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Factors Affecting Seed

Germination

Mark A. Bennett
Seed Biology Program
Dept. of Horticulture and Crop Science
Ohio State University
Columbus, OH 43210-1086
bennett.18@osu.edu

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Internal Factors Affecting
Seed Germination
Seed vitality
Genotype

Seed maturation

Seed dormancy

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Germinability During Seed Development

alfalfa tomato
Developing alfalfa
seeds (A) do not
germinate when
excised from the
pod until late
development
(maturation drying)
stages
Tomato seeds (B)
take about 60 DAP From Bewley, J.D. and M. Black. 1994.
to reach full Seeds Physiology of Development and
maturity, but Germination.
germinate at 90%
levels by about 40
DAP

ABA and Seed Development


Arabidopsis:
ABA
insensitive
ABA content of seed
commonly increases
during development;
declines during late ABA deficient
maturation (drying).
For seeds developing

within fleshy fruits,


ABA content + low
water potential of fruit Wild type
tissues (osmotic
stress) both prevent
precocious
germination. From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

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Vivipary (vp) Mutants and
Seed Development
Even if wild-type
kernels of vp maize
are sprayed with
fluridone (ABA +
carotenoid synthesis
inhibitor) during
seed development,
precocious
(viviparous)
germination occurs
Mutants can be
ABA-insensitive or
From Bewley, J.D. and M. Black. 1994.
ABA deficient Seeds Physiology of Development and Germination.

Acquisition of Desiccation Tolerance

Developing
castor bean
(Ricinus
communis)
seeds removed
from pod and
placed on water
do not germinate
until about 50
DAP
If excised seeds
are first
desiccated,
From Bewley, J.D. and M. Black. 1994.
germination is Seeds Physiology of Development and Germination.
seen by 25 DAP

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Soluble Sugars and Desiccation Tolerance
In maturing seeds
of many species
conc. of sucrose
(disaccharide) &
the
oligosaccharides
raffinose,
stachyose and
verbascose (RFOs)
increase.
Monosaccharides
(glucose,
mannose, fructose,
galatose)
predominate in
seeds at the
From Bewley, J.D. and M. Black. 1994. desiccation-intol.
Seeds Physiology of Development and stage.
Germination.

Proteins & Seed Development


LEA (late
embryogenesis
abundant) proteins
occur as 2 distinct
classes; one class
increases at
midpoint of cotton
seed development
(25-35 DAP),
coincident with ABA
peaks
The second LEA

class increases at
maturation drying
(45-50 DAP)
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

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Seed Dormancy

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

Seeds are dispersed from the parent plant with different degrees of
dormancy (polymorphism, heteromorphy, or heteroblasty)
This seed variation is often seen in color, size, and coat thickness

Seed Dormancy Types

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and Germination.

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Relative Dormancy Concept

Dormancy
expression strongly
linked to
temperature
The critical temp

frequently shifts
with seed age or
various treatment
(cytokinins, nitrate,
GAs)

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and
Germination.

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From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and Germination.

Environmental Factors and


Seed Germination

Water
Temperature

Oxygen

Light

Smoke

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Water Potential Differences
Seed vs. Soil
The rate of water
transfer from soil
to seed declines
with time.
Capillary and

vapor movement
of water near
seed influenced
by soil
compaction, soil
types.

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

Seed Water Uptake


Seed-soil contact
varies with seed
size, shape.
Small seeds
(+mucilaginous
types), and smooth-
coated seeds most
efficient in H2O
uptake, increased
soil contact; larger
surface area/volume
ratio

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From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

Seed Water Uptake


Permeability of
seed to H2O is
important for
rate of uptake.
Soybean
embryos with
higher initial
moisture
contents (25%;
40%) imbibe
faster than
embryo w/ lower
water contents.

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

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Soaking Injury and Seed
Solute Leakage
With imbibition,
rapid leakage of
solutes (sugars,
organic acids,
ions, amino
acids, proteins)
into the
surrounding
medium occurs.
These solutes
may stimulate
growth of soil
microbes (fungi,
From Bewley, J.D. and M. Black. 1994.
bacteria).
Seeds Physiology of Development and
Germination.

