.
ovary Ovaries
with seeds ripen into
o dry or fleshy fruits
fruits.

Fruit anatomy
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The ovules are fertilized in a process that starts with pollination, which involves the movement
of pollen from the stamens to the stigma of flowers. After double fertilization, these ovules will
become seeds. After pollination, a tube grows from the pollen through the stigma into the
ovary to the ovule and two sperm are transferred from the pollen to the megagametophyte.

Within the megagametophyte one of the two sperm unites with the egg, forming a zygote, and
the second sperm enters the central cell forming the endosperm mother cell, which completes
the double fertilization process. Later the zygote will give rise to the embryo of the seed, and
the endosperm mother cell will give rise to endosperm, a nutritive tissue used by the embryo.

As the ovules develop into seeds, the ovary begins to ripen and the ovary wall, the pericarp,
may become fleshy (as in berries or drupes), or form a hard outer covering (as in nuts). The
pericarp is often differentiated into two or three distinct layers i.e. exocarp (outer layer, also
called epicarp), mesocarp (middle layer) and endocarp (inner layer).

In some fruits, especially simple fruits derived from an inferior ovary, other parts of the flower
(such as the floral tube, including the petals, sepals, and stamens), fuse with the ovary and
ripen with it. In other cases, the sepals, petals and/or stamens and style of the flower fall off.
When such other floral parts are a significant part of the fruit, it is called an accessory fruit.

Fruit types

A. Simple fruit: A fruit that develop from a single ovary e.g. cherries, tomato.

B. Aggregate fruit: When several ovaries are formed within one flower e.g. Strawberry.

C. Multiple Fruit: In some plants such as pineapple, the simple fruit of many separate flower
fuse together to form multiple fruit e.g. pineapple.

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Stages of fruit development

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From ovule to seed

Mature seed

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 Cont…….

Cont…….
Seed development is triggered by a double-fertilization process resulting
in the differentiation of the embryo, endosperm, and seed coat that are
the major regions of the seed and essential for seed viability and
reproduction.
The genes required to program the differentiation and developmental
processes required for seed development are largely unknown.

Seed development

Modes of Reproduction: The modes of reproduction in crop plants may be broadly grouped into two
categories.

Asexual Reproduction: Asexual reproduction does not involve fusion of male and female gametes. New
plants may develop from vegetative parts of the plant (vegetative reproduction).

Sexual Reproduction: Sexual reproduction involves fusion of male and female gametes to form a zygote,
which develops in to an embryo. In crop plants, male and female gametes are produced in specialised
structures known as flowers.

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Sporogenesis: Productions of microspores and megaspores is known as sporogenesis. Microspores are
produced in anthers (microsporogenesis), while mega spores are produced in ovules
(megasporogenesis).

Microsporogenesis: Each anther has four pollen sacs, which contain numerous pollen mother cells
(PMCs). Each PMC undergoes meiosis to produce four haploid cells or microspores. This process is
known as microsporogenesis.

Megasporogenesis: Megasporogenesis occurs in ovules, which are present inside the ovary. A single cell
in each ovule differentiates into a megaspore mother cell. The megaspore mother cell undergoes
meiosis to produce four haploid megaspores. Three of the megaspores degenerate leaving one
functional megaspore per ovule. This completes megasporogenesis.

Gametogenesis

The production of male and female gametes in the microspores and the megaspores, respectively, is
known as gametogenesis.

Microgametogenesis

This refers to the production of male gamete or sperm. During the maturation of pollen, the microspore
nucleus divides mitotically to produce a generative and a vegetative or tube nucleus. Shortly after
pollination, the pollen germinates. The pollen tube enters the stigma and grows through the style. The
generative nucleus now undergoes a mitotic division to produce two male gametes or sperms. The
pollen, along with the pollen tube, is known as microgametophyte. The pollen tube finally enters the
ovule through a small pore, micropyle, and discharges the two sperms into the embryo sac.

Megagametogenesis.

