[Link].

Botany Sem-IV | Seed Technology | Unit 1 – Seed Science & Physiology

[Link]. Botany – Semester IV (NEP)

Botany Elective 8: SEED TECHNOLOGY
UNIT 1: Introduction to Seed Science and Seed
Physiology
Detailed Study Notes
Session: 2024–2025

References: Copeland & McDonald (2001) · Egli (2017) · Basra (2006) · Elias et al. (2012) ·
Winn (2014)

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1.1 Seed Biology Basics

1.1.1 Seed Structure and Types

A seed is a mature ovule that contains a miniature embryonic plant along with stored
nutritive tissue, enclosed within a protective seed coat. Seeds represent the primary unit of
sexual reproduction in spermatophytes (gymnosperms and angiosperms). Understanding
seed structure is fundamental to all aspects of seed science, technology, and agronomy
(Copeland & McDonald, 2001).

A. General Plan of a Seed

Every seed, regardless of species, consists of three fundamental components:

 Seed coat (testa + tegmen): The outermost protective layers derived from
integuments of the ovule. The testa is the outer coat and the tegmen is the inner
coat. Together they shield the embryo from mechanical damage, pathogens,
desiccation, and unfavourable temperatures.
 Embryo: The miniature sporophyte developing from the fertilised egg. It
comprises the embryonic axis (plumule + hypocotyl + radicle) and one or two seed
leaves (cotyledons). The embryo is the only living, metabolically active part of a
mature quiescent seed.
 Food reserve tissue: Nutritive tissue that supports early seedling growth. In
dicots, reserves accumulate mainly in the fleshy cotyledons; in monocots
(especially grasses), reserves are stored in a specialised starchy endosperm. In
endospermic seeds, the endosperm persists at maturity; in non-endospermic
(exalbuminous) seeds, the endosperm is fully absorbed by cotyledons during
development.

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Figure 1.1 – Longitudinal section of a Dicot Seed (Bean, Phaseolus vulgaris) showing all structural components

B. Structural Components in Detail

(i) Seed Coat (Testa)
The testa is derived from the outer integument of the ovule. Its thickness, texture, and
chemical composition vary enormously among species—from thin papery coats in garden
peas to the extremely thick, impermeable, lignified coats of hard-seeded legumes.
Structurally, the testa typically consists of:

 Epidermis: Outermost cell layer, often bearing a waxy cuticle or mucilaginous

cells. In Fabaceae, palisade (Malpighian) cells form a thick, highly lignified
epidermis that creates physical dormancy.
 Sub-epidermis (Hourglass cells): Columnar sclerenchymatous cells that provide
rigidity.
 Parenchyma layers: Inner ground tissue, sometimes compressed at maturity.

(ii) Hilum, Micropyle, and Raphe

Hilum is the scar left where the funicle (stalk) was attached; it acts as a hygroscopic valve
regulating water and gas exchange. The micropyle is a minute pore (remnant of the ovular
micropyle) through which the pollen tube entered; it serves as the primary water-entry

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point during imbibition. The raphe is a ridge of tissue corresponding to the fused funiculus,
present in anatropous ovules (most dicots).

(iii) Embryo Components

Component Description Future Fate

Plumule Apical meristem + embryonic Shoot system

leaves (eophylls)

Hypocotyl Axis between cotyledon Lower stem / elongates in

attachment & radicle epigeal germination

Radicle Embryonic root with root cap Primary root

Cotyledon(s) Seed leaf; 1 in monocots, 2 in Absorb or photosynthesise as

dicots seed leaves

Coleoptile Sheath protecting plumule in Pushes through soil, becomes

grasses leaf-sheath

Coleorhiza Sheath protecting radicle in Ruptured as radicle elongates

grasses

Scutellum Single cotyledon in grasses; Absorbs and digests endosperm

enzyme-secreting face towards reserves
endosperm

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Figure 1.2 – Longitudinal section of a Monocot Seed (Maize grain, Zea mays) showing fused pericarp-testa, starchy
endosperm, aleurone layer, and embryonic structures

C. Types of Seeds
Seeds are classified on multiple criteria:

(i) Based on endosperm persistence

 Albuminous / Endospermic seeds: Endosperm persists in the mature seed and
serves as the primary food reserve. Examples: monocots (wheat, maize, rice), and
some dicots (castor, coffee).
 Exalbuminous / Non-endospermic seeds: Endosperm is completely absorbed by
cotyledons during development; food reserves are stored in thick fleshy
cotyledons. Examples: most legumes (pea, bean), sunflower.

