Download Hormones 101 - WSU Tree Fruit

PLANT HORMONES 101
Greene, D.
[email protected]
University of Massachusetts, Stockbridge School of Agriculture, Amherst, MA
KEYWORDS
Plant growth regulators, Plant hormones, auxins, gibberellins, cytokinins, abscisic acid,
ethylene, directors of plant growth and development, naturally occurring compounds.
ABSTRACT
The growth and development of a plant is directed and guided by hormones produced
by the plant. Plant hormones generally do not participate directly in the growth of a
plant but they do send the signals that lead to gene expression and gene activation,
enzyme activation and/or synthesis that ultimately results in developmental changes.
There are five universally recognized major classes of hormones: auxins, gibberellins,
cytokinins, abscisic acid and ethylene. There are two hormones that have a less
dominant role in the regulation of plant growth and development: brassinosteroids and
the jasmonates.
Major Hormones
Auxins: This group of hormones was the first to be discovered in the 1930s. Indole-3acetic acid is the primary endogenous auxin found in most plants. While it is limited in
the number of field uses it is probably has the greatest influence on how trees are
grown and managed. Auxins do promote cell elongation.
Apical dominance. Auxins are produced in the apical buds of shoots and diffuse
downward inhibiting the growth and development of bud below. If the apical bud is
removed or pruned off then inhibition of growth is lifted and lateral buds start to develop.
These buds develop until they become dominate and reestablish the apical dominance.
Growth of lateral buds are allowed due to the buildup of sugars and the presence of
cytokinins.
Leaf abscission. Auxins produced in the leaf diffuse downward and prevent abscission.
As long as sufficient auxin is reaching the abscission zone abscission will not occur.
Phototropism and geotropism. Plants tend to bend and grow toward light and response
to gravity because of the movement of auxin resulting in asymmetric distribution of
auxin. Auxin mediated ethylene production.
Gibberellins (GA): This is a group of hormones that is largely responsible for stem
elongation and growth of shoots. A major goal in tree fruit production is to find cultural
and chemical means to reduce and counteract the growth-stimulating effects GAs have
on plants. There are over 135 known naturally occurring GAs. Each plant species has
just a few GAs that are particular to that plant. GA4 and GA7 are dominant GAs in apple.
Growth promotion. Limiting GA production in apple is often a goal in pome fruit
production.
Flower bud formation. GAs are known to be natural inhibitors of flower development in
pome fruit. It is largely accepted that GAs produced in the seeds play a major role in
inhibiting return bloom. GA7 is particularly inhibitory in apple.
Elongation of fruit. GAs as well as cytokinins are known to influence the shape
(elongation) of apples.
Cytokinins: This is a group of hormones that participates in many physiological
processes but their effects are less dramatic than observed for other hormones. They
are cell division factors and they help counteract stress in the plant and inhibit
senescence. Cytokinins interact with auxins in the regulation of apical dominance;
auxins reinforcing apical dominance and cytokinins help overcoming it.
Abscisic Acid: While abscisic acid does promote abscission it is far more dominant and
plays a more important role in other physiological events.
Stomatal Movement and Transpiration. ABA plays a dominant role in regulating
stomatal movement which in turn largely determines the rate of transpiration and water
loss from a plant.
Stress and Senescence. ABA is produced when plants are placed under stress. This
may lead to abscission and accelerate senescence.
Ethylene: Because ethylene is a gas its acknowledgement as an important plant
hormone was ignored for many years.
Advance fruit ripening. When fruit ripen they give off ethylene. The start of significant
ethylene production (or CO2 evolution) generally signals the physiological marker that
indicates that fruit are ripe. This ethylene then speeds the ripening process. Controlling
ethylene production and action is key to regulating ripening, fruit quality and storage
potential.
Promote Abscission. Exogenous application of ethylene or ethylene production in the
plant in response to stress will promote abscission. Controlling heat, water and cold
stress will alleviate premature leaf abscission.
Auxin induced ethylene production. Elevated rates of auxin increase ethylene
production in plants. Some responses attributed to auxins may be cause by ethylene
such as epinasty.
Less Known Plant Hormones
Brassinosteroids are required for vegetative growth and pollen tube growth, promote
cell division and cell expansion and are required for plant morphological development.
Jasmonates are best known for inducing plant defenses against injury due to insect
diseases of mechanical injury.
Hormones 101
Duane W. Greene
University of Massachusetts
What are Plant Hormones?
• Plant hormones are organic compounds,
produced by the plant that In low
concentration (10-3 M) regulate or direct
growth and development in a plant.
Plant Growth Regulators
• Plant growth regulators are very similar to
plant hormones except PGRs may or may not
be produced by the plant. Many are synthetic
or not universally found in all plants.
• Some plant hormones are also considered
PGRs (GA4+7, GA3 , ABA, ACC).
Plant Hormones
• PGRs act as signal givers or directors of
growth and development.
• In general, they are not directly involved in the
processes that they direct.
• One could look at PGRs as the first domino in
a series of event that ultimately results in a
physiological response.
• Another way of looking at this that they are the
trigger that initiates a physiological action.