The nucleus of a functional megaspore divides mitotically to produce four or more nuclei. In most of the
crop plants, megaspore nucleus undergoes three mitotic divisions to produce eight nuclei. Three of
these nuclei move to one pole and produce a central egg cell and two synergid cells; one synergid is
situated on either side of the egg cell. Another three nuclei migrate to the opposite pole to give rise to
antipodal cells. The two nuclei remaining in the centre, the polar nuclei, fuse to form a secondary
nucleus. The megaspore thus develops into a mature megagametophyte or embryo sac. The
development of embryo sac from a megaspore is known as megagametogenesis. The embryo sac
generally contains one egg cell, two synergids, three antipodal cells (all haploid), and one diploid
secondary nucleus.

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 Gene imprinting

Introduction:-

 People inherit two copies of their genes

genes—oneone from their mother and one from their
father.
 Usually bothh copies of each gene are active, or “turned on,” in cells.
 In some cases, however, only one of the two copies is normally turned on.
 Which copy is active depends on the parent of origin: some genes are normally active
only when they are inherited from a person’s father; others are active only when
inherited from a person’s mother.
 This phenomenon is known as genomic imprinting.

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Definition:- “Genomic imprinting is an epigenetic phenomenon leading to a change of gene expression
dependent on whether the gene was inherited from the maternal or the paternal parent (Reik and
Walter, 2001)”.

“Genomic imprinting refers to genes that are silent when maternally inherited but expressed when
paternally inherited, or vice versa”.

Gene imprinting is a differential expression of autosomal genes according to their parent of origin.

Paternal imprinting means that an allele inherited from the father is not expressed in offspring.
Maternal imprinting means that an allele inherited from the mother is not expressed in
offspring.

Epigenetics: “The study of changes in gene function that are heritable and that do not entail a
change in DNA sequence”

Discovery of genomic imprinting:

First report by Helen Crouse in 1960.
The first description of the imprinting phenomenon was given by McGrath and Solter
in 1984
The word “ Imprinting” was first used to describe events in insect Pseudococcus nipae.
In plants first demonstrated in maize by Kermicle (1970)
The study of gene specific imprinting during seed development in Arabidopsis.

What is imprinting ?
 Imprinted chromosomes are “marked”
 Imprinting differs in sperm and egg
 Imprinting turns off specific genes
 Imprinting does not change the nucleotide sequence.

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  It is an inheritance process independent of the classical mendelian inheritance.
 If the allele inherited from the father is imprinted and thereby silenced then only the
allele from the mother is expressed.(in the case of gene H19 or CDKN1C).
 If the allele from the mother is imprinted then only the allele from the father is
expressed (e.g. in the case of gene IGF-2).

Imprinting

 Most genes.
 Inherit working copies.
 One from mom, other from dad.
 Imprinting.
 Inherit one working copy.
 Depending on gene, copy from
mom or dad is epigenetically
silenced.
 Silencing occuring through
addition of methyl groups
during egg/sperm formation

Types of gene imprinting:

1. Allelic Imprinting: In which only alleles from a certain background are subject to parent-
of origin–specific gene expression.
Example: An imprinted angiosperm gene was in alleles of the maize R gene. The R gene
conditions anthocyanin accumulation in the aleurone (the outer cell layer of the endosperm) of
maize kernels. When an RR female (red) is mated to a rr male (colorless), all of the kernels have
a fully coloured aleurone. However, the reciprocal cross gives rise to kernels with mottled
aleurone pigmentation, indicative of irregular anthocyanin distribution (Kermicle, 1970). This
phenomenon is specific to the endosperm, and no reciprocal differences are observed in
embryos or seedlings (Brink et al., 1970).

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2. Locus imprinting: In which all known alleles from different backgrounds are under parent-
of origin control.

Example: The Arabidopsis MEDEA (MEA) gene

DNA methylation (Cont..)

Mechanism
 DNA methylation may stably alter the expression of gene in cells
as cell devide and differentiate from embryonic stem cells into # Methyl groups are transferred
specific tissues. from S-adenosyl methionine
in a reaction catalysed by a
 DNA methylation is typically removed during zygote formation DNA methyl transferases
and re-established through successive cell divisions during (DNMT).

development.
# SAM is then converted to
SAH(S-adenosyl
homocysteine).

