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A Sketch of an 8 Part
Plant Hormone Theory
"Fools have no interest in understanding; they only want to air their own
opinions." Proverbs 18:2 NLT "Whatever exists has already been named..."Ecclesiastes 6:10 NIV
Informally since 1986 and on the Web since 1996,
I have written several fairly different versions of comprehensive
speculations on the functions and behavior of plant hormones. In early
2008 I was reading the Wikipedia articles on Jasmonates, and the article
made me question the role I had made for Auxin as the indicator of excess sugar. This
was because by Jasmonic Acid is involved in tuber formation and
de-chlorophylling leaves actions we might expect from Auxin. Presumably both these events occur because of an
excess of sugar. So if this is true, my new eight hormone scheme
from 2007 needs rearranging again.
Furthermore on 08/12/2008 I read an article on Brassinosteroid,
causing me to reclassify again it as working with GA as a sugar deficiency
indicator and not as a mineral deficiency one. I now leave the mineral
deficiency signal up for question.
So here's the break down:
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Deficiency
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Abundance
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Sugar
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Gibberellin/Brassinosteroid
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Jasmonic Acid (Auxin too?)
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Gases
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Ethylene
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Auxin
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Minerals
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?
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Cytokinin
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Water
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Abscisic Acid
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Salicylic Acid
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One of the problems with this scheme is the lack of a mineral deficiency
hormone candidate. Also there appears to be some question about
whether Brassinosteroid increased root growth or inhibition. If it
inhibits it short term and increase it long term, than this understandable
as just the behavior we would expect from a sugar shortage message.
Roots don't make sugar, and should be the first place to experience sugar
deficiency. The hormone may be an attempt to restart root growth on the long term. However, on the short run it might want to
change the behavior of the shoot to bring down more sugar through the stem
to the roots. It might want also minimize any increase in deficiency
the root might be experiencing, through the means of inhibiting it's root
growth.
Another problem, is seen when I explain below in the big table below that going
against common belief, I believe Jasmonic and Salicylic Acid are needed to
be present in addition to Auxin and Cytokinin for cell division. However
from my now rather ancient researches into the matter I never found such a
reference or indication...Additionally those two hormones do not seem to be involved in crown gall formation as we
would anticipate if these hormones were additional crucial indicator "green lights" for cell division.
Keeping these in mind we might postulate two other ways to organize the overall roles of the
hormones. One is to think for each of the four major nutrients hormones are released when there is
not enough of the nutrient, a different one is released when there are
growable amounts and finally a still different
hormone is released when there is too much of any nutrient. You then
might end up with the following table:
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Nutrient
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Deficiency Hormone
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Growable Amount Hormone
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Excess Hormone
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Sugar
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Gibberellin/Brassinostreroid
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Auxin
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Jasmonic Acid
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Gases
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Ethylene
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Auxin
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?
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Water
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Abscisic Acid
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Salicylic Acid
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Ethylene
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Minerals
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Ethylene
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Cytokinin
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Abscisic Acid
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A third possible scenario is to return to a very simple system I postulated
some time ago. Auxin
would be released when a root or shoot meristematic cell finds that it
contains more than enough shoot derived nutrients mainly sugar, and all
other environmental conditions are favorable for growth. Cytokinin would be made when
meristematic cells are bathed in more than enough nutrients of the sort
normally provided by the root, mainly water and minerals and all other
conditions are favorable for growth. Conversely
Gibberellin/Brassinostreroid would be made when mature cells have less than enough shoot
nutrients, i.e. sugar and Oxygen to survive especially if environmental
conditions are poor. Finally Ethylene might be
released when mature cells are receiving less than enough nutrients normally
received from the roots, mainly minerals and water, to support life at all,
thus senescence of the cell is warranted. Again this effect may be
accentuated by poor environmental conditions.
In this scheme Abscisic Acid might fulfill the role akin to adrenaline or
cortisol in animals, signaling a need emergency
action under most kinds of rapidly developing environmental stress, not just
water shortages. Complimentarily, Salicylic Acid may be the hormone
released when things are running normally and no special rapid response is
needed from the plant. It might be the "feel good" hormone.
The problem with this scheme has been pointed out that GA is made by
meristematic cells not mature ones. This is not fatal to the
speculations, but does kind of make them a little less symmetrical and
compelling.
A different table might then emerge:
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Root Derived Nutrient Abundance + Good
Environmental Conditions
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Root Derived Nutrient Deficiency + Bad
Environmental Conditions
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Shoot Derived
Nutrient Abundance + Good Environmental Conditions
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Auxin & Cytokinin - produces cell division
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1st Cytokinin then 2nd Gibberellin/BA - produces 1st root
broadening then older root cell senescence
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Shoot Derived
Nutrient Deficiency + Bad Environmental Conditions
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1st Auxin then 2nd Ethylene - produces 1st stem lengthening then
older stem cell senescence
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GA/BA & Ethylene - produces cell senescence
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One thing not discussed so far is that root oxygen is probably mostly
obtained from the soil surrounding the roots, not from the leaves.
This resolves the perplexing property of Ethylene causing the senescence of
leaves because the shoot and leaves aren't the providers of O2 for the root.
So the plant wouldn't be shooting itself in the foot if it were to trim
older inefficient leaves and stems and the resources freed could be used for
making adventitious roots under anoxia conditions.
Hormone Table - Under Construction
Note the information in the table
is constructed to defend the speculations of the first table and hasn't yet
been modified to provide the equal treat all three ways of understanding
hormones may deserve.
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Abscisic Acid
(ABA)

ABA's main role is clearly resistance
to drought conditions -
A Wikimedia Commons Image
Speculative Overall Role:
Signal of Water Deficiency

ABA May Figure Prominently in
Desert Plants Like Cactus -
A Wikimedia Picture
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Induce by drought.[5]
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Closes stomata.[6]
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Inhibits shoot growth but will not
have as much affect on roots or may even promote growth of
roots.