(ii) Based on number of cotyledons

 Monocotyledonous: Single cotyledon (scutellum). Characteristic of Poaceae
(grasses), Liliaceae, Orchidaceae.
 Dicotyledonous: Two cotyledons. Characteristic of most non-grass angiosperms—
Fabaceae, Solanaceae, Brassicaceae, etc.

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 Polycotyledonous: Three or more cotyledons (polyembryony). Occurs in

gymnosperms (Pinus) and rarely in angiosperms.

(iii) Based on seed moisture at maturity (Roberts, 1973)

 Orthodox seeds: Tolerate desiccation to < 5% moisture content without loss of
viability; viability increases as moisture and temperature are lowered. Majority of
crop species (cereals, legumes, many vegetables).
 Recalcitrant seeds: Cannot tolerate desiccation; die if moisture drops below 20–
30%. These seeds have no dormant phase. Examples: Mangifera indica (mango),
Hevea brasiliensis (rubber), Avicennia species.
 Intermediate seeds: Intermediate tolerance; can withstand moderate desiccation
but not the extremes tolerated by orthodox seeds. Examples: Coffea arabica
(coffee), Carica papaya.

1.1.2 Seed Development and Maturation

Seed development (embryogenesis + reserve accumulation) is a tightly regulated process
that transforms the fertilised ovule into a dormant, desiccation-tolerant propagule.
According to Egli (2017), seed development passes through five overlapping stages.

Figure 1.3 – Five sequential stages of seed development from fertilisation to quiescence, showing relative embryo size
and reserve accumulation

Stage I – Post-fertilisation (Cell Division Phase)

Immediately after double fertilisation, the zygote undergoes a series of stereotyped cell
divisions. The first division is asymmetric, producing a smaller apical cell (gives the

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embryo proper) and a larger basal cell (gives the suspensor). Within 3–5 days in most crops,
a globular embryo is formed. The primary endosperm nucleus (triploid) divides rapidly to
form a liquid or cellular endosperm that bathes the embryo. Nutrient import from the
maternal tissue is initiated via the funiculus through phloem unloading. The testa begins
to develop from the integuments.

Stage II – Histodifferentiation (Embryo Organ Formation)

The embryo transitions through characteristic morphological shapes: globular → heart →
torpedo → walking-stick (in dicots). This transition is orchestrated by auxin gradients
(particularly PIN1-mediated polar auxin transport). By the torpedo stage, the shoot apical
meristem (SAM), root apical meristem (RAM), cotyledons, hypocotyl, and radicle are
distinct. In monocots, the scutellum, coleoptile, and coleorhiza become identifiable at this
stage. Endosperm cellularisation begins, and storage cells differentiate (protein bodies,
amyloplasts).

Stage III – Cell Expansion and Reserve Deposition (Filling Phase)

This is the most critical agronomic phase—seed dry weight accumulates predominantly
here. Storage compounds are deposited:

 Carbohydrates: Sucrose imported via phloem is converted to starch by ADP-

glucose pyrophosphorylase (AGPase) in amyloplasts. Starch is deposited as a
mixture of amylose (~25%) and amylopectin (~75%).
 Proteins: Amino acids (transported as asparagine, glutamine) are assembled into
storage proteins (albumins, globulins, prolamins, glutelins) deposited in protein
storage vacuoles (PSVs).
 Lipids: Fatty acids are synthesised in plastids, assembled as triacylglycerols
(TAGs) in the ER, and sequestered in oil bodies (oleosomes). Oleosins coat oil
bodies, preventing coalescence.

Accumulation rate follows a near-linear pattern; the filling period duration is influenced
by temperature (high temperatures shorten filling), source–sink ratio, and stress.

Stage IV – Maturation and Desiccation

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As seed filling concludes, seed moisture content falls dramatically from ~80% to < 15% in
orthodox seeds. Key events:

 ABA accumulation: Abscisic acid (ABA) peaks at late embryogenesis, inducing

Late Embryogenesis Abundant (LEA) proteins which protect cellular structures
during desiccation (protective glass-formation theory). ABA also suppresses
precocious germination by maintaining dormancy.
 Desiccation tolerance mechanisms: Sucrose vitrification (glass state), LEA
proteins, heat shock proteins (HSPs), reactive oxygen species (ROS) scavenging
enzymes (SOD, CAT, APX), and repair of DNA lesions.
 Seed colour change: Chlorophyll degradation and tannin/flavonoid synthesis alter
testa colour to species-specific shades—serving as visual maturity indicators.