Soaking Injury (continued)


Seeds of many legumes
(esp. Phaseolus sp., Pisum
sp., Glycine max) germinate
very poorly when hydrated
without their testas.
Some seeds (e.g. soybeans)
also leak proteinase
inhibitors and lectins upon
early imbibition may
protect against microbes,
insects.

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Temperature and Germination
Temperature
affects
capacity for
germination,
and rate of
germination.
Temperature

minimas,
maximas and
optimal ranges
vary by
species.
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

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Temperature Effects on
Dormancy, Germination
All the unchilled
seeds are dormant
at 20oC and above

The chilled
Delphinium seeds
have no dormancy;
temperature range
for germination
extends above 30oC

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and
Germination.

Temperature and Germination


Rates
Maximum rate
typically occurs
over a narrow
range (1-2oC)
This

heterogenous
response to
temperature
distributes
germination in
time

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and Germination.

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Germination Test Durations
and Temperature Optima
Highest
germination
rates around
25oC (per 2d
values) for
Brussels
sprouts.
Very good
eventual
germination
from about
10oC to 30oC,
per 14d values.
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

Oxygen and Germination


Embryo: Storage tissue:

Patterns of O2 consumption by the embryo (A)


differs late in germination from that by storage
tissues (B)
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and
Germination.

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Oxygen Availability in Soil
Found to vary from 19%
to 21% in a wide range of
conditions (time of year,
tillage, soil depth, soil
type) Sells, 1965
Exception may be when
seeds are encased in soil
particle aggregates,
sealing them off from gas
exchange with the soil
environment -(Pareja et
al., 1985)

Oxygen Solubility in Water


O2 solubility increases with decreasing
temperatures (twice as much 02
available to seeds at 5oC than at 40oC)
Literature on O2 role in seed dormancy
is sometimes conflicting, inconclusive
because of these solubility differences.

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Seed Respiration Anaerobic Conditions

Alcohol
dehydrogenase
(ADH) activity
parallels declines
in seed ethanol
levels.
Periods of natural

anaerobiosis in
germinating seeds
can last from a
few hours to
several days.
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination.

Variable Seed Coat


Permeability to Oxygen

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and Germination.

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Seed Coats also Linked to Germination Inhibitors

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and
Germination.

From Kigel, J. and G. Galili. 1995. Seed Development and Germination.

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O2 Concentrations and Tomato Seed
Germination

From Kigel, J. and G. Galili. 1995.


Seed Development and Germination.

Temperature Effects on O2
Requirements

From Kigel, J. and G.


Galili. 1995. Seed
Development and
Germination.

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Effects of Light on Seed
Germination
Light can promote or inhibit
germination
Sensitivity to light is important to seed
banks and other ecological responses,
providing a mechanism for optimal
timing of seedling establishment
The photoreceptor for most types of
seed responses is phytochrome

Species that Produce


Photosensitive Seeds

From Black, Bewley and Halmer, 2006. The Encyclopedia of Seeds

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Light and Germination
Levels of photodormancy vary not only
among genotypes, but also among
seed lots of the same cultivar (lettuce-
Wurr et al., 1986)

Photo conversions of Pr to Pfr usu. take


place when seed is hydrated above 8%

Critical periods for light environment


effects on seed germination or
storability occurred after physiological
maturity (PM) in lettuce (Contreras et
al., 2008

Lettuce - Normal
Seedlings after AA

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White light also inhibits
germination of some species
Illumination over many hours of germination
period generally required
Blue region of light spectrum (thru
cryptochrome) likely responsible
This type of photo inhibition is magnified for
seeds under water stress
Tomato seed studies -> ABA biosynthesis;
blue light increases sensitivity to ABA.
(M. Black, 2006)

Influences of smoke on
germination
Stimulatory effects of smoke on >100 Fynbos (heathland)
species in South Africa, Australia, N. America
Butenolide (3-methyl-2H-furo[2,3-c] pyran-2-one)
conforms to the necessary attributes of smoke from fires in
natural settings:
1) stable at high temps (melting pt. is ~119oC)
2) water-soluble
3) active at wide range of concentrations (1 ppm to ppt)
4) capable of promoting germination of a wide range of fire-
following species [Flematti, et al 2004 Science vol. 305]

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Chemical promotion of germination endogenous hormones

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and Germination.

