Newest Version of General Speculations

on Plant Hormones - 8 Hormone System

"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
By Paul D. Pruitt
M.A. University of Pennsylvania, Biology, 1986
Introduction
The 8 most widely studied plant hormones are Auxin (usually IAA), Cytokinin (CK), Ethylene
(ET), Gibberellin (GA), Abscisic Acid (ABA) and the lesser studied, Brassinosteroid (BA),
Salicylic Acid (SA), and Jasmonic Acid (JA).
If you look through the previous versions of this theory from the links above, they seem to
suggest some contradictory things but overall the suggestions have in common that there are in
general two classes of hormones, one which is made under good growing conditions, and the
other made when conditions do not warrant growth and in fact necessitate cutting back on size,
using stored resources and changing strategy for obtaining nutrients to more risky but also higher
return rate strategies.
In general also the past ideas tied specific hormones to specific nutrients. Thus Auxin was seen
tied to growth warranting levels of sugar and gases, Cytokinin was seen as made when growth
warranting levels of water and minerals. Gibberellin was seen as made when there were less than
enough sugar and gases for growth and Ethylene was seen as made when there were less than
enough minerals and water to warrant growth.
ABA and Salicylic Acid at least in some versions were left out and not tied to any specific
nutrients. With ABA, this flew in the face of conventional wisdom because it has widely been
seen as a water deficiency signal. Instead ABA was seen as an emergency signal of any rapidly
developing adverse situation and SA (Salicylic Acid) as its all clear signal.
Growth Hormones - Auxin, Cytokinin, Salicylic
Acid, and Unknown Hormone X (Jasmonic Acid?)
Upon reflection I now am ready to reconsider my ideas and instead postulate an 8 hormone
system instead of a six. Note as with all my previous versions, I speculate that "growth levels" of
nutrients in a meristematic cell are twice as much in the shoot than in the root if it is supposed to
procure that nutrient (i.e. the shoot is supposed to procure sugar). This is because the shoot cell
will eventually have to support both it and a similar sized root cell in say sugar production. Also
then in the root, a meristematic cell needs to procure twice as much minerals as a similar sized
meristematic cell, before it will start producing Cytokinin.
Also note, if they have growth levels of the nutrient of which they are an indicator, Cytokinin,
Auxin and any other growth hormone should continue to be made by any plant cell as they
mature but it much lesser amounts. Perhaps the only reason for continuing to make the hormones
in mature cells, is to avoid senescence as small growth hormone production indicates productive
"profitable" cells (having nutrient levels higher than needed for the survival of that cell and any
cell dependent on it for production of that nutrient). Thus it is a good thing for instance, for a
mature leaf making more than enough sugar and taking in more gases than are needed for the
survival of both it and a similar sized root cell, that this shoot cell continue to receive a
guaranteed amount of water and maybe minerals too and for the leaf not to senesce and its
resources be reused for younger cell growth. This preservation for senescence is done at least in
part by the Auxin and Unknown Hormone X (see below - may be Jasmonate), it makes.
Auxin - made when there are growth levels of sugar in any meristematic cell.
Cytokinin - made when there are growth levels of minerals in any meristematic cell.
Salicylic Acid - made when there are growth levels of water in meristematic cells.
Unknown Hormone X (Jasmonic Acid?) - made when there are growth levels of needed gases,
Oxygen and Carbon Dioxide.
Senescent Hormones - Gibberellin, Unknown
Hormone Z (Brassinosteroid?), Abscisic Acid, Ethylene
There has been criticism of my work in the past in regards to which cells make of the
deficiency/senescence hormones. This was because I classify Gibberellin as a deficiency signal,
but the according the Plant Physiologists, GA (Gibberellin) is made more often in abundance in
meristematic tissue than mature tissue. In my earlier versions, I thought it was more beautiful
esthetically to see the deficiency hormones as made in higher amounts in mature tissue and
dropping off the younger the age of a cell is. I don't want to get in a discussion about that here
but suffice it to say, either way does not effect most of the content of my versions. In the 2007
version I did concede to the critics and made a version with all the hormones including those I
label as deficiency hormones (except Abscisic Acid and Salicylic Acid) being made at high levels
by meristematic tissue and dropping off as the cells mature.
Gibberellin - Sugar deficiency.
Unknown Hormone Z (Brassinosteroid?) - Mineral deficiency.
Abscisic Acid - Water deficiency. This is well accepted.
Ethylene - Oxygen and maybe gas deficiency. Ethylene has been accepted in the past by some as
an Oxygen deficiency signal.
Discussion and Ramifications
Most of the features discussed in previous version of the paper continue to hold in a similar way
as they did before.