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Induced by salt stressed plants.
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Synthesized under heat stress.
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Induced by cold stress.
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Inhibits the uptake of Kinetin
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Down regulates enzymes needed for photosynthesis[7]
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Actions of ABA attempt to mitigate drought effects.
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Water loss is slowed by closing of the stomata.
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Shoot growth does not rectify water deficiency, however, new
root growth may.
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High salt levels may water stress plants.
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Heat stress may induce water stress.
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Cold may make water less available to plants.
Transpiration cools plants, so this might be avoided by the ABA
induced stomata closing.
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The photosynthesis process uses twice as much water as it
makes.[8]
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Synthesis: Droughted plants should have
high levels of ABA, well watered plants, low levels. Like
abundance signals ABA may be mostly made in meristematic cells
and much less so as cells mature. (Or for real theoretical
beauty, deficiency hormones should be made mostly in mature
cells and much less so in meristematic cells). ABA should
be made when a cell has less than enough water to support both
it any cell dependent on it for water acquisition. Thus
ABA is an indication that water exists in less than enough
amounts to continue the plant at its current size, thus the
plant must use emergency stores of water, find new sources of
the liquid and cut down on water sinks.
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Exogenous Treatment: High levels of
exogenously applied ABA should induce SA synthesis, because many
of ABA's effects may be to increase water levels within the
plant, if only temporarily. This may include making
dormant reactions that are normally dependent on water.
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Inhibition and Stimulation: ABA should
encourage root and new root growth, but inhibit shoot growth and
even encourage shoot and leaf senescence.
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Storage: ABA should cause the emptying of
stored water reserves found in vacuoles or tubers.
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Nutrient and Hormone Attraction and Repulsion:
ABA should generally push all nutrients and abundance
signals/hormones out of cells. ABA should attract the
deficiency signals/hormones, GA, ET and BA, leading to positive
feedback and cell senescence.
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Apical Dominance: Should break root apical
dominance because low water levels are an indication of poor
performance by the currently dominant apical root. ABA may
strengthen the currently dominant shoot apex in order not to
encourage any new shoot growth which would be a further sink on
water levels.
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Hormone Transport: Water deficiency, on
average should be detected in the leaves first, the point
furthest from the source of water. Water may be repelled
from tissues high in ABA, thus ABA may be built up in the roots,
in order to force water toward the shoots.
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Cell Division: Although it may encourage it
in the roots, if it is inducing new ones, ABA should generally
inhibit cell division, as a water deficient plant is in no
condition to expand.
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Senescence: Just as I am hypothesizing that
SA, JA, IAA and CK all need to be present to induce cell
division, ABA, GA, ET and BA may all need to be present for cell
senescence to proceed. ABA should encourage senescence,
particularly of shoot tissue whose nutrients can be cannibalized
and used to make more water absorbing root tissue.
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Specific to a Water Deficiency Signal:
Since GA causes lengthening, ET broadening, BA lengthening,
what's left is ABA should cause cell and tissue broadening when
it induces growth if it does. (I believe I may have seen such a
finding but I have to find the reference again).
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Jasmonates
(JA)

Jasmonates are similar chemically to
prostaglandins, which are chemical messengers found in animals-
A Wikimedia Commons Image
Speculative Overall Role:
Signal of Sugar Abundance

JA prompts plants to store excess sugar in tubers like the Potato -
A Wikimedia Commons Image
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"High levels of JA are also found in
flowers and pericarp tissues of developing reproductive
structures..."
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High levels of JA exist in the chloroplasts of illuminated
plants;[1]
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"JA levels also increase rapidly in response to mechanical
perturbations such as tendril coiling and when plants suffer
wounding."
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"High levels of JA encourage the accumulation of storage
proteins; genes encoding vegetative storage proteins are JA
responsive and tuberonic acid (a JA derivative) has been
proposed to play a role in the formation of tubers[4][5]"
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"JA application can induce
chlorosis and inhibition of genes encoding proteins involved
in photosynthesis, although the purpose of this response is
unknown it is proposed that this response to JA could help
reduce the plant's capacity for carbon assimilation under
conditions of excess light or carbon[1]"
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"The role of JA accumulation in flowers and fruit is
unknown; however, it may be related to fruit ripening (via
ethylene), fruit
carotenoid composition, and expression of genes encoding
seed and vegetative storage proteins[1]"
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Flowers have high amounts of sugar in order to produce
nectar.
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Chloroplast have high amounts of sugar due to
photosynthesis.
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Bruising and wounding ruptures cells and releases vacuole
sequestered sugars. Also neighboring plant cell reaction
to wounding may be to release amylases (by inducing GA?) which
increases sugar levels in the inter cell spaces.
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Storage proteins and tubers are induced by JA in order to
store excess sugar.
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Chlorosis induced by JA, is a negative feedback loop to
cannibalize excess photosynthesis machinery. JA is an
indication of excess photosynthesis capacity.
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Fruits and developing seeds may use JA to store necessary
sugar reserves in seeds and fruit.
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Synthesis: Well lighted plants should have
high JA levels, darkened plants should have low levels. JA
should be mostly made in meristematic cells and much less so as
cells mature. JA should be made when a cell has more than
enough sugar to support both it any cell dependent on it for
sugar acquisition. Thus JA is always an indication that
growth amounts of sugar exist and if conditions warrant, that
the plant has enough sugar to grow at least in the specific cell
where the JA is. (Shoot cells are responsible for acquiring
sugar for both it and similar size cells in the root whereas a
root cell is only responsible to itself for it own sugar level).
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Exogenous Treatment: Should induce GA,
because JA up regulates various processes limited by sugar
levels. Exogenously applying JA leads the plant to falsely
believe that it has high levels of sugar, thus engaging all
sorts of reactions that use sugar, thus further depleting what
may simply be a homeostatic level of existing sugar and moving
this level into the deficiency range.