Stage V – Quiescence / Readiness for Dispersal

The metabolic rate of the seed drops to near-zero (quiescent state). Moisture content
stabilises at 10–14% in most crop seeds. Seed mass is at its maximum (physiological
maturity) and viability is at its peak. The seed is now ready for dispersal, harvest, or
storage.

Key Concept: Physiological maturity ≠ Harvest maturity. At physiological maturity, seed

achieves maximum dry weight and viability; harvest maturity is when the seed/crop can be
mechanically harvested without damage losses.

1.2 & 1.3 Seed Germination and Physiology

1.2.1 Definition and Types of Germination

Germination is the resumption of metabolic activity and growth of a viable, non-dormant
seed following imbibition of water, leading to the emergence of the seedling from the seed.
More precisely (ISTA, 2021), germination is "the emergence and development from the seed
embryo of those essential structures which, for the type of seed in question, are indicative
of the ability to develop into a normal plant under favourable conditions."

A. Phases of Germination

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Germination follows three distinct phases of water uptake, characterised by the triphasic
imbibition curve (Bewley & Black, 1994; cited in Copeland & McDonald, 2001):

1. Phase I – Imbibition: Rapid, passive water uptake driven by matric potential of

seed colloids (proteins, starch, cell walls). Occurs even in dead seeds. Causes rapid
rehydration, membrane restructuring (transient leakage of solutes), and
reactivation of existing enzymes. Duration: minutes to hours.
2. Phase II – Activation (Lag phase): Water uptake plateau; metabolic activities are
resumed. mRNA repair and de novo transcription begin; mitochondria are
reconstituted; respiratory rate rises; lipid mobilisation commences. The cell cycle is
re-entered: S-phase DNA synthesis prepares for the first cell division. Duration:
hours to days.
3. Phase III – Radicle Protrusion: Renewed water uptake accompanies cell
elongation in the radicle. Turgor-driven expansion forces the radicle through the
micropyle/endosperm cap. This marks completion of germination sensu stricto.

B. Types of Germination Based on Cotyledon Position

Figure 1.4 – Comparison of epigeal and hypogeal germination showing cotyledon position relative to soil surface

(i) Epigeal Germination

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Definition: The hypocotyl elongates rapidly and carries the cotyledons above the soil
surface. Cotyledons turn green and photosynthesise for a brief period before withering.
The hypocotyl initially appears as a hook (hypocotyl hook / plumular hook) that protects
the delicate plumule during soil penetration.

 Examples: Bean (Phaseolus vulgaris), Castor bean (Ricinus communis), Onion

(Allium cepa), Mustard (Brassica).
 Significance: The emerged cotyledons contribute to photosynthesis early on,
supplementing reserve mobilisation.

(ii) Hypogeal Germination

Definition: The epicotyl elongates, and cotyledons remain below the soil surface (at or
below the point of sowing). The cotyledons function only as storage organs and are
absorbed/remain in soil.

 Examples: Pea (Pisum sativum), Chickpea (Cicer arietinum), all cereals—wheat,

rice, maize, barley.
 Significance: Cotyledon reserves are safer from surface temperature fluctuations
and herbivory; important for deep-sown seeds.

1.2.2 Factors Affecting Germination

A. Internal (Intrinsic) Factors
(i) Seed Viability
Viability refers to the ability of a seed to germinate and produce a normal seedling. It is
determined by genetic makeup, degree of maturation at harvest, and ageing during
storage. Seeds lose viability due to lipid peroxidation, protein denaturation, membrane
degradation, and DNA damage during storage.

(ii) Seed Vigour

Vigour is defined by ISTA as the sum total of those properties which determine the activity
and performance of seed lots of acceptable germination in a wide range of environments.
High-vigour seeds germinate rapidly, uniformly, and under sub-optimal conditions; they

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give stronger seedling establishment. Vigour differs from viability in that a seed may be
viable (capable of germinating under ideal conditions) but lack vigour (fails under stress).

(iii) Dormancy
Dormant seeds will not germinate even under ideal external conditions until dormancy is
broken (see Section 1.4). Dormancy is a major intrinsic impediment to germination and is
discussed in detail in later sections.

(iv) Seed Size and Quality

Larger seeds with more stored reserves germinate more vigorously and produce stronger
seedlings, especially under stress. Damaged, diseased, or immature seeds show reduced
germination.

B. External (Environmental) Factors

(i) Water (Moisture / Imbibition)
Water is the single most critical factor for germination initiation. The minimum water
potential (Ψ) required for germination (base water potential, Ψb) is species-specific,
typically ranging from −0.5 to −1.5 MPa. Mechanisms:

 Hydration of colloids and cytoplasm enables enzyme activation.