DNA methylation regulates imprinted genes in plants

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Imprinted genes in plants:

 In flowering plants (angiosperms).

 During fertilisation of the egg cell, a second, separate fertilization event gives rise to the
endosperm, an extra-embryonic structure that nourishes.
 Unlike the embryo, the endosperm is often formed from the fusion of two maternal
cells with a male gamete. This results in a triploid genome.
 The 2:1 ratio of maternal to paternal genomes appears to be critical for seed
development.
 It has been suggested that these imprinted genes are responsible for the triploid block
effect in flowering plants that prevents hybridization between diploids and auto-
tetraploids.

Role of genomic imprinting in seed development

 Seed development in flowering plants is initiated by double fertilization.
 One haploid sperm fuses with the haploid egg to produce the diploid embryo and the
second haploid sperm fuses with the diploid central to form the triploid endosperm.
 The embryo has a maternal: paternal parental genomic ratio of 1:1 (1m:1p) whereas
the ratio in the endosperm is 2m:1p.
Conclusion
 Genetic imprinting is found to have major role in many key developmental processes
and genome dosage is one of the factors contributing to the imprinting.
 To date, the scientific community is still debating on its role in evolution and significance
in the process of crop improvement.
 Imprinted gene regulates the transfer of nutrients to the developing progeny.
 Advanced technologies like genome-wide approaches may contribute in helping the
researchers to unravel the potential mechanism of genetic imprinting and its possible
benefits to crop improvement.

Seed and Fruit Abortion, Proximate Mechanism of Seed and Fruit Abortion

Fruit abortion is nonrandom with respect to damage (such as that caused by seed predators
and bad weather) and with respect to the number of seeds within a fruit (the more the better).
The immature fruits abort when the number of pollinated flowers exceeds the resources
available for fruit production; that aspects of fruit abortion have been investigated by
physiologists, horticulturalists, foresters, and others in the plant sciences since the turn of the
century.
Seed abortion, a common phenomenon in flowering plants, is the mortality of immature seeds
between fertilization and seed maturation (Bawa & Webb 1984).
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What Happens To A Flower After Fertilization ?
• The flower’s job is done after fertilization. Much of the plant’s energy is used to develop
the ovary and the ovules.
• In several weeks, the fertilized ovule ripens and b
becomes a seed.
• The ovary also ripens and becomes a fruit.
• If an ovary contains several ovules, each ovule can form a seed.
• The petals and stamens of the flower are no longer necessary, and they often wither.
• In many plants, however, the receptacle and the calyx become part of the fruit.

FRUIT AND SEED

• A fruit is a mature (ripened) ovary that contains one

or more seeds.

• A seed is a mature ovule consisting of the following parts:

– Seed coat (for protection)

– Embryo (immature plant)

– Endosperm (for food)

• An embryo can consist of the following parts:

– One cotyledon (monocot plants)

– Two cotyledons (dicot plants)

– Epicotyl that forms leaves

– Hypocotyl that forms stem and root

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 Factors that limit fruit and seed
production
Number of fruits number of female flowers

Number of seeds number of ovules

This reproductive potential depend upon:

• number of pollinated flowers

• number of fertilized ovules

• fruit/seed predation

• weather conditions

• ability of the maternal parent to provide the necessary

resources for development

Growth hormones:
Growth hormones (auxins, gibberellins, and cytokinins) play important roles in the growth and
development of fruit and seeds. This transition from a flower to a developing fruit is termed
"fruit set" and is usually accompanied by the wilting or abscission of petals and stamens. In
some species, exogenous applications of hormones allow fruit set in the absence of pollen.
Hormones produced by the seeds also play a leading role in the mobilization of resources into
the developing fruits.