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Inhibition and Stimulation: Should Induces
new root growth, just like Auxin. If Auxin is also present, JA
should inhibit shoot growth because high JA and IAA levels are
an indication of at least a short term lack of need to expand
the shoots.
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Storage: should cause sugar to be stored in
proteins and tubers for less propitious times.
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Nutrient and Hormone Attraction and Repulsion:
Should attract all nutrients and abundance signals to a
cell and repulse deficiency signals.
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Apical Dominance: Should induce shoot
apical dominance along with Auxin, however the possibility
exists for two dominant apices if one is particularly good at
sugar production (in the light) and one good at oxygen
harvesting (in the wind). May break root apical dominances
under conditions of low CK and SA.
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Hormone Transport: May be expected to
travel in the direction of the roots, away from the shoots and
particularly the shoot meristems. Regions of a cell or
tissue or plant part that contains high JA, may particularly
attract sugar and transport of sugar may follow active JA
transport down a plant.
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Cell Division: Is actually necessary for
cell division along with Auxin, Cytokinin and Salicylic
acid. If there are some plant callus lines that will
divide with only Auxin and Cytokinin present it is because these
cell lines are mutants that produce SA and JA natively, or these
other hormones are unknowingly being included with the "other"
nutrients/vitamins that are also added to calluses to get them
to divide.
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Senescence: Should protect plant tissue
from senescence, particularly root tissue.
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Specific to a Sugar Abundancy Signal:
Because ET and IAA show complimentary growth patterns with ET
broadening and IAA lengthening and the same is true for CK
(broadening) and BA (lengthening), we might expect that JA
should show a complimentary growth pattern to GA's cell
lengthening, thus JA should broaden cells and plant tissue.
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Gibberellin
(GA)

GA1 - there are many different
gibberellin molecules -
A
WIkimedia Commons Image
Speculative Overall Role: Signal of Sugar Deficiency

The so called "Foolish Seedling" a
disease first characterized in Japan which is due to too much GA -
Image from
here
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"Gibberellins also reverse the inhibition of shoot growth
and dormancy induced by ABA.[13]"
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Dissolves stored starch.
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Inhibits root growth.
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Promotes shoot lengthening especially in the dark.
Promotes new shoot growth.
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GA induces shoot growth and breaks shoot growth dormancy to
increase sugar levels.
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GA temporarily increase sugar levels by dissolving stored
starch.
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GA inhibits root growth which is counterproductive to
increased sugar levels.
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GA increases stem length in the dark, to move shoots out of
the shade or the ground, into the light and into sugar
production.
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Synthesis: Darkened plants should have high
levels of GA, well lighted plants, low levels. Like abundance
signals GA may be mostly made in meristematic cells and much
less so as cells mature. (Or for real theoretical beauty,
deficiency hormones should be made mostly in mature cells and
much less so in meristematic cells). GA should be made
when a cell has less than enough sugar to support both it any
cell dependent on it for sugar acquisition. Thus GA is an
indication that sugar exists in less than enough amounts to
continue the plant at its current size, thus the plant must use
emergency stores of starch, find new sources of the molecule and
cut down on its sinks.
-
Exogenous Treatment: High levels of
exogenously applied GA should induce JA synthesis, because many
of GA's effects may be to increase sugar levels within the
plant, if only temporarily. This may include making
dormant reactions that normally depend on sugar.
-
Inhibition and Stimulation: GA should
encourage shoot and new shoot growth, but inhibit root growth
and even encourage root senescence. This may be a
particularly apparent when ethylene levels are high and ABA and
BA levels low as this is an indication that resources need to
rerouted from the root to the shoot.
-
Storage: GA should cause the emptying of
stored sugar reserves found in vacuoles or tubers.
-
Nutrient and Hormone Attraction and Repulsion:
GA should generally push all nutrients and abundance
signals/hormones out of cells. GA should attract the
deficiency signals/hormones, ABA, ET and BA, leading to positive
feedback and cell senescence.
-
Apical Dominance: GA
should break shoot apical dominance because low sugar levels are
an indication of poor performance by the currently dominant
apical shoot. GA may strengthen the currently dominant
root apices in order not to encourage any new root growth which
would be a further sink on sugar levels.
-
Hormone Transport: Sugar deficiency, on
average should be detected in the lroots first, the point
furthest from the source of wsugar. Sugar may be repelled
from tissues high in GA, thus GA may be built up in the shoots,
in order to force sugar toward the roots.
-
Cell Division: Although it may encourage it
in the shoots, if it is inducing new ones, GA should generally
inhibit cell division, as a sugar deficient plant is in no
condition to expand.
-
Senescence: Just as I am hypothesizing that
SA, JA, IAA and CK all need to be present to induce cell
division, ABA, GA, ET and BA may all need to be present for cell
senescence to proceed. GA should encourage senescence,
particularly of root tissue whose nutrients can be cannibalized
and used to make more sugar producing shoot and leaf tissue.
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Auxin
(IAA and others)

Indol-3-ylacetic Acid , the most
common Auxin -
A Wikimedia Commons Image
Speculative Overall Role: Signal of Oxygen and Maybe Carbon Dioxide
Abundance

Crown galls are caused by
Agrobacterium tumefaciens bacteria; they produce and excrete Auxin
and Cytokinin and I argue Salicylic acid and jasmonic acid, which
interfere with normal cell division and cause largely
undifferentiated calluses of cells -
A
Wikimedia Image (Caption is also partially from Wikipedia - see
here)
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Induce new roots and also promote lateral and adventitious
root development and growth
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Made mostly in meristematic cells decreasing as cells
mature.
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Made in highest amounts shoot apices.