 Softening of seed coat allows physical expansion.
 Dissolved oxygen and mineral transport require an aqueous medium.
 Water is essential for hydrolytic enzyme activity (amylases, proteases, lipases).

However, excess water can inhibit germination by reducing oxygen availability (anaerobic
respiration is insufficient for normal germination).

(ii) Temperature
Temperature influences the rate of enzymatic reactions and membrane fluidity, governing
metabolic rates during germination. Three cardinal temperatures are defined:

Temperature Effect

Minimum (T_min) Below this, germination ceases; enzymes inactive

Optimum (T_opt) Fastest and most complete germination

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Maximum (T_max) Above this, germination inhibited; protein

denaturation

T_opt for cereals Approximately 20–30 °C

T_opt for cool-season crops 10–20 °C (wheat, barley, pea)

T_opt for warm-season crops 25–35 °C (maize, sorghum, cotton)

Some seeds require alternating temperatures (thermophotoperiodism); others require

stratification (prolonged cold, 2–5°C) for dormancy relief.

(iii) Oxygen
Germinating seeds are highly aerobic. Oxygen is essential for:

 Oxidative phosphorylation (ATP production via electron transport chain).

 Beta-oxidation of fatty acids in lipid-storing seeds.
 Biosynthetic reactions (protein synthesis, membrane biogenesis).

Some semi-aquatic plants (rice) can germinate under low oxygen using anaerobic
fermentation pathways, producing ethanol as a by-product.

(iv) Light
Seeds can be classified by their light requirements as:

 Positively photoblastic: Germination stimulated by light (red light, 660 nm).

Mediated by phytochrome (Pfr form is active). Examples: Lactuca sativa (lettuce),
Nicotiana tabacum (tobacco).
 Negatively photoblastic: Germination inhibited by light. Require darkness.
Examples: Allium cepa (onion), Cucurbita (pumpkin), Phlox.
 Neutral (light-independent): Most crop seeds—maize, wheat, sorghum, soybean.

The phytochrome system (Pr ⇌ Pfr interconversion) is the primary photoreceptor. Red
light (660 nm) converts Pr → Pfr (active); far-red light (730 nm) converts Pfr → Pr (inactive).
The ratio R:FR in the light environment thus determines germination behaviour.

(v) Soil Conditions and Depth of Sowing

Soil texture, structure, and pH influence water availability and aeration. Optimal sowing
depth varies; too deep placement leads to seedling depletion of reserves before emergence.

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Soil microbiome can either promote (mycorrhizae, PGPR) or inhibit (pathogens)

germination.

1.3.1 Biochemical Changes During Germination

A. Carbohydrate Mobilisation
In starchy seeds (cereals), the aleurone layer is activated by gibberellin (GA3) secreted by
the embryo. GA3 induces de novo synthesis of alpha-amylase and other hydrolytic
enzymes in aleurone cells. These enzymes are secreted into the endosperm where they
hydrolyse starch:

Starch (amylose/amylopectin) → Maltose → Glucose

Alpha-amylase cleaves internal α-1,4-glycosidic bonds; beta-amylase removes maltose

from non-reducing ends; limit dextrinase/pullulanase debranches α-1,6-bonds. Glucose
and sucrose are transported to the growing embryo via the scutellar epithelium. The central
role of GA in this cascade was elucidated by Paleg (1960) and later confirmed at molecular
level.

B. Protein Mobilisation
Storage proteins (vicilin, legumin in legumes; hordein in barley; gliadin/glutenin in wheat)
are hydrolysed by endopeptidases, carboxypeptidases, and aminopeptidases sequentially.
The resulting amino acids fuel protein synthesis in the growing embryo. Protease
activation is also partially under GA and ABA regulation.

C. Lipid Mobilisation
In lipid-storing seeds (castor bean, sunflower, cotton), triacylglycerols stored in oil bodies
are mobilised via:

4. Lipase activity: TAG is hydrolysed to glycerol + 3 fatty acids by seed lipases.

5. Beta-oxidation: Fatty acids are activated (acyl-CoA synthetase), transported into
glyoxysomes, and oxidised via beta-oxidation to acetyl-CoA.
6. Glyoxylate cycle: Two acetyl-CoA molecules are condensed via the glyoxylate
cycle (unique to plants and microbes) into succinate, exported to mitochondria,

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and converted to oxaloacetate → phosphoenolpyruvate → sucrose. This net

conversion of fats to sugars enables gluconeogenesis in germinating seeds—a
unique biochemical capability.