Factors That Limit Fruit Set:

Flowers must be pollinated in order to set fruits except in apomictic and parthenocarpic
species. This does not imply that only unpollinated flowers abscise or that pollination
necessarily limits fruit set. The number of fruits that set is usually determined by resources
rather than the number of morphologically female flowers, and Natural levels of pollination
exceed fruit set in many species.
Evidences:
• In Wisconsin, a six-year investigation of the sour cherry, Prunus cerasus, revealed the
presence of pollen tubes in the styles of 82-100% of the abscised flowers.
• Among milkweeds, 95% of the flowers of Asclepias syriaca had a pollinium inserted into
at least one of the five stigmatic chambers, but only 9.6% of the flowers set fruits.
And the reason suggested for this that some of these species may have abscised flowers that
received incompatible pollen.
In general, the proportion of pollinated flowers that set fruit decreases as the number of
pollinated flowers increases. (A. G. Stephenson, 1981)

Factors affecting seed and fruit abortion:

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 Water deficit during reproduction

Lack of pollination

Weather conditions

Seed predation

Growth hormones

Abiotic and Biotic agents

Factors That Promote Fruit Abortion

The proportion of juvenile fruits and seeds that mature is dependent upon extrinsic factors,
such as
• Weather conditions
• Seed predation,
• and the ability of the maternal parent to provide the resources necessary for growth
and development.

Shedding of damaged fruits:

• Many abiotic and biotic agents dam
damage fruits and promote abscission.
• Late frosts during the spring are occasionally a principal cause of fruit mortality.
• Hard (1963) observed that Pinus resinosa trees located on lower ground lost all of their
first-year
year conelets to a late spring frost, whe
whereas
reas trees on higher ground abscised
conelets from only the lower parts of their crowns.

• Seed predation by insects also causes selective abscission of young fruits. For example, a
stand of Cassia grandis aborted approximately 95 % of the initiated fruits anda 81 % of
these were insect damaged.

Shedding of ubdamaged fruits:

• Some undamaged juvenile fruits abscise because of genetic or developmental

abnormalities.
• However, most abortions of undamaged fruits seem to be a response to limited
resources.
• If resources
ces are limited, competition among fruits and subsequent abortion are likely to
increase with the number of fruits initiated. In most cases, the proportion of fruits that
survive is a decreasing function of the number of fruits initiated.

1. Water deficit during

ring reproductive development was shown to be a dominant
environmental factor accelerating the rate of abortion (Kato, 1964; Westgate &

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 Peterson, 1993). Based on these findings, Raper & Kramer (1987) concluded that water
stress imposed during flower development reduces photosynthesis and the amount of
photoassimilates allocated to reproductive tissues, thereby accelerating the rate of
abortion.
2. Another possible physiological factor affecting flower abortion in soybean is the
availability of plant hormones (Huff & Dybing, 1980; Heindl et al., 1982; Carlson et al.,
1987). There are a number of reports showing that the application of 6-
benzylaminopurine (BA) to racemes reduced the rate of abortion and thereby increased
pod set.
3. Role of phytohormones in flower development in soybean

The application of BA to racemes reduced seed abortion and increased pod number (Crosby
et al., 1981). soybean abortion is primarily caused by deficiency in or competition for
photoassimilates and nutrients among growing organs (Brevedan et al., 1978; Brun & Betts,
1984; Antos & Wiebold, 1984).

Environmental causes:
1. Temperature: Extreme temperature such as high daytime temperatures (above 85 0F /
29 0C), or high nighttime temperatures (above 70 0F / 21 0C), or low nighttime
temperatures (Below 55 0 F / 13 0 C) tomato plants will drop the flowers.
2. Humidity: The ideal humidity range is between 40-70%. If humidity is either too high or
too low, it interferes with the release of pollen and with pollen’s ability to stick to the
stigma. So pollination will not occur.
3. Other potential sources of blossom drop:

a. Lack of pollination: Tomatoes need some help to pollinate. Insects, wind or hand shaking of
the flowers is necessary to carry the pollen from the anthers to the stigma. During extremes
weather conditions, there are often no insect pollinators.

b. Nitrogen: High or low application rates of N fertilizer can cause blossom drop. High rates of
nitrogen fertilization encourage lush vegetative growth and inhibit flower production and/or
pollination, resulting in poor fruit set. Low N produces spindly vines with low food reserves that
cannot support a crop.

c. Lack of water: Shallow watering will stress and weaken the plants. The root zone should be
uniformly moist throughout the growing season to develop a large, healthy root system.

d. Insect damage or disease: Use good cultural practices and treat for disease as soon as
symptoms appear.