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Inhibits secondary buds below site of synthesis producing
apical dominance.
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Auxins decrease in light and increase where its dark.
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Induce cell lengthening.
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Induce secondary xylem differentiation.
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Auxins promote flower initiation, converting stems into
flowers.
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When Auxins are no longer produced by the growing point of a
plant, this initiates leaf abscission
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Auxin induces new root growth to compliment oxygen
abundance. Since oxygen is mostly taken in by the leaves,
abundance of it shifts growth away from the leaves to roots.
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? All the abundance signals are indications that certain
meristems are worth sending nutrients to, e.g. investing in, and
the strongest candidate in many species wins out to the
exclusion of all others. Although possibly there is one
dominant shoot apex for all four nutrient groups, water,
minerals, sugar and gases. Perhaps most of the time, the
mineral and water apex and the sugar and gas apices are the same
making two dominant apices, one for the root and one for the
shoot.
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Oxygen abundance would be most apparent in that part of the
plant producing and harvesting the most amount of oxygen, the
leaves.
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? Cuts down on excess growth of the shoot. Excess
oxygen already exists, so the mission is that it needs to be
complimented by excess sugar, minerals and water. Perhaps
producing these complimentary nutrients is not accomplished by
letting secondary shoot buds grow in an uncontrolled way, but by
a second shoot apex establishing itself as the best possible
site of sugar producing and putting all other resources into
establishing productive roots (basically your guess is as good
as mine :-).
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? Opposite what is expected if speculative overall role is
true. It is known the Auxin is made in the highest levels
in the shoot apex and this is often the most light exposed part
of the plant. Perhaps light induces an increase in Auxin
biosynthesis, due to the photosynthesis reaction producing
oxygen, but that light induces a greater active transport
away of Auxin than darkness such that real levels appear to fall
in illuminated plant parts.
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? Cell lengthening requires processes that require a lot of
oxygen? Excess oxygen is sequestered in vacuoles, blowing a cell
up like a balloon?
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? Opposite of what is expected, unless secondary xylem
remains a hollow tube for the transport of oxygen, not the up
flow of water. However, maybe xylem differentiation may be
to bring water and minerals to compliment the oxygen indicated
by Auxin and the sugar indicated by JA.
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? Perhaps flower initiation normally takes a lot of
respiration, so point of highest concentration of Auxin, the
dominant oxygen apex, is the best place to initiate a flower.
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A leaf no longer producing Auxin, may no longer be "pulling
its weight" in terms of oxygen harvesting and thus needs to be
excised.
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Synthesis: Well aerated plants should have
high IAA levels, anoxia treated plants should have low levels.
IAA should be mostly made in meristematic cells and much less so
as cells mature. IAA should be made when a cell has more
than enough oxygen to support both it any cell dependent on it
for oxygen acquisition. Thus IAA is always an indication
that growth amounts of oxygen are being procured by the plant
and if conditions warrant, that the plant has enough oxygen to
grow at least in the specific cell where the IAA is. (Shoot
cells are responsible for acquiring oxygen for both it and some
of the oxygen for a similar size cell in the root. Whereas
a root cell is only responsible to itself for it own oxygen
level and may even obtain some oxygen from spaces between soil
particles).
-
Exogenous Treatment: Should induce ET,
because IAA up regulates various processes limited by
oxygen. Exogenously applying IAA leads the plant to
falsely believe that it has high levels of oxygen, thus engaging
all sorts of reactions that use O2,
thus further depleting what may simply be a homeostatic level of
existing O2 and moving
this level into the deficiency range.
-
Inhibition and Stimulation: Should Induces
new root growth, just like JA. Especially if JA is also present,
IAA should inhibit shoot growth because high JA and IAA levels
are an indication of at least a short term lack of need to
expand the shoots.
-
Storage: should cause O2
to be stored in proteins and tubers for less
propitious times.
-
Nutrient and Hormone Attraction and Repulsion:
Should attract all nutrients and abundance
hormones/signals to a cell and repel deficiency
hormones/signals.
-
Apical Dominance: Should induce shoot
apical dominance along with JA, however the possibility exists
for two dominant apices if one is particularly good at sugar
production (in the light) and one good at oxygen harvesting (in
the wind). May break root apical dominances under
conditions of low CK and SA.
-
Hormone Transport: May be expected to
travel in the direction of the roots, away from the shoots and
particularly the shoot meristems. Regions of a cell or
tissue or plant part that contains high IAA, may particularly
attract O2 and transport
of O2 may follow active
IAA transport down a plant.
-
Cell Division: Along with Cytokinin
and JA and Salicylic acid, IAA should be necessary for
cell division. If there are some plant callus
lines that will divide with only Auxin and Cytokinin present it
is because these cell lines are mutants that produce SA and JA
natively. Alternatively these latter two hormones are
unknowingly being included with "other" nutrients/vitamins that
are also added to calluses to get them to divide.
-
Senescence: Should protect plant tissue
from senescence, particularly root tissue.
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Ethylene
(ET)

Ethylene is an extremely simple gas -
A
Wikimedia Commons Image
Speculative Overall Role: Signal of Anoxia/Oxygen Maybe Also Carbon
Dioxide Deficiency

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Broadens/thickens plant parts.
-
Inhibits leaf expansion.
-
Inhibits geotropism.
-
Ethylene is produced at a faster rate in rapidly-growing and
-dividing cells, especially in darkness.
-
Induces leaf abscission.
-
Induced by high levels of Auxin, especially in the roots.
-
Induced by flooding.
-
Produces the epinasty reaction where leaf surfaces
deliberately grow from a position perpendicular to the stem to
one which is more horizontal.
-
Induces root hair growth.
-
Induces prop roots in flooded plants.
-
Induces air spaces in roots during flooding.
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? Opposite of what might be expected as generally thin
things have more surface area per volume than thick things.
