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Imbibition and germination
Seed dormancy, Imbibition and germination
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Seed development and germination
Vivipary: No dormancy the seed continues growth. Orthodox seeds: The seed enters dormancy, dehydrated seed- tolerance to water deficiency Recalcitrant seeds: the seed enters dormancy but is not tolerant to water deficiency
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Mangrove tree- Rhizophora- vivipary
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Dormancy is an adaptive trait The consequence of natural selection
Drought and stress resistance Maximal germination under existing environmental conditions
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Dispersal in time of germination between different
seeds on the same plant in order to maximize survival a. reduced risk of mortality b. Reduced sibling competition. Dormancy break as a consequence of sensing favorable environmental conditions, seasonality, water And nutrients availability
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Seed dormancy Dormant seed: a seed that does not germinate under optimal water ; oxygen and temperature conditions Germination: Growth of embryo, breakage of seed coat, Water absorption and cell division in the embryo.
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Breaking seed coat barriers by the root
Endosperm Cersis canadensis
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Types of dormancy - Non-dormant seeds
- Primary dormancy – gradual time dependant Release from dormancy – post ripening - Secondary dormancy – seeds do not germinate under unfavorable environmental conditions - Enforced dormancy – temp. water stress
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Dormancy classification
Seed dormancy Types of dormancy Exogenous Coat imposed dormancy Mechanical dormancy: growth constraint. Physical dormancy: permeability to H2O , O2, inhibitors. Chemical dormancy: inhibitors in endosperm, seed coat. Endogenous Embryo dormancy Morphological dormancy: Under-developed embryo. Physiological dormancy : Physiological inhibition. Dormancy classification Nikolaeva (1977), Black & Bewley (1980) Baskin & Baskin (1998)
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Harsh seed coat inhibits germination. Oleaceae
Coat imposed dormancy Mechanical dormancy Harsh seed coat inhibits germination. Oleaceae (Rosaceae) (Juglandaceae)
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Morpho-physiological dormancy
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Coat imposed dormancy:
Chemical dormancy ABA Phenols (coumarin, ferulic acid) Na Cl: Atriplex , Zygophyllum , - Leaching
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water permeability barrier
3) Coat imposed dormancy: Physical dormancy water permeability barrier , Musaceae, Cannaceae.
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Bottom cavities in legumes
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Relation between air humidity
% seed water content Relation between air humidity and seed water content RH %
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Relation between air humidity
and seed water content % % RH X -hard seeds O -scarified
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Barrier to gas exchange
1. Seed coats are impermeable to oxygen the difusion coefficient of oxygen in water is very low, could be a limiting factor to germination Excess water can inhibit germination 2. Seed coats contain phenolic compounds that fix oxygen molecules
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% Effect of oxygen concentration on seed germination % O2
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Effect of Hillum plugging on germination
in melon varieties
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Effect of oxygen concentration on germination of melon seeds
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Techniques for breaking seed dormancy
1. Scarification - mechanical abrasion 2. Stratification -. Chemical treatments Treatment with concentrated sulfuric acid (15 min.) 3. Hot water or dry air
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Physiological dormancy Non-deep physiological dormancy
The dormancy can be broken by a short (days) exposure to either low or high temperatures, light or hormonal treatments GA ו- CK. - The requirements is nullified gradually in storage
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Lettuce Embryo in endospem
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Datura Tomato Lettuce lateral endosperm micropylar endosperm
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Physiological dormancy
Intermediate/deep physiological dormancy Seeds require 1-5 months of stratification to break dormancy During stratification hormonal changes will occur in the embryo axis
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Factors that determine dormancy
Factors related to seed coats Water penetration Mechanical resistance for radical break Gas exchange 2. Factors related to immature embryos Stratification by heat and cooling 3. Physiological factors related to imbibition and germination Hormonal treatments
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A classification system for seed dormancy
Baskin JM & Baskin CC A classification system for seed dormancy Seed Science Research 14:
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Seed Imbibition Phase I (imbibition)- Water uptake is a physical process driven by water potential gradient between the seed and its environment. Phase II (lag phase)- Seed water reaches a plateau level and remains constant. Metabolic processes are activated. Duration depends on Temp. Phase III (Embryo growth)- Point of no return, seed absorbs large quantities of water. Sensitive to dehydration. Embryo growth and radicle emergence.
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Imbibition Temperature
Temperature at imbibition is critical especially in large seeds. Low temp. at Imbibition cause imbibition injury. Membrane deterioration- leakage of cellular contents
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PHASE TRANSITION DIAGRAM
Phospholipid Phospholipid Free Fatty Acid
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Phospholipid Phospholipid Free Fatty Acid Gel Phase Liquid Crystalline
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Respiration and metabolic activation
Mitochondria – oxidative phosphorylation Generation of ATP prior to radicle emergence. Anaerobic respiration is possible. Protein and nucleic acid formation Repair of DNA damaged in dehydration
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Weakening of covering tissue-embryo growth
Cell division and cell expansion Cell division does not occur before radicle emergence Weakening of the cover by water and physical pressure of the radicle Cell wall hydrolases are expressed Activation is controlled by higher GA lower ABA levels.
