Germination

Germination is the process by which an

organism grows from a seed or similar
structure. The term is applied to the
sprouting of a seedling from a seed of an
angiosperm or gymnosperm, the growth
of a sporeling from a spore, such as the
spores of fungi, ferns, bacteria, and the
growth of the pollen tube from the pollen
grain of a seed plant.
Sunflower seedlings, three days after germination.

Sunflower time lapse with soil cross section,

showing how the roots and the upper part of the
plant grow
Seed plants

A seed tray used in horticulture for sowing and

taking plant cuttings and growing plugs

Germination glass (glass sprouter jar) with a plastic

sieve-lid
Brassica campestris germinating seeds

Germination is usually
the growth of a plant
contained within a seed;
it results in the Play media
formation of the
seedling. It is also the process of
reactivation of metabolic machinery of
the seed resulting in the emergence of
radicle and plumule. The seed of a
vascular plant is a small package
produced in a fruit or cone after the
union of male and female reproductive
cells. All fully developed seeds contain
an embryo and, in most plant species
some store of food reserves, wrapped in
a seed coat. Some plants produce
varying numbers of seeds that lack
embryos; these are empty seeds which
never germinate. Dormant seeds are
viable seeds that do not germinate
because they require specific internal or
environmental stimuli to resume growth.
Under proper conditions, the seed begins
to germinate and the embryo resumes
growth, developing into a seedling.

-Step 1- Water imbibition, the uptake of water,

results in rupture of seed coat.
 p
-Step 2-The imbibition of the seed coat results in
emergence of the radicle (1) and the plumule(2), the
cotyledons get unfolded(3).
-Step 3-This marks the final step in the germination

of the seed where the cotyledons are expanded

which are the true leaves/peasNote- Temperature
must be kept at an optimum level.

Disturbance of soil can result in vigorous

plant growth by exposing seeds already
in the soil to changes in environmental
factors where germination may have
previously been inhibited by depth of the
seeds or soil that was too compact. This
is often observed at grave sites after a
burial.[1]
Seed germination depends on both
internal and external conditions. The
most important external factors include
right temperature, water, oxygen or air
and sometimes light or darkness.[2]
Various plants require different variables
for successful seed germination. Often
this depends on the individual seed
variety and is closely linked to the
ecological conditions of a plant's natural
habitat. For some seeds, their future
germination response is affected by
environmental conditions during seed
formation; most often these responses
are types of seed dormancy.
Water is required for germination.
Mature seeds are often extremely dry
and need to take in significant
amounts of water, relative to the dry
weight of the seed, before cellular
metabolism and growth can resume.
Most seeds need enough water to
moisten the seeds but not enough to
soak them. The uptake of water by
seeds is called imbibition, which leads
to the swelling and the breaking of the
seed coat. When seeds are formed,
most plants store a food reserve with
the seed, such as starch, proteins, or
oils. This food reserve provides
nourishment to the growing embryo.
When the seed imbibes water,
hydrolytic enzymes are activated which
break down these stored food
resources into metabolically useful
chemicals.[2] After the seedling
emerges from the seed coat and starts
growing roots and leaves, the
seedling's food reserves are typically
exhausted; at this point photosynthesis
provides the energy needed for
continued growth and the seedling
now requires a continuous supply of
water, nutrients, and light.
Oxygen is required by the germinating
seed for metabolism.[3] Oxygen is used
in aerobic respiration, the main source
of the seedling's energy until it grows
leaves.[2] Oxygen is an atmospheric
gas that is found in soil pore spaces; if
a seed is buried too deeply within the
soil or the soil is waterlogged, the seed
can be oxygen starved. Some seeds
have impermeable seed coats that
prevent oxygen from entering the seed,
causing a type of physical dormancy
which is broken when the seed coat is
worn away enough to allow gas
exchange and water uptake from the
environment.
Temperature affects cellular metabolic
and growth rates. Seeds from different
species and even seeds from the same
plant germinate over a wide range of
temperatures. Seeds often have a
temperature range within which they
will germinate, and they will not do so
above or below this range. Many seeds
germinate at temperatures slightly
above 60-75 F (16-24 C) [room-
temperature in centrally heated
houses], while others germinate just
above freezing and others germinate
only in response to alternations in
temperature between warm and cool.
Some seeds germinate when the soil is
cool 28-40 F (-2 - 4 C), and some when
the soil is warm 76-90 F (24-32 C).
Some seeds require exposure to cold
temperatures (vernalization) to break
dormancy. Some seeds in a dormant
state will not germinate even if
conditions are favorable. Seeds that
are dependent on temperature to end
dormancy have a type of physiological
dormancy. For example, seeds
requiring the cold of winter are
inhibited from germinating until they
take in water in the fall and experience
cooler temperatures. Cold stratification
is a process that induces the
dormancy breaking prior to light
emission that promotes germination
.[4] Four degrees Celsius is cool enough
to end dormancy for most cool
dormant seeds, but some groups,
especially within the family
Ranunculaceae and others, need
conditions cooler than -5 C. Some
seeds will only germinate after hot
 temperatures during a forest fire which
cracks their seed coats; this is a type
of physical dormancy.