However, perhaps broadening of leaf surface area increases
Oxygen uptake, as does growing roots in a lateral direction
rather than down.
-
? Opposite of what might be expected as leaf expansion
should increase oxygen absorption.
-
? Geotropism makes a plant grow upright. Perhaps
growing in a more lateral way could increase oxygen absorption
in some circumstances, but I don't know how.
-
Anoxia is most likely to occur in the dark and in
metabolically highly active cells which may be typified by
rapidly dividing cells.
-
? Seems counterintuitive. Should instead preserve
leaves since they are the main organ of oxygen production and
harvesting.
-
Auxin induces many respiration requiring processes, thus
depleting oxygen if Auxin levels get too high.
-
Flooding deprives roots of oxygen they normally take in from
between soil particles. Well aerated soil is best for
growing. Standing water has much less oxygen than well
aerated soil.
-
Epinastic leaves act like the handles of water pumps to
actively pump up flooded water from roots so that it can be
evaporated from the leaves and the roots move away more quickly
from being water logged. Also or may be instead, leaves
parallel to the stem have a greater amount of transpiration than
leaves perpendicular to the stem because they flap up and down
in the wind, which increases evaporation rates.
-
Root hairs absorb more oxygen from the soil more readily.
Also root hairs increase absorption of minerals so we might
expect that BA or a better candidate acting as a mineral
deficiency hormone should induce this.
-
Prop roots are like snorkels. They are hollow and let
the water logged roots more easily get oxygen.
-
The air spaces induced in roots by flooding serve the same
function as above, increasing aeration of the roots.
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Synthesis: Plants experiencing anoxia
should high levels of ET, well aerated plants, low levels. Like
abundance signals ET may be mostly made in meristematic cells
and much less so as cells mature. (Or for real theoretical
beauty, deficiency hormones should be made mostly in mature
cells and much less so in meristematic cells). ET should
be made when a cell has less than enough O2
to support both it any cell dependent on it for O2
acquisition. Thus ET is an indication that O2
exists in less than enough amounts to continue the plant at its
current size, thus the plant must use emergency stores of
oxygen, find new sources of the molecule and cut down on its
sinks.
-
Exogenous Treatment: High levels of
exogenously applied ET should induce IAA synthesis, because many
of ET's effects may be to increase O2
levels within the plant, if only temporarily. This may
include making dormant reactions that normally depend on O2.
-
Inhibition and Stimulation: ET should
encourage shoot and new shoot growth, but inhibit root growth
and even encourage root senescence. This may be a
particularly apparent when GA levels are high and ABA and BA
levels low as this is an indication that resources need to
rerouted from the root to the shoot.
-
Storage: ET should cause the emptying of
stored O2 reserves found
in vacuoles or tubers if such things exist.
-
Nutrient and Hormone Attraction and Repulsion:
ET should generally push all nutrients and abundance
signals/hormones out of cells. ET should attract the
deficiency signals/hormones, ABA, GA and BA, leading to positive
feedback and cell senescence.
-
Apical Dominance: ET
should break shoot apical dominance because low O2
levels are an indication of poor performance by the currently
dominant apical shoot. ET may strengthen the currently
dominant root apices in order not to encourage any new root
growth which would be a further sink on O2
levels.
-
Hormone Transport: O2
deficiency, on average should be detected in the
roots first, the point furthest from the source of the greatest
amount of O2
(although some O2
may be absorbed from
around the roots). O2
may be repelled from tissues high in ET, thus ET may be built up
in the shoots, in order to force O2
toward the roots.
-
Cell Division: Although it may encourage it
in the shoots, if it is inducing new ones, ET should generally
inhibit cell division, as an O2
deficient plant is in no condition to expand.
-
Senescence: Just as I am hypothesizing that
SA, JA, IAA and CK all need to be present to induce cell
division, ABA, GA, ET and BA may all need to be present for cell
senescence to proceed. ET should encourage senescence,
particularly of root tissue whose nutrients can be cannibalized
and used to make more oxygen harvesting shoot and leaf tissue.
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Cytokinin
(CK)

Zeatin - the most common Cytokinin -
A Wikimedia
Commons Image
Speculative Overall Role: Signal of Mineral Abundance e.g. Nitrogen,
Phosphate, Potassium etc.

Cytokinin is synthesized in
the greatest amounts in root tips -
A Wikimedia Commons Image
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Induces new shoots.
-
Induces cell broadening.
-
Made mostly in dividing meristematic cells.
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"Inhibits plant tissue senescence, perhaps more in the
shoots than in the roots. New growth and newly-germinated
seedlings produce more ethylene than can escape the plant, which
leads to elevated amounts of ethylene, inhibiting leaf
expansion. As the new shoot is exposed to light, reactions by
photochrome in the plant's cells produce a signal for
ethylene production to decrease, allowing leaf expansion."
(Adedipe,
Hunt, & Fletcher)
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Attempts to compliment root derived mineral
abundance with shoot derived nutrients of sugars and gases.
-
? If BA is CK's compliment this follows the
other hormone compliment pairs that show complimentary growth
patterns. CK broadens, BA lengthens, IAA lengthens, ET
broadens.
-
CK is a signal that certain meristems cells
are good bets for sending resources too. CK is in effect
an indication that these cells are producing the amounts of
minerals they supposed to and enough to warrant growth.
-
Preserves shoots in order to compliment
minerals with sugar and gases. May actually inhibit root
growth and induce root senescence, just as Jasmonic Acid induces
chlorosis.
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-
Synthesis: Well fertilized
plants should have high CK levels, plants living in poor soils
should have low levels. CK should be mostly made in meristematic
cells and much less so as cells mature. CK should be made
when a cell has more than enough essential minerals to support
both it any cell dependent on it for mineral acquisition.