Composition and location


of Auxin
Major auxin in
developing seeds is
IAA indole-3-
acetic acid
IAA is most
abundant in young
plant tissues
(immature seeds,
fruits)
IAA commonly
conjugated to
sugars, sugar
alcohols, and amino
acids (asp, glu, ala)
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination

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Gibberellins (GAs)
First GA identified (GA3 gibberellic acid) was
isolated from fungus Gibberella fujikuroi in the
1930s
The first plant gibberellin (GA1) was isolated in
1957; since then over 125 naturally occurring GA-
like compounds have been isolated from higher
and lower plants (mosses, algae)
GAs play a key role in degradation of storage
reserves -> e.g. barley aleurone system

GAs (continued)
Studies with mutants of Arabidopsis and tomato
basis for hormone balance theory; germination
timing coordinated by action of GAs and ABA
Some types of seed dormancy also linked to
changes in GA, ABA contents or sensitivity
Germination of photoblastic seeds (light
promoted or inhibited) such as lettuce,
Arabidopsis also linked to red light effect on
synthesis of GA

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Abscisic acid - ABA
Found in seeds of
many species;
promotes maturation
and dormancy,
synthesis of certain
storage proteins, and
inhibits germination
ABA generally rises in
concentration during
seed development,
reaches 1 or 2 peaks,
then (usually)
declines rapidly at the
time of seed
drydown.

From Bewley, J.D. and M. Black. 1994.


Seeds Physiology of Development and
Germination

Cytokinins (CK)
CKs have
diverse effects
on plants;
notable in
promoting cell
division
Synthesized 1 in
o
root tips and
transported thru
xylem to aerial
plant parts
CKs also occur
in developing
embryos (e.g.
zeatin from
maize kernels)
From Bewley, J.D. and M. Black. 1994.
Seeds Physiology of Development and Germination

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Ethylene (C2H4) and
germination
Eth is made in most plant tissues; also induced by
various stresses (e.g. wounding, drought, flooding)
Eth may be induced by auxin, which can be further
enhanced by CKs or ABA; eth may also self-
regulate synthesis (+ or -)
Eth is biosynthesized from ACC (a-
aminocyclopropane-1-carboxylic acid), which is
synthesized from SAM (S-adenosyl-L-methionine)
Exogenous Eth-generating compounds (e.g.
ethephon) can be used in seed enhancements
(Encycl of Seeds; 2006)

From Bewley, J.D. and M. Black. 1994. Seeds Physiology of Development and Germination

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Hydrogen peroxide (H2O2) and
germination
Early literature H2O2 as a fungicidal seed
treatment stimulated germination of certain
seeds
Mechanism(s) unknown; H2O2 may
accelerate early respiration phase of
germination?
H2O2 test for seed analysts results in more
rapid root protrusion from seed (SCST/Seed
Technol. Training manual; pgs. 10:14-15)

Other chemical promoters of


germination
Potassium nitrate (KNO3) commonly
used in germ testing; linked to relieving
light dormancy by increasing sensitivity
to light in many species
Respiratory, other metabolic inhibitors
(e.g. cyanide)
Anaesthetics (e.g. chloroform, acetone,
ethanol)

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Other chemicals that stimulate
germ/break dormancy

From Black, Bewley and Halmer, 2006. The Encyclopedia of Seeds

Chemicals which inhibit


germination

From Black, Bewley and Halmer, 2006. The Encyclopedia of Seeds

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