Most common annual vegetables have

optimal germination temperatures
between 75-90 F (24-32 C), though many
species (e.g. radishes or spinach) can
germinate at significantly lower
temperatures, as low as 40 F (4 C), thus
allowing them to be grown from seeds in
cooler climates. Suboptimal
temperatures lead to lower success rates
and longer germination periods.

Light or darkness can be an

environmental trigger for germination
and is a type of physiological
 dormancy. Most seeds are not
affected by light or darkness, but many
seeds, including species found in
forest settings, will not germinate until
an opening in the canopy allows
sufficient light for growth of the
seedling.[2]

Scarification mimics natural processes

that weaken the seed coat before
germination. In nature, some seeds
require particular conditions to
germinate, such as the heat of a fire (e.g.,
many Australian native plants), or
soaking in a body of water for a long
period of time. Others need to be passed
through an animal's digestive tract to
weaken the seed coat enough to allow
the seedling to emerge.[2]

Malted (germinated) barley grains

Dormancy …

Some live seeds are dormant and need

more time, and/or need to be subjected
to specific environmental conditions
before they will germinate. Seed
dormancy can originate in different parts
of the seed, for example, within the
embryo; in other cases the seed coat is
involved. Dormancy breaking often
involves changes in membranes, initiated
by dormancy-breaking signals. This
generally occurs only within hydrated
seeds.[5] Factors affecting seed
dormancy include the presence of certain
plant hormones, notably abscisic acid,
which inhibits germination, and
gibberellin, which ends seed dormancy.
In brewing, barley seeds are treated with
gibberellin to ensure uniform seed
germination for the production of barley
malt.[2]

Seedling establishment …
In some definitions, the appearance of
the radicle marks the end of germination
and the beginning of "establishment", a
period that utilizes the food reserves
stored in the seed. Germination and
establishment as an independent
organism are critical phases in the life of
a plant when they are the most
vulnerable to injury, disease, and water
stress.[2] The germination index can be
used as an indicator of phytotoxicity in
soils. The mortality between dispersal of
seeds and completion of establishment
can be so high that many species have
adapted to produce large numbers of
seeds.
Germination rate and
germination capacity

Germination of seedlings raised from seeds of

eucalyptus after 3 days of sowing.

In agriculture and gardening, the

germination rate describes how many
seeds of a particular plant species,
variety or seedlot are likely to germinate
over a given period. It is a measure of
germination time course and is usually
expressed as a percentage, e.g., an 85%
germination rate indicates that about 85
out of 100 seeds will probably germinate
under proper conditions over the
germination period given. Seed
germination rate is determined by the
seed genetic composition, morphological
features and environmental factors. The
germination rate is useful for calculating
the number of seeds needed for a given
area or desired number of plants. For
seed physiologists and seed scientists
"germination rate" is the reciprocal of
time taken for the process of
germination to complete starting from
time of sowing. On the other hand, the
number of seed able to complete
germination in a population (i.e. seed lot)
is referred as germination capacity.
Repair of DNA damage …

Seed quality deteriorates with age, and

this is associated with accumulation of
genome damage.[6] During germination,
repair processes are activated to deal
with accumulated DNA damage.[7] In
particular, single- and double-strand
breaks in DNA can be repaired.[8] The
DNA damage checkpoint kinase ATM has
a major role in integrating progression
through germination with repair
responses to the DNA damages
accumulated by the aged seed.[9]

Dicot germination …
The stages of germination of a pea plant. A.Seed
Coat B. Radicle C. Primary Root D. Secondary Root
E. Cotyledon F. Plumule G. Leaf H. Tap Root

The part of the plant that first emerges

from the seed is the embryonic root,
termed the radicle or primary root. It
allows the seedling to become anchored
in the ground and start absorbing water.
After the root absorbs water, an
embryonic shoot emerges from the seed.
This shoot comprises three main parts:
the cotyledons (seed leaves), the section
of shoot below the cotyledons
(hypocotyl), and the section of shoot
above the cotyledons (epicotyl). The way
the shoot emerges differs among plant
groups.[2]