The glyoxylate cycle (isocitrate lyase + malate synthase) is active only in germinating lipid-
rich seeds and green algae. It bypasses the two carbon-loss steps of TCA cycle, allowing net
synthesis of carbohydrates from acetyl-CoA. (Bewley & Black, 1994 in Basra, 2006)

D. Respiratory Changes
Dry seeds have extremely low respiration rates (< 1 μL O2/g/h). Upon imbibition,
respiration rises exponentially:

 Phase I: Anaerobic fermentation predominates initially (low O2 in dry seed

interior).
 Phase II: Transition to aerobic respiration as oxygen diffuses in; pentose phosphate
pathway (PPP) also activated to provide NADPH for reductive biosynthesis.
 Phase III: Full aerobic TCA cycle and oxidative phosphorylation—necessary for
active growth.
 ATP generation powers membrane repair, mRNA transcription/translation, and
cell division.

E. Nucleic Acid Metabolism

Within minutes of imbibition, mRNA repair and polysomal assembly begin using stored
maternal mRNAs (long-lived mRNAs stored in dry seeds). Within hours, de novo
transcription of germination-specific genes (GA-signalling genes, amylase genes,
aquaporins) begins. The cell cycle is re-entered during Phase II; DNA replication (S-phase)
precedes the first mitotic division that marks radicle elongation.

F. Hormone Dynamics
Hormone Role during Germination

GA (Gibberellins) Induces alpha-amylase synthesis in aleurone;

promotes cell elongation; relieves dormancy

ABA (Abscisic acid) Maintains dormancy in mature seed; levels

decline upon imbibition; inhibits germination at
high levels

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Auxin (IAA) Stimulates root elongation; cell elongation in

hypocotyl

Cytokinin (CK) Promotes cell division in embryo; nutrient

mobilisation

Ethylene Can stimulate germination; relieves

thermoinhibition (e.g., lettuce); interacts with
ABA signalling

Brassinosteroids (BR) Promote seedling emergence; synergise with GA;

BR-deficient mutants have reduced germination
rates

1.4 Seed Dormancy

1.4.1 Definition and Ecological Significance

Dormancy is a state of suspended germination in a viable seed even when placed under
apparently favourable environmental conditions (adequate moisture, temperature,
oxygen, and light). It is an adaptive mechanism that prevents premature germination,
spaces seedling emergence over time (bet-hedging strategy), and ensures germination is
synchronised with favourable seasons.

Fenner and Thompson (2005, cited in Basra, 2006) distinguish between dormancy and
quiescence:

 Quiescence: Simple inactivity due to unfavourable environmental conditions;

germination resumes immediately when conditions become favourable.
 Dormancy: Active inhibition of germination imposed by the seed itself, not
relieved by merely providing external requirements.

1.4.2 Types of Dormancy

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Figure 1.5 – Classification of seed dormancy types showing physical, physiological, morphological, and secondary
dormancy pathways

A. Physical Dormancy (Coat-imposed / Hard Seed Dormancy)

Mechanism: The seed coat or fruit coat (pericarp) is impermeable to water and/or gases
due to the presence of a layer of palisade cells with heavily lignified, suberised, or
cuticularised walls. Water can only enter through a specialised structure called the "water
gap" (a lens, chalaza, or hilum-associated structure) which opens upon specific
temperature, humidity, or scarification stimuli.

 Families: Fabaceae (many legumes), Convolvulaceae, Malvaceae, Geraniaceae,

Cucurbitaceae.
 Agricultural relevance: Hard seeds in pasture legumes (Trifolium, Medicago)
cause uneven germination; scarification before sowing is required.

B. Physiological Dormancy (PD)

This is the most common and agriculturally important type of dormancy. It is mediated by
the physiological state of the embryo or endosperm, not by physical barriers.

 Non-deep PD: Most common; broken by 3–12 weeks of cold stratification (2–5°C)
or dry after-ripening. Seeds from many temperate crops.

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 Intermediate PD: Requires 3–6 months of cold stratification. Examples: Viburnum,

Taxus.
 Deep PD: Embryo cannot germinate even when excised; requires prolonged cold
stratification (3–6 months) and warm treatment. Examples: many Rosaceae genera.

Biochemical basis: The ABA/GA ratio is the central regulator. High ABA levels promote
dormancy by: (a) inhibiting alpha-amylase synthesis, (b) promoting expression of ABI3,
ABI4, ABI5 transcription factors that repress germination genes, and (c) maintaining
vacuolar pH that inactivates germination enzymes. GA opposes ABA at every level.