Thus CK is always an indication that growth amounts of minerals
are being procured by the plant and if conditions warrant, that
the plant has enough minerals to grow at least in the specific
cell where the CK is. (Root cells are responsible for acquiring
minerals for both it and similar size cells in the root.
Conversely a shoot cell is only responsible to itself for it own
mineral nutrition levels).
-
Exogenous Treatment: Should
induce BA, because CK up regulates various processes limited by
minerals. Exogenously applying CK leads the plant to
falsely believe that it has high levels of minerals, thus
engaging all sorts of reactions that use or are normally limited
by mineral levels, thus further depleting what may simply be a
homeostatic level of the existing fertilizers and moving this
level into the deficiency range.
-
Inhibition and Stimulation:
Should induces new shoot growth, just like SA. Especially if SA
is also present, CK should inhibit root growth because high SA
and CK levels are an indication of at least a short term lack of
need to expand the roots.
-
Storage: CK should cause
excess minerals
to be
stored in vacuoles, storage proteins and tubers for less
propitious times.
-
Nutrient and Hormone Attraction and
Repulsion: Should attract all nutrients and abundance
hormones/signals to a cell and repel deficiency
hormones/signals.
-
Apical Dominance: Should
induce root apical dominance along with SA, however the
possibility exists for two dominant apices if one is
particularly good at fertilizer absorption (in good soil) and
one good at water harvesting (in the moist part of the soil).
CK may break shoot apical dominances under conditions of low JA
and IAA.
-
Hormone Transport: CK may be
expected to travel in the direction of the shoots, away from the
roots and particularly away from root meristems. Regions
of a cell or tissue or plant part that contains high CK, may
particularly attract fertilizer type minerals and transport of
important minerals may follow active or passive CK transport up
a plant in the xylem or other tissue.
-
Cell Division: Along with
IAA and JA and Salicylic acid, CK should be necessary
for cell division. If there are some plant callus
lines that will divide with only Auxin and Cytokinin present it
is because these cell lines are mutants that produce SA and JA
natively. Alternatively these latter two hormones are
unknowingly being included with "other" nutrients/vitamins that
are also added to calluses to get them to divide.
-
Senescence: Should protect
plant tissue from senescence, particularly shoot tissue.
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Brassinolides
(BA)

Brassinolide was the first
Brassinosteroid discovered -
A
Wikimedia Commons Image
Speculative Overall Role:
Signal of Mineral Deficiency e.g. Nitrogen, Phosphate, Potassium
etc.

Brassinosteroids were
first discovered in Brassinus napus, Rapeseed, from which comes
Canola Oil (from a variety). Brassins are from the Mustard Family -
A
Wikimedia Commons Image
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"One well-supported hypothesis is that all tissues produce
BRs, since BR biosynthetic and signal transduction genes are
expressed in a wide range of plant organs, and short-distance
activity of the hormones also supports this.[4][5]"
-
Promote differentiation of xylem tissues
-
Inhibit leaf abscission.[14]
-
Plants found deficient in brassinolides suffer from
dwarfism.
-
Promotes cell expansion and cell elongation;[4]
works with
Auxin
to do so[6]
-
Accelerates senescence in dying tissue cultured cells;
delayed senescence in BR mutants supports that this action may
be biologically relevant[4]
-
Can provide some protection to plants during chilling and
drought stress[4]
-
BA is transported acropetally.
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-
?
-
Stimulates phloem growth in order to route
sugar to a plant part experiencing deficiency.
-
BA does this to maximize the level of sugar
coming from the leaves. Leaf abscission would cut down on some
sugar production.
-
Brassinolide encourages stem lengthening
just like Gibberellin, because the signal could be an
indication the plant's leaves are in the shade and thus the stem
needs to lengthened to move it back into the sun.
-
?
-
All the deficiency hormones should speed
senescence in those plant parts that are sinks and net users of
their respective nutrients, but should preserve those plant
parts that are harvesters or net producers of the nutrients they
represent.
-
All the deficiency hormones may do this by
causing plant cells to either senesce (if they are in bad shape)
of go into hibernation.
-
Most sugar deficiencies should occur in the
roots, so this make sense.
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Note BA actually appears to be involved in the GA pathway
whether as the principal and GA being a secondary messenger or vice
versa. From the evidence BA is like GA, actually appears be a
sugar deficiency hormone.
-
Synthesis: If BA is a sugar deficiency
signal, plants grown in the shade or with a lot of darkness, should have high
levels of BA. Alternatively well illuminated plants should
show low levels.
BA should be made when a cell has less than enough sugar to
support both it any cell dependent on it for sugar
acquisition. Thus BA is an indication that sugar levels exists
in less than enough amounts to continue the plant at its current
size, thus the plant must use emergency stores of sugar (starch), find
new sources of light and cut down on the sugar sinks, by for
instance inhibiting root growth.
-
Exogenous Treatment: If BA is truly a
sugar deficiency hormone, high levels of
exogenously applied BA should induce JA synthesis, because many
of BA's effects may be to raise sugar levels within the plant,
if only temporarily.
-
Inhibition and Stimulation: BA should
inhibit root and new root growth and even encourage root
senescence. It should encourage shoot growth, preserve
leaves from senescing and
even encourage new shoot growth initiation. This may be a
particularly apparent when ABA levels are low and GA and ET
levels high as this is an indication that resources need to
rerouted from the shoot to the root.
-
Storage: If BA is really a sugar
deficiency signal, it should cause the emptying of stored
starch reserves found in vacuoles or tubers if such things
exist.
-
Nutrient and Hormone Attraction and Repulsion:
BA should generally push all nutrients and abundance
signals/hormones out of cells. BA should attract the
deficiency signals/hormones, ABA, GA and ET, leading to positive
feedback and cell senescence.