Epigeal …

In epigeal germination (or epigeous

germination), the hypocotyl elongates
and forms a hook, pulling rather than
pushing the cotyledons and apical
meristem through the soil. Once it
reaches the surface, it straightens and
pulls the cotyledons and shoot tip of the
growing seedlings into the air. Beans,
tamarind and papaya are examples of
plants that germinate this way.[2]

Hypogeal …
Germination can also be done by
hypogeal germination (or hypogeous
germination), where the epicotyl
elongates and forms the hook. In this
type of germination, the cotyledons stay
underground where they eventually
decompose. Peas, gram and mango, for
example, germinate this way.[10]

Monocot germination …

In monocot seeds, the embryo's radicle

and cotyledon are covered by a
coleorhiza and coleoptile, respectively.
The coleorhiza is the first part to grow
out of the seed, followed by the radicle.
The coleoptile is then pushed up through
the ground until it reaches the surface.
There, it stops elongating and the first
leaves emerge.[2]

Precocious germination …

When a seed germinates without

undergoing all four stages of seed
development, i.e., globular, heart shape,
torpedo shape, and cotyledonary stage, it
is known as precocious germination.

Pollen germination
Another germination event during the life
cycle of gymnosperms and flowering
plants is the germination of a pollen grain
after pollination. Like seeds, pollen grains
are severely dehydrated before being
released to facilitate their dispersal from
one plant to another. They consist of a
protective coat containing several cells
(up to 8 in gymnosperms, 2–3 in
flowering plants). One of these cells is a
tube cell. Once the pollen grain lands on
the stigma of a receptive flower (or a
female cone in gymnosperms), it takes
up water and germinates. Pollen
germination is facilitated by hydration on
the stigma, as well as by the structure
and physiology of the stigma and style.[2]
Pollen can also be induced to germinate
in vitro (in a petri dish or test tube).[11][12]
During germination, the tube cell
elongates into a pollen tube. In the
flower, the pollen tube then grows
towards the ovule where it discharges
the sperm produced in the pollen grain
for fertilization. The germinated pollen
grain with its two sperm cells is the
mature male microgametophyte of these
plants.[2]

Self-incompatibility …

Since most plants carry both male and

female reproductive organs in their
flowers, there is a high risk of self-
pollination and thus inbreeding. Some
plants use the control of pollen
germination as a way to prevent this self-
pollination. Germination and growth of
the pollen tube involve molecular
signaling between stigma and pollen. In
self-incompatibility in plants, the stigma
of certain plants can molecularly
recognize pollen from the same plant
and prevent it from germinating.[13]

Spore germination
Germination can also refer to the
emergence of cells from resting spores
and the growth of sporeling hyphae or
thalli from spores in fungi, algae and
some plants.
Conidia are asexual reproductive
(reproduction without the fusing of
gametes) spores of fungi which
germinate under specific conditions. A
variety of cells can be formed from the
germinating conidia. The most common
are germ tubes which grow and develop
into hyphae. The initial formation and
subsequent elongation of the germ tube
in the fugus Aspergillus niger has been
captured in 3D using holotomography
microscopy. Another type of cell is a
conidial anastomosis tube (CAT); these
differ from germ tubes in that they are
thinner, shorter, lack branches, exhibit
determinate growth and home toward
each other. Each cell is of a tubular
shape, but the conidial anastomosis tube
forms a bridge that allows fusion
between conidia.[14][15]

3D-visualization of Aspergillus niger spore

germination. This image has been captured using
holotomography microscopy

Resting spores …

In resting spores, germination involves

cracking the thick cell wall of the
dormant spore. For example, in
zygomycetes the thick-walled
zygosporangium cracks open and the
zygospore inside gives rise to the
emerging sporangiophore. In slime
molds, germination refers to the
emergence of amoeboid cells from the
hardened spore. After cracking the spore
coat, further development involves cell
division, but not necessarily the
development of a multicellular organism
(for example in the free-living amoebas
of slime molds).[2]

Ferns and mosses …

In plants such as bryophytes, ferns, and a

few others, spores germinate into
independent gametophytes. In the
bryophytes (e.g., mosses and liverworts),
spores germinate into protonemata,
similar to fungal hyphae, from which the
gametophyte grows. In ferns, the
gametophytes are small, heart-shaped
prothalli that can often be found
underneath a spore-shedding adult
plant.[2]

Bacteria …

Bacterial spores can be exospores or

endospores which are dormant
structures produced by a number of
different bacteria. They have no or very
low metabolic activity and are formed in
response to adverse environmental
conditions.[16] They allow survival and
are not a form of reproduction.[17] Under
suitable conditions the spore germinates
to produce a vital bacterium. Endospores
are formed inside the mother cell, and,
exospores are formed at the end of the
mother cell as a bud.[18]