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Reserves mobilization
Starch, other carbohydrates and lipids are converted to sucrose. Sucrose is transported to the growing axis. Proteases breakdown storage proteins into free amino acids. Phytase is activated to break down phytin and release of minerals.
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Germination, dormancy and Hormonal control
Temperature – Cardinal temperatures for dormancy break and germination Water potential – At low water potential seeds will not germinate regardless of temperatures. Light – Stimulate germination of many species. Red light (Phytochrome response) GA required for seed germination. Induces Beta mananase and other cell wall hydrolyzing enzymes. Ethylene – stimulates germination. Nutrients – Most seeds do not require nutrients for germination except for nitrates. KNO3 induces germination.
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Control of seed germination by environmental factors
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Temperature effects on the germination of melon seeds
Min. temp. Optimal temp. Max. temp. P-202 NY
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Seed categorization into temperature-requiring groups
1) Cool-temperature tolerant Range: 5-30/40oC. Optimal: 25-30oC. 2) Cool-temperature requiring Range: < 20-25oC. 3) Warm-temperature requiring Range: > 10-15oC. 4) Alternating temperatures Day/night temp. amplitude about 10oC.
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Temperature can affect germination in different ways:
1- Dry - High temperature (seed coat permeability) 2- Wet- Low temperature : chilling, stratification. 3- Temperature shifts (low => high temperature) 4- Temperature alternation (daily cycles) Temperature effects on germination interact frequently with light effects.
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Effect of stratification temperature on germination of apple seeds
85 days at low temp => to 20oC
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Duration of stratification at 5oC on seed germination
of Corylus avellana and Acer spp. Germ 5 => 200C A. pseudoplatanus אדר דולבני אדר Corylus A. platanoides ש Germ at 50C
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Effects of stratification on germination of Rumex obtusifolius at 25oC
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Chilling effects on germination temperature of
Delphinium דורבנית 6oC x 2 weeks
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Euonymus europaea
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Effect of number of temperature cycles on seed
germination of Rumex obtusifolius
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Temperature amplitude for 50% germination
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Temperature effects on germination
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Light can affect temperature in several ways
1- Light sensitive seeds (photoblastic seeds) a) positive photoblastic: light promoted germination. b) negative photoblastic: light inhibited germination. c) dual effect. 2) Light insensitive seeds (non-photoblastic seeds) - Effect of light depends on duration (sec to days), intensity and spectral quality of the light (R/FR). Phytochrome is the main photo-system involved in light control of germination.
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Effects of light on germination of Oryzopsis
(דגן-נשרן הדוחן): dual effect Short irradiation – 5 min promotion Darkness Continuous light inhibition
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Phytochrome photosystem
R m FR 730 m R FR Pfr is the promotive form of phytochrome
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a
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Dark germination - In light requiring seeds germination in the dark
depends on Pfr accumulated in the seeds during their development on the mother plant. - Decay of Pfr to Pr reduces germination in darkness.
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Effects of light and temperature on germination of Datura stramonium
Continuous light 20oC => 30oC Short light irradiation Darkness
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Escape time (escape from inhibition by FR)
R => dark period => FR % Pr Pfr Lettuce seed germination Hours between R and following FR irradiation
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Pr Pfr Red 20oC => 30oC => 20oC Darkness Dark reversion
Destruction Red 20oC => 30oC => 20oC Darkness Red light Pr Pfr Action Dark reversion (thermal process) Thermal dark reversion of Pfr explains inhibition of light promotion by high temperature.
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חסה, סלרי,(summer sowings)
Secondary dormancy seeds that after primary dormancy fail to germinate can reenter dormancy => secondary dormancy. Thermodormancy: Induced by imbibition at high temperatures (> 25oC). In contrast to thermal inhibition, will not geminate if transferred to lower temperature חסה, סלרי,(summer sowings) b) Skotodormancy: Induced by prolonged dark . c) Continuous light, anoxia
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Seed priming Controlled seed hydration treatment to reduce time to germination and seedling emergence, and improve rate and uniformity of germination. Seeds are kept in hydrated condition (priming solution) that allows metabolic processes, but prevents cell elongation and radicle emergence. After a period of time seeds are dried to original water content. Growth regulators can be added to the priming solution to promote processes.
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Processes during germination and priming
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Effects of priming on rate and uniformity of germination
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Priming requirements Seed hydration under controlled conditions that prevent germination low water potential: -0.5 to –1.5 MPa 9 (-5 to 15 bars) low temperature : oC during a few days (2-5 days) Seeds are activated metabolically but do not germinate.
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Techniques for seed priming
1) Osmotic priming (osmopriming, osmoconditioning) by imbibing seeds in osmotic solutions: salts , PEG. Then dried. washed 2) Matrix priming (matriconditioning, solid matrix priming) solid imbibed matrix: vermiculite, silica, mixed with seeds. Better for larger seeds. 3) Drum priming: hydrates seeds with measured amount of water in a tumbling drum.
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