-
Apical Dominance: BA should break shoot apical dominance because low
sugar levels are an indication of
poor performance by the currently dominant apical shoot. BA
may strengthen the currently dominant root apice in order not
to encourage any new root growth which would be a further sink
on essential sugar levels.
-
Hormone Transport: sugar
deficiency, on average should be detected in the
roots first, the point furthest from the source of mineral
harvesting,
the
shoot.
-
Cell Division: Although it may encourage it
in the shoot, if it is inducing new ones, BA should generally
inhibit cell division, as a sugar deficient plant is in no
condition to expand.
-
Senescence: Just as I am hypothesizing that
SA, JA, IAA and CK all need to be present to induce cell
division, ABA, ET , BA and GA (with BA and GA working together) may all need to be present for cell
senescence to proceed. BA should encourage senescence,
particularly of root tissue whose nutrients can be cannibalized
and used to make more shoots.
-
Specific to a Sugar Deficiency Signal:
Should induce C4 photosynthesis which is a more efficient but
more risky form of photosynthesis/sugar making. It's more
risky because it causes the buildup of poisons in the plant.
-
GA and BA may be part of the same chemical hormone cascade
pathway.
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Rai et
al., 1986, Journal of
Experimental Botany 37 : 129-134.
-
http://en.wikipedia.org/wiki/Salicylic_acid#History.
-
SMITH, Donald, L., PRITHIVIRAJ, Balakrishnan , ZHOU, Xiaomin, KHAN,
Wajahat, Salicyl Acid And Related Phenolic Compounds For Increasing
Photosynthesis In Plants, World Intellectual Property Organization
(WO/2001/026464).
-
K. Grossmann1 and T. Schm�lling, 1995, Plant Growth
Regulation 16 : 2 -
http://www.springerlink.com/content/q5000p3854l72wxk/.
-
http://en.wikipedia.org/wiki/Abscisic_Acid.
-
Zhang, J., U. Schurr, and W.J. Davies, Control of Stomatal Behaviour
by Abscisic Acid which Apparently Originates in the Roots. Journal
of Experimental Botany, 1987. 38(7): p. 1174.
-
Plant, A.R., GENE EXPRESSION REGULATED BY ABSCISIC ACID AND ITS
RELATION TO STRESS TOLERANCE. Annu. Rev. Plant Physiol. Plant Mol.
Biol, 1994. 45: p. 113-141.
-
http://en.wikipedia.org/wiki/Photosynthesis#Overview
Further Reading
References pulled from my own library
research started in the mid 1980's (and some say limited to this period and
before). Nevertheless I believe the information is relevant to today
as the earliest findings are crucial to a general theory here, not the more
detailed ones of later years.
Abeles, F. B., Holm, R. E., & Gahagan, H. E. Abscission: the role of aging. Plant Physiology 42, 1251-56, 1967.
Adedipe, N. O., Hunt, L. A., & Fletcher, R. A. Effects of Benzyladenine on Photosynthesis growth and senescence of the bean plant. Phys. Plant. 25, 151-53, 1979.
Addicott, F. T., Carns, H. R., Lyon, J. L., Smith, O. E., & McMeans, J. L. On the physiology of Abscisins. Recrulateurs Naturels de la Croissance Vegetale, pp. 687-703. Paris: C.N.R.S., 1964.
Barrington, E. J. W. Hormones. The New Encyclopaedia Britannica. Macropaedia v. 8, pp. 1074-88. Chicago: Encyclopaedia Britannica, Inc., 1975.
Beevers, L., Loveys, B., Pearson, J. A., & Wareing, P. F. Phytochrome and hormonal expansion and greening of etiolated wheat leaves. Planta 90, 286-94, 1970.
Black, M. Abscisic Acid in seed germination and dormancy. Abscisic Acid, ed. F. T. Addicott, pp. 331-364. New York: Praeger, 1983.
Booth ? Nature London 194-204
Brown, A. W., Reeve, D. R., & Crozier, A. The effect of light on the Gibberellin metabolism and growth of Phaesolus coccineus seedlings. Planta 126, 83-91, 1975.
Burg, S. P., & Burg, E. A. The interaction between
Auxin and Ethylene and its role in plant growth. PKAS 55, 262-69, 1966.
Davis & Wareing ? Planta 65 p. 129
Engelke, A. L., Hamzi, H. Q., & Skoog. F. Cytokinin-Gibberellin regulation of shoot development and leaf form in tobacco plantlets. Amer. J. of Botany 60, 491-95, 1973.
Esashi, Y., & Leopold, A. C. Plant Physiology 44, 1470, 1970.
Goeschl, J. D., Pratt, H. K., & Bonner, B. An effect of light on the production of Ethylene and the growth of the plumula portion of the etiolated pea seedling. Plant Physiology 42, 1077-80, 1967.
Goldthwaite, J. J. Further studies of hormone regulated senescence in Rumex leaf tissue. Plant Growth Substances, 1970, ed. D. J. Carr, pp. 581-88. Berlin: Springer, 1972.
Hayes, P. M., & Patrick J. W. Photosynthate transport in stems of Phaesolus vulgaris treated with Gibberellic Acid, Indole 3-Acetic Acid or Kinetin. Effects at the14site of hormone application. Planta 166: 371-79, 1985.
Hewett, E. W., & Wareing, P. F. Cytokinins in Populus x robusta Schneid: Light effects on endogenous levels. Planta 114, 119-129, 1973.
Houck, D. H., & Lamotte, C. E. Primary phloem regeneration without concomitant xylem regeneration--its hormone control in Coleus. Amer. J. Botany 64, 799-809, 1977.
Imaseki, H. Hormonal control of wound-induced responses. Encyclopedia of Plant Physiology. v. 11, ed. R. P. Pharis & D. M. Reid, p. 504, Heidelberg: Springer Verlag, 1985.