Plant hormones signal to initiate a
response which is then passed on and
results in a response.
PGR
Response
Send signals that lead to gene expression,
gene activation and activation and/or
synthesis of enzymes
Major Classes of Hormones
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Auxins
Gibberellins
Cytokinins
Abscisic Acid
Ethylene
(Brassinosteroids)
(Jasmonates)
Auxins
• This is the first hormone that was isolated
and identified- 1930’s.
• Indole-3-acetic acid is the primary
endogenous auxin.
• While limited in the number of field
applications, auxin probably has the
greatest influence on how you grow and
manage your trees.
Auxins control or influence:
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Apical Dominance
Phototropism
Geotropism
Abscission: leaves, fruit and flowers
Cell elongation
Cambial activity
Root formation
Increase flowering
Auxin induced ethylene production
Apical Dominance
Apical Dominance
Leaf Abscission
Fruit Abscission
Auxins present in the king fruit
suppress the auxins coming from the
lateral fruit.
This reduction on auxin triggers the
production of enzymes that weaken
and destroy the abscission zone.
We suspect that the reduction in
auxin is carbohydrate (CHO)
mediated. The greater the CHO
deficit the more abscission.
Phototropism
Geotropism
Geotropism
Gibberellins (GA)
• Stem Elongation
• Stimulate seed germination
• Flowering
– Promotion
– Inhibition
• Stimulate pollen tube growth
• Elongates fruit
• Improves fruit finish
Stem Elongation
• Generally vigorous vegetative
growth is undesirable for tree
fruit.
• Inhibitors of GA biosynthesis
are used to retard growth and
make trees more efficient,
productive and carry higher
quality fruit.
Inhibition of Flowering
• GA inhibits flower bud formation in tree fruit.
• Seeds in developing fruit are rich sources of
GA.
• Since trees generally set more fruit than is
desirable, attempts are made to cause a
large number of fruit containing seeds to
abscise early in the season so that flower bud
formation for the following year will not be
inhibited or at least inhibited minimally.
• Inhibition may occur early.
Elongation of Fruit
Cytokinins
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Cell division and cell growth
Deferral of senescence
Apical dominance, promote lateral bud break
Cell differentiation
Nutrient mobilization
Morphogenesis
Fruit elongation
Cell Division
• Cytokinins were discovered in the late
1940s because they caused cell division.
• The cytokinins were given their name
because they did promote cytokinesis (cell
division).
Cytokinins inhibit scenesence
• Cytokinins delay and defer
senescence by regulating enzymes
that encourage cytokinin synthesis
and inhibit their degradation.
Apical Dominance
• Cytokinins interact with auxin to regulate
apical dominance.
• Sugars are also invovled.
Abscisic Acid (ABA)
• BA was discovered in large part because
it caused leaf abscission in cotton.
• However, ABA only plays a secondary role
in abscission. Both auxins and ethylene
are more prominent hormones in
regulation of abscission.
Abscisic Acid
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Controls stomatal movement
Inhibition of vegetative growth
Promotes abscission
Involved in seed and seed dormancy
Produced in stressed plants
Stomatal Movement & Transpiration
• ABA plays a pivotal role in the regulation
of stomatal movement water loss.
• In response to water stress ABA increase
in the leaves which results in cations
moving out of the guard cells esp. K+.
• The lower osmotic pressure in the guard
cell results in osmotic loss of water
causing the stomates to close.
Ethylene
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Advances fruit ripening
Promotes senescence
Enhances flowering in pome fruit
Causes epinasty
Frequently produced as result of auxin
application
• Auxin induced ethylene production
Ethylene the Hormone
• Ethylene was ignored as a legitimate plant
hormone for many years.
• Scientist could not understand how a small
gas molecule could be a hormone
• It was not until ethylene could be detected
a very low rate with a gas chromatograph
that its importance as recognized.
Ripening of Fruit
• When fruit ripen they give off large
amounts of ethylene.
• Efforts are made either inhibit ethylene
production (ReTain) or block its response
(SmartFresh, Harvista).
• Ethylene management is a key to extend
storage life.
Ethylene Promotes Abscission
• Ethylene along with auxins are key
hormones that regulate leaf abscission.
• Ethylene production is increased in
stressed plants.
• This in turn results in a decrease in auxin
content which can then lead to abscission
Auxin Induced Ethylene Production
• Auxins stimulate ethylene production.
Frequently, responses in a plant that
appear ethylene-like are actually triggered
by auxin.
• Examples:
– Epinasty of leaves following NAA application
– Leaf curling following 2,4-D application
Jasmonates
• Produced in response to wounding as a
natural defense compound.
• The can be growth retardants and retaerd
root and coleoptile growth
• Promote senescence.
• Most common are methyl jasmonate and
jasmonic acid.
Brassinosteroids
• Steroids that promote and are necessary
for normal stem elongation.
• Plants that lack this hormone are dwarfed
and this can be reversed by providing
brassinosteriods.
Summary
• Plant hormones are the organic compound
produced by the plant that regulate the
growth and development of the plant.
• They control all aspect of growth and
development of aa plant.
• They should be considered signal givers
that control the growth in an ordered
sequenced series of events.