Light-stimulated
germination
As mentioned earlier, light can be an
environmental factor that stimulates the
germination process. The seed needs to
be able to determine when is the perfect
time to germinate and they do that by
sensing environmental cues. Once
germination starts, the stored nutrients
that have accumulated during maturation
start to be digested which then supports
cell expansion and overall growth.[19]
Within light-stimulated germination,
Phytochrome B (PHYB) is the
photoreceptor that is responsible for the
beginning stages of germination. When
red light is present, PHYB is converted to
its active form and moves from the
cytoplasm to the nucleus where it
upregulates the degradation of PIF1.
PIF1, phytochrome-interaction-factor-1,
negatively regulates germination by
increasing the expression of proteins
that repress the synthesis of gibberellin
(GA), a major hormone in the germination
process.[20] Another factor that promotes
germination is HFR1 which accumulates
in light in some way and forms inactive
heterodimers with PIF1.[21]

Although the exact mechanism is not

known, nitric oxide (NO) plays a role in
this pathway as well. NO is thought to
repress PIF1 gene expression and
stabilizes HFR1 in some way to support
the start of germination.[19] Bethke et all
(2006) exposed dormant Arabidopsis
seeds to NO gas and within the next 4
days, 90% of the seeds broke dormancy
and germinated. The authors also looked
at how NO and GA effects the
vacuolation process of aleurone cells
that allow the movement of nutrients to
be digested. A NO mutant resulted in
inhibition of vacuolation but when GA
was later added the process was active
again leading to the belief that NO is
prior to GA in the pathway. NO may also
lead to the decrease in sensitivity of
Abscisic acid (ABA), a plant hormone
largely responsible for seed dormancy.[22]
The balance between GA and ABA is
important. When ABA levels are higher
than GA then that leads to dormant
seeds and when GA levels are higher,
seeds germinate.[23] The switch between
seed dormancy and germination needs
to occur at a time when the seed has the
best chances of surviving and an
important cue that begins the process of
seed germination and overall plant
growth is light.

See also
Lily seed germination types
Oldest viable seed
Pot farm
Seedling
Seed tray
Sprouting
Urban horticulture
Pyrophyte for germination after fire.
 Vivipary when seeds or embryos begin
to develop inside or before they detach
from the parent.