C. Morphological Dormancy (MD)

Seeds with morphologically underdeveloped embryos at dispersal. The embryo is
differentiated but requires further growth inside the seed before germination can occur.
Examples: Apiaceae (carrot, parsley), Ranunculaceae, Liliaceae. Treatment: warm moist
stratification to complete embryo development.

D. Morphophysiological Dormancy (MPD)

Combination of underdeveloped embryo (morphological) plus physiological inhibition.
Up to eight levels of complexity described by Baskin & Baskin (2004). Requires complex
temperature sequences (e.g., warm followed by cold stratification) for germination.
Examples: Trillium, many forest herbs.

E. Chemical Dormancy
Presence of germination-inhibiting chemicals in seed tissues, seed coat, or surrounding
fruit. Examples include:

 Abscisic acid (ABA): Endogenous inhibitor present in high concentrations in

freshly harvested seeds of many species.
 Phenolic compounds: Tannins, ferulic acid, coumarin (in Avena fatua).
 Essential oils: Aromatic compounds in some species.
 Allelopathic chemicals: In fruits of some species (Citrus, tomato)—washing seeds
removes these inhibitors.

F. Mechanical Dormancy

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Hard, rigid fruit wall or endocarp that physically resists embryo expansion. Differs from
physical dormancy in that the wall is permeable to water but mechanically too strong for
the embryo to push through. Examples: Coconut (Cocos nucifera), Walnut (Juglans regia).

G. Secondary Dormancy
Dormancy induced in non-dormant or previously dormant (now after-ripened) seeds upon
exposure to unfavourable conditions (high temperature, darkness, flooding, CO2). Of
major weed science importance—weed seeds that after-ripen and lose primary dormancy
during summer may re-enter secondary dormancy in autumn or when buried.

1.4.3 Mechanisms to Overcome Dormancy

A. After-ripening (Dry Storage)
Freshly harvested seeds of many crop species exhibit primary dormancy that is relieved
upon dry storage at low-to-moderate temperature for weeks to months. Mechanism
involves: slow oxidative reactions, lipid peroxidation, reactive oxygen species (ROS) that
act as dormancy-breaking signals, and reduction in ABA levels. Temperature-dependent;
15–35°C accelerates after-ripening, < 10°C slows it.

B. Stratification (Moist Chilling)

Seeds are placed in moist medium (sand, vermiculite, filter paper) at 2–5°C for required
duration (weeks to months). This mimics winter conditions. Biochemical changes:
reduction in ABA, increase in GA, activation of beta-1,3-glucanase (weakens endosperm
cap), and alteration of ABA:GA ratio favouring germination.

C. Scarification
Type Method Examples

Mechanical scarification Sandpaper, abrasion, knife Legume seeds, Morning glory

nicking of testa

Chemical scarification Concentrated H2SO4 (10–30 Clover, alfalfa, Ipomoea

min); NaOH; HCl

Hot water treatment Immersion in hot water (75– Acacia, Cassia

80°C) for 10–30 s then cool

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Thermal scarification Dry heat at 50–90°C for specific Fire-following species

duration

D. Light Treatments
Exposure to red light (660 nm) converts inactive Pr to active Pfr form of phytochrome,
promoting germination in positively photoblastic seeds. Continuous fluorescent light, leaf
canopy light gaps (high R:FR), and light breaks during stratification break dormancy in
many weed seeds.

E. Chemical Treatments to Break Dormancy

Chemical Mechanism Used for

Gibberellic acid (GA3) Induces alpha-amylase; Cereals, cool-season grasses

overcomes ABA inhibition

Potassium nitrate (KNO3) 0.2% Provides NO3- as N-signalling Standard ISTA germination test
molecule; activates germination additive
pathways

Thiourea (0.5–1%) Stimulates germination; reduces Apple, Prunus seeds

ABA levels

Ethylene / Ethephon Promotes germination; reduces Lettuce, celery

thermodormancy

Sodium hypochlorite Surface sterilisation + testa Weed seed studies

softening

Hydrogen peroxide (H2O2) Oxidative signals mimic after- Dormant weeds, barley
ripening

F. Temperature Treatments
 Warm stratification (20–30°C): For morphological dormancy; completes embryo
development.
 Fluctuating temperatures: Alternating 15/25°C or 20/30°C (day/night) breaks
dormancy in many weed seeds and turfgrasses by mimicking diurnal temperature
fluctuations of the soil surface.