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e. Heavy fruit set: When a tomato plant has too many blossoms, the resulting fruits are all
competing for the limited food supplied by the crop. The plant will automatically abort some
flowers. Once the initial crop is harvested, the problem should subside.

Conclusion

The yield is determined by the number of pods (seeds) produced per unit area and individual
seed weight. The seed number depends upon the number of floral buds that initiate pods and
attain maturity. Generally, plants produce an abundance of floral buds, but a large proportion
of them abort during development. The rates of flower and pod abscission/abortion were
estimated to reach 80% (Shibles et al., 1975). Alleviation of this abortion rate should increase
pod and seed number, and thereby increase yield.

Chemical Composition of Seeds, Seed storage Protein

The plant seed is not only an organ of propagation and dispersal but also the major plant
tissue harvested by humankind. The amount of protein present in seeds varies from 10% (in
cereals) to 40% (in certain legumes and oilseeds) of the dry weight, forming a major source
of dietary protein. Although the vast majority of the individual proteins present in mature
seeds have either metabolic or structural roles, all seeds also contain one or more groups of
proteins that are present in high amounts and that serve to provide a store of amino acids
for use during germination and seedling growth. These storage proteins are of particular
importance because they determine not only the total protein content of the seed but also
its quality for various end uses. In the case of wheat, the storage proteins form the gluten
fraction, whose properties are largely responsible for the ability to use wheat flour to make
bread, other baked goods, and pasta. These properties are not shared by the storage
proteins of other cereals.

Seeds may be divided into those whose main storage materials is carbohydrates and those
whose main storage material is lipid. Seed containing proteins can belong to either group.
Almost no seeds are known in which the predominant storage material is protein, although
there are exceptions, such as soybean that has been reported to contain 66% protein.

Seed storage Protein: According to their solubility in water and salt solutions protein were
classified as follows:

Protein type Solubility

1. Albumins * In water and salt solution

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 2. Globulins * In salt solution

3. Prolamins * Soluble in 70-80% ethanol but not soluble in water

or absolute alcohol.

4. Glutelins * Soluble only in acids and alkaline

Characteristics of Seed Storage Proteins

 First, they are synthesized at high levels in specific tissues and at certain stages of
development.
 Many seeds contain separate groups of storage proteins, some of which are rich in
sulfur amino acids and others of which are poor in them.
 Second common property of seed storage proteins is their presence in the mature seed
in discrete deposits called protein bodies, whose origin has been the subject of some
dispute and may in fact vary both between and within species.
 Finally, all storage protein fractions are mixtures of components that exhibit
polymorphism both within single genotypes and among genotypes of the same species.
 This polymorphism arises from the presence of multigene families and, in some cases,
proteolytic processing and glycosylation.

(a) 2s ALBUMIN STORAGE PROTEINS

The 2s albumins were initially defined as a group on the basis of their sedimentation
coefficients (S20.wo) f -2 (Youle and Huang, 1981). They are widely distributed in dicot
seeds and have been most widely studied in the Cruciferae, notably oilseed rape (in which
they are called napins) and Arabidopsis. The napins consist of two polypeptide chains with
Mr values of 9000 and 4000, which are linked by interchain disulfide bonds. This appears to
be the most typical 2s albumin structure: similar heterodimeric proteins are present in
species as diverse as pumpkin (Hara-Nishimura et al., 1993), Cotton. castor bean.

(b)PROLAMIN STORAGE PROTEINS

Whereas the 2s albumin and globulin (see later discussion) storage proteins are widely
distributed in flowering plants, the prolamins are restricted to one family, the grasses.
These include the major cereals, in which prolamins usually account for approximately half
of the total grain nitrogen. Exceptions to this general rule are oats and rice, in which the
major storage proteins are 11s globulin-like and prolamins are present at low levels (5 to
10Vo of the total grain protein). Prolamins are traditionally recognized as a group on the
basis of their solubility in alcohol hater mixtures (usually 60 to 70% ethanol) and their high
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 levels of glutamine and proline. However, comparisons of amino acid sequences have
shown that this definition must be widened to include components that are insoluble in
aqueous alcohols in the native state due to the presence of inter chain disulfide bonds and
to recognize that all prolamins, even those that are insoluble in aqueous alcohols, are
related, except for the a-zeins of maize (and their homologs present in related Panicoid
cereals).