References
1. Forensic Botany. Wiley-Blackwell.
2012. p. 10.
2. Raven PH, Evert RF, Eichhorn SE
(2005). Biology of Plants (7th ed.).
New York: W.H. Freeman and
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ISBN 978-0-7167-1007-3.
3. Siegel SM, Rosen LA (1962). "Effects
of Reduced Oxygen Tension on
Germination and Seedling Growth".
Physiologia Plantarum. 15 (3): 437–
 444. doi:10.1111/j.1399-
3054.1962.tb08047.x .
4. Baskin CC, Baskin JM (2014).
Variation in Seed Dormancy and
Germination within and between
Individuals and Populations of a
Species. Seeds: Ecology,
Biogeography, and, Evolution of
Dormancy and Germination.
Burlington: Elsevier Science. pp. 5–
35. ISBN 9780124166837.
5. Bewley JD, Black M, Halmer P
(2006). The encyclopedia of seeds:
science, technology and uses Cabi
Series . p. 203. ISBN 978-0-85199-
723-0.
6. Waterworth WM, Bray CM, West CE
(June 2015). "The importance of
safeguarding genome integrity in
germination and seed longevity" .
Journal of Experimental Botany. 66
(12): 3549–58.
doi:10.1093/jxb/erv080 .
PMID 25750428 .
7. Koppen G, Verschaeve L (2001). "The
alkaline single-cell gel
electrophoresis/comet assay: a way
to study DNA repair in radicle cells of
germinating Vicia faba". Folia
Biologica. 47 (2): 50–4.
PMID 11321247 .
8. Waterworth WM, Masnavi G,
Bhardwaj RM, Jiang Q, Bray CM,
West CE (September 2010). "A plant
DNA ligase is an important
determinant of seed longevity" . The
Plant Journal. 63 (5): 848–60.
doi:10.1111/j.1365-
313X.2010.04285.x .
PMID 20584150 .
9. Waterworth WM, Footitt S, Bray CM,
Finch-Savage WE, West CE (August
2016). "DNA damage checkpoint
kinase ATM regulates germination
and maintains genome stability in
seeds" . Proceedings of the National
Academy of Sciences of the United
States of America. 113 (34): 9647–
 52. doi:10.1073/pnas.1608829113 .
PMC 5003248 . PMID 27503884 .
10. Sadhu MK (1989). Plant
propagation . New Age International.
p. 61. ISBN 978-81-224-0065-6.
11. Martin FW (June 1972). "In vitro
measurement of pollen tube growth
inhibition" . Plant Physiology. 49 (6):
924–5. doi:10.1104/pp.49.6.924 .
PMC 366081 . PMID 16658085 .
12. Pfahler PL (January 1981). "In vitro
germination characteristics of maize
pollen to detect biological activity of
environmental pollutants" .
Environmental Health Perspectives.
37: 125–32. doi:10.2307/3429260 .
 JSTOR 3429260 . PMC 1568653 .
PMID 7460877 .
13. Takayama S, Isogai A (2005). "Self-
incompatibility in plants". Annual
Review of Plant Biology. 56 (1): 467–
89.
doi:10.1146/annurev.arplant.56.0326
04.144249 . PMID 15862104 .
S2CID 1196223 .
14. Roca MG, Davide LC, Davide LM,
Mendes-Costa MC, Schwan RF,
Wheals AE (November 2004).
"Conidial anastomosis fusion
between Colletotrichum species".
Mycological Research. 108 (Pt 11):
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 CiteSeerX 10.1.1.463.3369 .
doi:10.1017/S0953756204000838 .
PMID 15587065 .
15. Roca MG, Arlt J, Jeffree CE, Read ND
(May 2005). "Cell biology of conidial
anastomosis tubes in Neurospora
crassa" . Eukaryotic Cell. 4 (5): 911–
9. doi:10.1128/EC.4.5.911-
919.2005 . PMC 1140100 .
PMID 15879525 .
16. J.-M. Ghuysen; R. Hakenbeck (9
February 1994). Bacterial Cell Wall .
Elsevier. pp. 167–. ISBN 978-0-08-
086087-9.
17. Eldra Solomon; Linda Berg; Diana W.
Martin (15 September 2010).
 Biology . Cengage Learning.
pp. 554–. ISBN 978-0-538-74125-5.
18. Encyclopedia Britannica (2002).
Encyclopedia britannica .
Encyclopedia Britannica. p. 580.
ISBN 978-0-85229-787-2.
19. Penfield S (September 2017). "Seed
dormancy and germination" . Current
Biology. 27 (17): R874–R878.
doi:10.1016/j.cub.2017.05.050 .
PMID 28898656 .
20. de Wit M, Galvão VC, Fankhauser C
(April 2016). "Light-Mediated
Hormonal Regulation of Plant
Growth and Development". Annual
Review of Plant Biology. 67: 513–37.
 doi:10.1146/annurev-arplant-
043015-112252 . PMID 26905653 .
21. Li R, Jia Y, Yu L, Yang W, Chen Z,
Chen H, Hu X (February 2018). "Nitric
oxide promotes light-initiated seed
germination by repressing PIF1
expression and stabilizing HFR1".
Plant Physiology and Biochemistry.
123: 204–212.
doi:10.1016/j.plaphy.2017.11.012 .
PMID 29248678 .
22. Bethke PC, Libourel IG, Aoyama N,
Chung YY, Still DW, Jones RL (March
2007). "The Arabidopsis aleurone
layer responds to nitric oxide,
gibberellin, and abscisic acid and is
 sufficient and necessary for seed
dormancy" . Plant Physiology. 143
(3): 1173–88.
doi:10.1104/pp.106.093435 .
PMC 1820924 . PMID 17220360 .
23. Shu K, Meng YJ, Shuai HW, Liu WG,
Du JB, Liu J, Yang WY (November
2015). "Dormancy and germination:
How does the crop seed decide?".
Plant Biology. 17 (6): 1104–12.
doi:10.1111/plb.12356 .
PMID 26095078 .

Further reading
Rajjou L, Duval M, Gallardo K, Catusse J,
Bally J, Job C, Job D (2012). "Seed
germination and vigor". Annual Review of
 Plant Biology. 63: 507–33.
doi:10.1146/annurev-arplant-042811-
105550 . PMID 22136565 .
Deno NC (1980). Seed Germination: Theory
and Practice. State College, PA.
OCLC 918148836 . "An extensive study of
the germination rates of a huge variety of
seeds under different experimental
conditions, including temperature variation
and chemical environment"

External links

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Germination
 Wikisource has the text of the 1906
New International Encyclopedia
article "Germination".

Sowing Seeds A survey of seed

sowing techniques.
Germination time-lapse ≈1 minute HD
video of mung bean seeds germinating
over 10 days. Hosted on YouTube.

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