1.5 Seed Storage and Longevity

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1.5.1 Seed Aging and Deterioration

Seed ageing (or seed deterioration) refers to the progressive decline in seed quality—
viability, vigour, germination rate, and seedling performance—over time during storage.
It is an irreversible process accelerated by high temperature and moisture content
(Copeland & McDonald, 2001).

A. Mechanisms of Seed Deterioration

(i) Lipid Peroxidation
Free radical chain reactions initiated by reactive oxygen species (ROS: O2•−, •OH, H2O2)
attack polyunsaturated fatty acids (PUFAs) in membrane phospholipids. Products of lipid
peroxidation (MDA, 4-HNE, acrolein) are cytotoxic and further damage proteins and DNA.
Malondialdehyde (MDA) levels serve as a reliable biochemical marker of seed ageing.

(ii) Membrane Integrity Loss

Lipid peroxidation and phospholipase activation alter membrane fluidity and selective
permeability. Aged seeds exhibit increased electrolyte leakage upon imbibition
(measurable by electrical conductivity tests). Loss of membrane integrity disrupts
compartmentation of metabolic activities.

(iii) Protein Damage

Carbonylation, cross-linking, and denaturation of storage and metabolic proteins
(enzymes) reduce their activity. Glutamine synthetase, alcohol dehydrogenase, and
catalase are especially sensitive markers. Maillard reactions (non-enzymatic browning)
between reducing sugars and free amino groups also damage proteins.

(iv) DNA Damage

Single-strand and double-strand breaks, base oxidation (8-oxo-deoxyguanosine), and
strand cross-links accumulate in aged seeds. These lesions must be repaired during early
germination via base excision repair (BER) and nucleotide excision repair (NER) pathways;
seeds with severe DNA damage fail to repair and die.

(v) Enzyme Inactivation

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Key enzymes of germination (alpha-amylase, lipase, glutamate dehydrogenase, catalase,

superoxide dismutase) lose activity with seed age due to oxidative damage and structural
modifications.

(vi) Reserve Depletion and Toxic Product Accumulation

Slow respiration during storage consumes seed reserves. Fermentation metabolites
(ethanol, acetaldehyde) and lipid oxidation products accumulate and are toxic to the
embryo.

Figure 1.6 – Left: Seed viability curves under different storage conditions; Right: Effect of temperature and moisture
content on seed longevity (Harrington's rule)

1.5.2 Harrington's Rule of Thumb

Proposed by Harrington (1963) and empirically validated, this rule states:

"For every 1% decrease in seed moisture content, the storage life of the seed is doubled. For
every 5°C (10°F) decrease in storage temperature, the storage life of the seed is doubled." –
Harrington (1963)

These rules apply for seed moisture between 5–14% and temperatures between 0–50°C.
Combined: a 1% reduction in MC and a 5°C reduction in temperature together quadruple
storage life.

James' Rule (the "100 rule"): The sum of % MC and storage temperature (°F) should not
exceed 100 for safe storage. E.g., at 50°F (10°C), MC should not exceed 50%—but this is a
very rough approximation; actual safe MCs for most seeds are 8–12%.

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1.5.3 Storage Conditions and Methods

A. Factors Determining Storage Life
Factor Effect on Longevity Optimal Value

Seed moisture content (MC) Most critical; exponential effect < 5–8% for orthodox

Storage temperature Highly critical; exponential < 10°C (freezing for long-term)
effect

Relative humidity (RH) Determines equilibrium MC of < 35% RH

seeds

Oxygen partial pressure High O2 accelerates oxidative Low O2 / inert atmosphere

damage

Initial seed quality Storage cannot improve quality High vigour seeds

Light Promotes oxidation and Dark storage

chlorophyll breakdown

Insect/fungal contamination Consumption and toxin Pest-free; fungicide treatment

production

B. Types of Storage
(i) Short-term / Working Storage (Active Collections)
 Temperature: 10–20°C; RH: 35–45%.
 Containers: permeable bags (cloth, paper) or ventilated bins.
 Duration: weeks to 1–2 years (for most crops).
 Used by seed companies and farmers for current season use.

(ii) Medium-term Storage

 Temperature: 4–10°C; RH: 30–40%; MC: 6–8%.
 Containers: sealed tins, poly-bags, or moisture-proof containers.
 Duration: 5–15 years.
 Used in national seed banks and seed certification agencies.

(iii) Long-term / Cryogenic Storage

 Temperature: −18°C (standard gene bank) to −196°C (liquid nitrogen).
 MC: 3–7% (must be achieved before cooling to prevent ice crystal damage).
 Containers: sealed, hermetically closed foil pouches or cryo-vials.