Jacobs, W. P. Comparison of the movement and vascular differentiation effects of the endogenous
Auxin and of phenoxyacetic weed killers in stems and petioles of Coleus and Phaesolus. Ann. N.Y. Acad. Sci. 144, 102-117, 1967.
Jahardhan, K. V., Vasudeva, N., & Gopel, N. H. Diurnal variation of endogenous
Auxin in arabica coffee leaves. J. Plant Crops 1 (Suppl), 93-95, 1973.
Kawase, M. Effects of flooding on Ethylene concentration in horticultural plants. J. Am. Soc. Hortic. Sci. 97, 584-88, 1972.
Lecoq, C., Koukkari, W. L., & Brenner, M. L. Rhythmic changes in Abscisic Acid (Abscisic Acid) content of soybean leaves. Plant Physiology 72 (suppl.), 52, 1983.
Manos, P.J., & Goldthwaite, J. A kinetic analysis of the effects of Gibberellic acid, Zeatin, and Abscisic Acid on leaf tissue senescence in Rumex. Plant Physiology 55, 192-98, 1975.
Marre, E. Effects of fusiccocin and hormones on plant cellmembrane activities, observations and hypothesis. Regulation of Cell Membrane Activities in Plants. eds. Marre, E. & Caffer, O. Amsterdam: Elsevier/North Holland Biomedical Press, pp. 175-202, 1977.
McMichael, B. L., & Hanny, B. W. Endogenous levels of Abscisic Acid in Water stressed cotton leaves. Agron. J. 69, 979-82, 1982.
Mitsuhashi-Kato, M., Mishibaoka, H., & Shimokoriyama, M. Anatomical and physiological aspects of developmental processes of adventitious root formation. Plant and Cell Physiology 19, 393-400, 1978.
Nooden, L. D. Senescence in the whole plant. Senescence in Plants, ed. K. V. Thimann, Boca Raton, FL: CRC Press, 1980.
Palmer, J. H., & Phillips, I. D. J. The effect of the terminal bud indole acetic acid and nitrogen supply on the growth and orientation of the petiole of the Helianthus. Annus. Physiol. Plant 16, 572-84, 1963.
Pooviah, B. W., and Leopold, A. C. Deferral of leaf senescence with calcium. Plant Physiology 52, 236-39, 1973.
??, Phys. Plant v. 51 375-79
Reid, D. M., and Wample, R. L. Water relations and plant hormones. Chapter 14 in Volume 11, Hormonal Regulation of Development III, eds. A. Pirson and M. H. Zimmerman. Heidelberg: Springer Verlag, 1985.
Reinhold, L. Phytohormones and the orientation of growth. Phytohormones and Related Compounds a Comprehensive Treatise, v. II, ed. by D. S. Letham, P. B. Goodwin, and T.J. V. Higgins. Amsterdam: Elsevier/North Holland Biomedical Press, 1978.
Ross, E. L. Growth regulators and conifers: their physiology and potential uses in forestry. Plant Growth Regulating Chemicals, v. II, ed. by L. G. Nickell. Boca Raton, FL: CRC.
Sachs, T., and Thimann, K. V. The Role of Auxin and Cytokinin in the Release of Buds from Dominance. Amer. J. Bot. 54, 126-44, 1967.
Sembdner, G., Gross, D., Liebisch, H. W., and Schneidner, G. Bio-synthesis and metabolism of plant hormones. Hormonal Regulation of Development 1, ed. J. MacMillen, Heidelberg: Springer Verlag, 1980.
Skoog, F., and Miller, W. Chemical regulation of growth and organ formation in plant tissues cultured in vitro. Symp. Soc. Exp. Biol. 11, 118, 1957.
Snow, R. Plagiotropism and correlative inhibition. New Phytologist 44, 110-117, 1945.
Sutcliffe, J. F. Regulation in the whole plant. In Transport in Plants v. 2. Part B: Tissues and Organs. Encyclopedia of Plant Physiology, ed. U. Luttge and M. G. Pitman. Heidelberg: Springer Verlag, 1976.
Thimann, K. V. Cell Enlargement and Growth. Hormone Action in the Whole Life of the Plant Amherst: Univ. of Mass. Press, 1977.
Thompson, M. J., Meudt, W. J., Mardava, N. B., Dutky, S. R.1Lusby, W. R., and Spaulding, D. W. Synthesis of Brassinosteroid and relationship of structure to plant growth promoting effect. Steroid 39 #1, 89-105, 1982.
Torrey, J. G. Auxin control of vascular pattern formation in regenerating pea root meristems grown in vitro. Amer. J. Bot. 44, 859-870, 1957.
Van Staden, J., and Smith, A. R. The synthesis of Cytokinin in excised roots of maize and tomato under aseptic conditions. Annals Bot. 42, 751-753, 1978.
Varner, J. E. GA-controlled synthesis of alpha-amylase in barley endosperm. Plant Physiology 39, 412-415, 1964.
Wain, R. L. Some development in research on plant growth inhibitors. Proc. Roy, Soc. B. 191, 335-352, 1975.
Wareing, P. F., and Phillips, I. D. J. Growth and Differentiation in Plants. Great Britain: Pergamon Press, 1981.
Webb, D. P., Van Staden, J., and Wareing, T. F. Seed dormancy in Acer. In J. Exp. Bot. 24, 105-106, 1973.
Wright, S. T. C. The effect of plant growth regulator treatments on the levels of ethylene emanating from excised turgid and wilted wheat leaves. Planta 148, 381-88, 1980.
ZeevABArt, J. A. D. Giberellin and flowering. The Biochemistry and Physiology of Giberellin, v. 2, ed. A. Crozier, New York: Praeger, 1983.
Zimmerman, P. W., and Hitchcock, A. E. Initiation and stimulation of adventitious roots caused by unsaturated hydrocarbon Gases. Contributions to the Boyce Thompson Institute 5, 351-369, 1933.
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