(c) GLOBULIN STORAGE PROTEINS

The globulins are the most widely distributed group of storage proteins; they are present not
only in dicots but also in monocots (including cereals and palms) and fern spores (Templeman
et al., 1987). They can be divided into two groups based on their sedimentation coefficients
(S20.w):

(1) 7S vicilin-type globulins and

(2) 11S legumin-type globulins.

Both groups show considerable variation in their structures, which results partly from post
translational processing. The globulin storage proteins have been studied in most detail in
legumes, soybean, broad bean, and French bean (Phaseolus vulgaris).

1. The 7S Globulins

7S globulins are typically trimeric proteins of Mr ~150,000 to 190,000 that lack cysteine
residues and hence cannot form disulfide bonds. Their detailed subunit compositions vary
considerably, mainly because of differences in the extent of post-translational processing. For
example, the vicilin subunits of pea are initially synthesized as groups of polypeptides of Mr
~47,000 and ~50,000, but post-translational proteolysis and glycosylation then give rise to
subunits with Mr values between 12,500 and 33,000.

2. The 11S Globulins

The 11S legumins are the major storage proteins not only in most legumes but also in many
other dicots (for example, brassicas, composits, and cucurbits) and some cereals (oats and rice).
The mature proteins consist of six subunit pairs that interact noncovalently. Each of these
subunit pairs consists in turn of an acidic subunit of Mr ~40,000 and a basic subunit of Mr
~20,000, linked by a singlepair is synthesized as a precursor protein that is proteolytically
cleaved after disulfide bond formation. Each subunit Although the 11S globulin of Brazil nut was
one of the first proteins to be crystallized (Maschke, 1859), the crystals of this and other 11S
globulins have generally been small and disordered and have failed to provide any details of
protein structure.

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Experimental Plan of seed storage proteins

1. Isolating and grinding the endosperm.

2. Total Endosperm Protein Isolation
3. Zein Protein Isolation
4. Preparing samples for PAGE
5. Visualizing the Zein Proteins

For example:- Total Endosperm Protein Isolation

a. Weigh 100 mg of the powdered endosperm meal and transfer to a microfuge tube.
b. Add 1 ml of 3X sample buffer. Vortex vigorously for 1 minute.
c. Put the microfuge tube at 65o C for 15 minutes, vortexing to resuspend the meal every 5
minutes.
d. Pellet the insoluble material in the micro-fuge for 5 minutes and transfer the
supernatant (avoid the pellet) to a clean micro-fuge tube.
e. To resolve the proteins on the basis of their size by SDS-PAGE (Poly-Acrylamide-Gel-
Electrophoresis), each lab group should prepare two samples of wild-type and two of
opaque-2 total proteins as follows:
µl of Extract µl 3X Sample Buffer µl of Water

30 20 100

50 0 100

Later, these samples must be boiled before loading on the gel to completely denature
the protein in the presence of the SDS. For now, place these aside until the zein protein
fraction is also ready to boil.

Conclusion

Much of the recent work on seed storage proteins was performed to provide a basis for
improving the nutritional and processing properties of crops using genetic engineering. The
recent development of reliable transformation procedures form maize, small grain cereals
(wheat and barley), and grain legumes means that this is now possible. For example, much of
the work on engineering 2s albumins has been based on manipulation of a variable “loop”
region identified by sequence comparisons, whereas Wallace et al. (1988) used a structural
mode for zein (Argos et al., 1982) to identify sites for the addition of lysine residues.

