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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.