Growth Hormone Nutrient Attraction - Because hormones also cause the attraction of nutrients
(and even other hormones), growth levels of hormones ensure the continued flow of
complimentary nutrients to a nutrient producer. Thus if a leaf cell is making Auxin, it draws
minerals, water and gases to itself to continue production of profitable amounts of sugar.
It may turn out Auxin even causes sugar to be attracted to site of Auxin synthesis and even more
strangely, sugar may relatively speaking be the nutrient which it attracts the most strongly to
where the Auxin is. The reason for this are two fold. First meristematic cells need to grow
quickly and may not produce enough sugar to grow at the rates the plant wants. That may be why
Auxin attracts sugar to meristematic cells in the first place. Secondly Auxin is transported down
the plant, so sugar follows this downward traveling of this. Jacobs and Gilbert first showed
around 1984 that Auxin transport pumps are located at the bottoms of the sugar transporting
phloem. So maybe Auxin is part of the engine that drives this sugar transport.
Similarly then Cytokinin is transported upward by the Xylem and it may more strongly attract
minerals to it from surrounding tissue, than other nutrients.
Perhaps the pre-eminent nutrient attracting properties of the growth hormones, for the very
nutrient of which they are an indicator of, occurs much more strongly in mature cells than
meristematic cells. Conversely then in the younger meristematic cells, then the nutrients the the
specific hormone attracts, would be more strong for the three categories of complimentary
hormones, and only weakly for the nutrient of which it is an indicator. The reason for the weak
attraction of the nutrient at all may be as indicated, to have a meristematic cell grow faster than it
could without the supplementation of the nutrient of which it is an indicator.
Growth Hormone Level Peak During the Day, Deficiency/Senescence Peak During the Night -
There is some evidence that this is true, but I saw this mostly in older journal articles. Suffice it
to say, that with colder temperatures and lack of light affecting the processes that require energy,
we might expect that there is less nutrient procurement at night and more of the requirement to
rely on stores. Also at night perhaps housekeeping and pruning of less efficient older plant parts
is more easily carried out (although it may be difficult to know if a leaf is an efficient sugar
producer at night!). This is all to say, deficiency/senescence hormones may be part of the normal
activity of a plant and not simply around when unusually bad environmental conditions exist.
There are other ramifications which you can read about in other versions, that should hold to be
still true with this 8 hormone theory too. For instance maybe now all four growth hormones,
Auxin, Cytokinin, Salicylic Acid and Unknown Hormone X are required for before a cell will
divide, and all four hormones Gibberellin, Unknown Hormone Z, Abscisic Acid and Ethylene are
needed before a cell will senesce.
I will finish the rest of this paper at a later date...10/17/2007
Further Reading
Barrington, E. J. W. Hormone. In The New Encyclopedia Britannica, Macropaedia v. 8, pp.
1074-88. Chicago: Encyclopedia Britannica, Inc., 1975.
Brown, A. W., Reeve, D. R., and Crozier, A. The effect of light on the Gibberellin metabolism
and growth of Phaesolus coccineus seedlings. Planta 126, 83-91, 1975.
Burg, S. P., and Burg, E. A. The interaction between Auxin and Ethylene and its role in plant
growth. PNAS 55, 262-69, 1966.
Engelke, A. L., Hamzi, H. Q., and Skoog. F. Cytokinin-Gibberellin regulation of shoot
development and leaf form in tobacco plantlets. Amer. J. of Botany 60, 491-95, 1973.
Goeschl, J. D., Pratt, H. K., and 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.
Hewett, E. W., and Wareing, P. F. Cytokinins in Populus x robusta Schneid: Light effects on
endogenous levels. Planta 114, 119-129, 1973.
Jahardhan, K. V., Vasudeva, N., and Gopel, N. H. Diurnal variation of endogenous Auxin in
arabica coffee leaves. J. Plant Crops 1 (Suppl), 93-95, 1973.
Lecoq, C., Koukkari, W. L., and Brenner, M. L. Rhythmic changes in abscisic acid (ABA)
content of soybean leaves. Plant Physiology 72 (suppl.), 52, 1983.
McMichael, B. L., and Hanny, B. W. Endogenous levels of abscisic acid in Water stressed cotton
leaves. Agron. J. 69, 979-82, 1982.
Mitsuhashi-Kato, M., Mishibaoka, H., and Shimokoriyama, M. Anatomical and physiological
aspects of developmental processes of adventitious root formation. Plant and Cell Physiology 19,
393-400, 1978.
Sembdner, G., Gross, D., Liebisch, H. W., and Schneidner, G. Biosynthesis and metabolism of
plant hormones. In Hormonal Regulation of Development I, ed. J. MacMillen, Heidelberg:
Springer Verlag, 1980.
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.
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.