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 Duration: centuries (theoretically indefinite for very dry, very cold storage).
 Example: Svalbard Global Seed Vault (Norway) – deposits from >1,300 gene banks
worldwide; stores orthodox seeds at −18°C as a global failsafe backup.

(iv) Controlled Atmosphere Storage

Modification of gas composition around stored seeds:

 CO2-enriched atmosphere (60–80% CO2): inhibits insect pests and fungi; slows
oxidative reactions.
 N2 or Ar (inert gas) atmosphere: eliminates O2-dependent deterioration; used for
very high-value germplasm.
 Hermetic storage (airtight containers/cocoons): CO2 produced by seed respiration
displaces O2; practical for tropical farmers.

C. Seed Packaging Materials

Material Water Vapour Permeability Suitable For

Cloth / Gunny bags High (permeable) Short-term; cool, dry climates

Multi-wall paper bags Moderate Short-medium term; temperate

Polyethylene bags (HDPE) Low to moderate Medium term; tropical

conditions

Aluminium foil laminates Very low (0.02 g/m2/day) Long-term; gene bank storage

Metal tins (sealed) Near zero Long-term; gene banks

Cryogenic vials / ampoules Near zero Ultra-long-term; LN2 storage

D. Seed Priming as a Storage Management Tool

Seed priming involves controlled imbibition to Phase II of germination followed by re-
drying before Phase III (radicle protrusion). Major priming techniques:

 Hydropriming: Soaking in water and re-drying.

 Osmopriming (osmoconditioning): Imbibition in osmotic solutions (PEG-6000,
KNO3, KH2PO4) at defined water potential (−0.5 to −1.5 MPa) for 24–48 h at 15–
25°C.
 Hydropriming + biopriming: Inoculation with PGPR (Pseudomonas fluorescens,
Trichoderma) during priming to improve vigour and resistance.

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Priming benefits: faster and more uniform germination, improved seedling establishment
under stress, partial repair of DNA and membrane damage (improves vigour of aged
seeds).

Unit Summary: Key Concepts at a Glance

Topic Key Points

Seed Structure Testa, tegmen, embryo (plumule, radicle,

hypocotyl, cotyledon(s)), food reserve
(endosperm or cotyledons)

Seed Types Orthodox vs recalcitrant; albuminous vs

exalbuminous; mono- vs dicotyledonous

Seed Development 5 stages: post-fertilisation → histodifferentiation

→ reserve filling → maturation drying →
quiescence

Germination Types Epigeal (hypocotyl elongates; cotyledons above

soil) vs Hypogeal (epicotyl elongates; cotyledons
below soil)

Germination Factors Water, temperature, oxygen, light; internal:

viability, vigour, dormancy

Biochemical Changes Starch→sugars (amylase/GA axis); proteins→AA

(proteases); fats→sugars (glyoxylate cycle);
aerobic respiration; hormone dynamics

Dormancy Types Physical, physiological (non-

deep/intermediate/deep), morphological,
chemical, mechanical, secondary

Breaking Dormancy After-ripening, stratification, scarification

(mech/chem/heat), light, GA3, KNO3

Seed Ageing Lipid peroxidation, membrane damage, protein

carbonylation, DNA strand breaks, enzyme
inactivation

Storage (Harrington) ↓ 1% MC = 2× life; ↓ 5°C = 2× life. Optimal: < 8%

MC, < 10°C, < 35% RH

Storage Types Short-term (ambient), medium-term (4–10°C),

long-term (−18°C), cryogenic (−196°C, LN2)

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References
Copeland, L.O. & McDonald, M.B. (2001). Principles of Seed Science and Technology. 4th ed.
Springer, New York.

Egli, D.B. (2017). Seed Biology and Yield of Grain Crops. 2nd ed. CAB International,
Wallingford.

Basra, A.S. (Ed.) (2006). Handbook of Seed Science and Technology. CRC Press, Boca Raton.

Elias, S.G., Adkins, S.O. & Smith, J.P.C. (2012). Seed Testing Principles and Practices. CAB
International.

Winn, R.L. (2014). Seed Technology Laboratory Manual. 1st ed. CRC Press.

Bewley, J.D. (1997). Seed germination and dormancy. Plant Cell, 9, 1055–1066.

Baskin, C.C. & Baskin, J.M. (2014). Seeds: Ecology, Biogeography, and Evolution of
Dormancy and Germination. 2nd ed. Academic Press, San Diego.

ISTA (2021). International Rules for Seed Testing. International Seed Testing Association,
Bassersdorf, Switzerland.

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