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 Seed respiration

Seed is a ripen ovule containing an embryo in its arrested state of development. Respiration is a
process of oxidizing food to release energy inside cells. Plants are known for their ability to
convert Carbon dioxide into Oxygen. However, all aerobic organisms take in Oxygen and give
off Carbon dioxide as long as they are alive. During germination, seeds use sugars and other
molecules as a substrate for respiration.

Seeds don't breathe in the same way mammals do. Instead, they breathe at a cellular level. In
cellular respiration, the seed uses stored sugars, water and oxygen to burn energy at a cellular
level and germinate, or sprout. Respiration increases dramatically as the seed sprouts. The seed
continues to breathe until the plant can make its own food via the process of photosynthesis.
The oxygen comes from tiny pockets of air in the soil. Most loose soil has plenty of air for seeds,
but if the seed is surrounded by water, it will not be able to obtain enough oxygen to
germinate.

Seeds are alive

To function in propagation, seeds must be alive. Seeds respire, a bit slowly. Consume O2,
produce CO2 and H2O. Seeds have a finite life span cannot be stored indefinitely.

Respiration

Respiration Glycolysis
• Occurs in all living organisms
 reactions are catalyzed by enzymes • Only stage which can occur without oxygen
• Oldest stage of respiration
 main food substance which oxidized in cells
– operated for billions of years in anaerobic organisms
is glucose • Converts glucose to 2 pyruvates in cytosol
enzymes
C6H12O6 + 6O2 6CO2 + 6H2O + energy – with O2 goes on to TCA cycle
– without O2 pyruvate is converted to lactate or ethanol
 as it takes place in all living cells, it is called (fermentation)
cellular respiration which is used to produce • Yields 2ATP/mole glucose in the absence of O2
energy for cells to use

3 Stages of Respiration

• Glycolysis
• TCA Cycle
• Electron Transport Chain

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 Glycolysis
TCA cycle
Glucose (6C)

2 Pyruvate (3C)

CO2 -O2 -O2

+O2
Ethanol Lactate
TCA Cycle

Electron Transport System

NADH and FADH2

e-
H+

ATP
e-
H+

4e- + 4H+ + O2 2H2O

cyt. oxidase

Mitochondria
3 Stages of Respiration
• Spherical to oval
• Glycolysis
– about 1 micron diameter
– cytoplasm
– with or without oxygen present – # mito./cell increases with demand for
– breaks glucose (6C) into 2 pyruvates (3C) respiration; 300-1000/root tip cell cristae
• TCA Cycle • Double-membrane bound
– mitochondrial matrix – outer smooth
– only if oxygen present – inner folds forming cristae
– converts pyruvate via acetyl CoA into CO2; generates • controls movement in/out
NADH and FADH2
• site of electron transportm
• Electron Transport Chain matrix
• Matrix
– mitochondrial membranes = cristae
– transfers electrons from NADH and FADH2 to reduce – soluble phase
O2 to H2O and generate ATP – site of TCA cycle; DNA, RNA, ribosomes

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 Alternate Fates of Glucose C Factors Affecting Respiration
• Not all C respired to CO2 • Kind of Cell or Tissue
• Intermediates of respiration branch off: – Young and Developing Cells (Meristematic Areas)
– amino acids usually Have Higher Respiration Rates
– pentoses for cell wall structure – Developing and Ripening Fruit and Seeds, too
– nucleotides – Older Cells and Structural Cells Respire at Lower
– porphyrin biosynthesis Rates
– fatty acid synthesis
– lignin precursors
– precursors for carotenoid synthesis,
hormones

Factors Affecting Respiration Factors Affecting Respiration

• Temperature
– Respiration generally Has Higher Optimum and • Oxygen
Maximum Temps than PS Rxs – Low O2 Can Reduce Aerobic Respiration
– Can Have Net Dry Matter Loss at High Temps and Increase Anaerobic Respiration
where Respiration Exceeds PS – Low O2 Can Reduce Photorespiration
– Temp Refers to Temp Inside Plant or Animal Cell,
not Air Temp
– Using Irrigation to Help Cool the Plant Can Keep
the Plant in Net Gain Range

• Light

– Can Enhance Rate of Photorespiration

– Does not Directly Affect other Forms of Respiration

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