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Plant hormones: ideal metabolic marker molecules.
S. Van Laer email@example.com www.plantevolution.be published on the web on 5/12/2010 version 1.0
Plant hormones or plant growth regulators are the signal molecules responsible for directing the growth of plants. The function of the different hormones are diverse and have been described in detail. The unravelling of the signalling pathways is ongoing. The same accounts for the biosynthetic pathways. Every year the amount of knowledge on plant hormones is growing. There is however one question which was not addressed until now. Why did these specific molecules became plant hormones? In this article an evolution theory will be presented that gives a possible answer to this question. According to the theory, the origin of the plant hormones is linked to the metabolic changes that occur just before or during the process for which the hormone signals. As many metabolic changes occur simultaneously, it is clear that a selection process has taken place selecting the most suitable molecule. In the theory presented, this selection process is linked to the energy loss or energy use of different metabolic processes. In the case of growth promoting signal molecules (auxin, cytokinin, gibberellins and brassenosteriods), a derivative of or a side product of the compound or building stone which acquires the most energy to build is the most suitable signal molecule. To minimize energy loss and optimize the energy efficiency of cells, these building stones will only be produced abundantly when the energy status of the cell is high. The same rule of energy efficiency as a selecting procedure applies also for the stress signal molecules (ethylene, absisic acid, salicylic acid and jasmonic acid). However in this case, it is not only the energy cost for the production of a building stone that can serve as a selecting procedure. The importance of the role a molecule plays in the stress response holds the key. For both the growth promoting hormones and the stress hormones it can be assumed that these molecules became a hormone because they were the most suitable metabolic marker molecules.
From one cell to multicellular
The evolution of a one cellular organism to a multiple cellular organism assumes coordinated growth. Coordinated growth is only possible when signal molecules are available that can be exchanged between the cells. Not only signal molecules are a prerequisite, but also the enzymes that produce the signal molecules and the receptors that respond to these molecules have to be present before a signal can turn into a response. It is however unlikely that all this appeared at the same time during the course of evolution. In order for the receptor to function, the signal molecule had to be present before the receptors. And in order for a signal molecule to occur, the enzymes that help to synthesis these molecules have to be there. In the beginning of the evolution of the multicellular organism, the cells however had no reason to produce these molecules and certainly had no reason to produce specific enzymes for the production of a molecule that still had to receive a function. Therefore it can be assumed that in the beginning the “to be signal” molecules were the products of a-specific conversion by enzymes of other metabolic pathways. The byproducts of these metabolic pathways were later recognized by new receptors as being the perfect
S. Van Laer 1 www.plantevolution.be
The evolution and selection of the different types of growth and stress signals are looked upon in the following paragraphs. namely tryptophan. Natural selection selects the solutions which require the smallest amount of energy and/or minimize energy loss. they will only be produced abundantly when the cell has enough energy to grow. Besides being essential components of the cell structure. The molecule had to be able to pass the membrane and cell wall without specific transporters as non were available at the beginning of the evolution of signal molecules. Based on the presumptions made above. proteins or derivatives of proteins are no candidates for becoming a growth signal molecule.plantevolution. Indol acetic acid is derived from tryptophan and functions as a plant hormone regulating the growth and enlargement of cells in S. Furthermore the stability of the candidate signal molecules was very important. Tryptophan. Because the synthesis of these amino acids is highly regulated. The ability to produce enough proteins perfectly indicates the growth status of the cell. (version 1. the correlation between the growth status and the production of an amino acid will be the greatest with the amino acid that requires the most energy to be build. The synthesis of these amino acids is highly regulated. even the amino acid composition of proteins is affected. however due to their size. The candidate signal molecule do not only had to represent the status of the cell. but also had to fulfil other requirements. natural selection has selected these cells which minimize energy loss. It can be assumed that during the course of evolution. However in this case the molecule cannot be the molecule that requires the least energy input because the energy input of the molecule must also reflect the high energy status of the cell. the signal will last longer than the status it represents. Proteins itself will only be produced in sufficient amounts if the building stones of the proteins. Other candidates are the derivatives of the building blocks of proteins. they are also necessary for the production of all the other building stones of the cell and the energy production. namely the amino acid. being a building block of proteins. Natural selection not only favoured a highly regulated amino acid production.0) 5/12/2010 molecules to represent the status of the multicellular organism. cannot be a signal molecule itself. Auxin Growth in a unicellular organism is determined by the ability to produce enough energy and building stones to allow cell enlargement. A lot of proteins fulfil this requirement. If a molecule is too stable. The candidate molecule therefore may only be produced abundantly when the energy status of the cell is high and when the cell is growing. the molecule also had to had unique structural properties that enables a receptor to recognize this specific molecule. namely the amino acids. Proteins play the central role in this process.Plant hormones: ideal metabolic marker molecules. Amino acids with high energy building costs are found more sparsely and seem to be present only when their special characteristics are necessary for the function of the protein. Beside all this. The above criteria all had to be fulfilled before a molecule could become a signal molecule reflecting the growth and stress status of cells. Derivatives of tryptophan do not have this restriction. Highly unstable molecules will react before transported to another cell. The selection of a molecule that indicates the growth status of the cell and which can function as a growth signal for other cells will be based on the same principal. can be produced in sufficient amounts. An overproduction of amino acids equals energy loss for the cell. especially the synthesis of these amino acids which require a large energy input during the production. Van Laer 2 www.be .
growth cannot only be based on metabolic building activity and status of division because both processes fluctuate. The question remains. A rise in the concentration of the nucleic acids reflects that the cell is going to divide. Cell division requires the cells to produce large quantities of nucleic acids. As the energy status is reflected by the amount of indole acetic acid. Cytokinin In multicellular organisms cell growth occurs coordinated. cells have to be able to signal their growth status to other neighboring cells and likewise have to be able to sense the growth status of the other cells. it is logical to assume that cell division can only occur when the level of indole acetic acid is high enough. requires a more advanced status marker. A higher status. which is the ideal metabolic marker reflecting the growth status and/or the capability for a cell to grow. Weather conditions can change radically and photosynthesis. it can be presumed that the dividing status is presented the best by a derivative of the nucleic acid that requires the most energy to be build. A rice in the concentration of free nucleic acids excreted by the dividing cells would be the perfect signal for signalling division if they were not a common building stone present in every cell.plantevolution.0) 5/12/2010 plants. a distinction can be made between cell enlargement and cell division. However. Auxins and cytokinins. Fluctuating signals are not the best signals for regulating growth. Furthermore. The nucleic acid with the highest building costs is adenine. for example signalling the presence of a seed to the surrounding tissue to induce the growth of fruit. Besides energy.be . which derivative of which nucleic acid is the most suited for presenting the dividing status of the cell. In order to coordinate growth. Van Laer 3 www. Cell division can only occur when abundant energy is available. two signal molecules derived from fluctuating processes. a derivative of adenine. This conclusion is reached by following the same logical selection procedure for signal molecules explained above for the signal representing cell enlargement. As mentioned above. Neighbouring cells have to be able to sense this. Because the production of nucleic acids requires a lot of energy and the cell uses its energy in an economical way avoiding energy loss and energy overconsumption. Derivatives of nucleic acids do not have this restriction and could therefore better fulfil this role. Auxins only represent a status of high metabolic building activity and cytokinins represent a status of cell divisions. Gibberellins and brassenosteriods Basic growth can be signalled by the metabolic markers indole acetic acid and zeatine. Otherwise irregular structures will be formed. fluctuates during the day and between days. Only then the cell will start duplicating its DNA. can S. cell enlargement is reflected by the production of the tryptophane derivative indole acetic acid. which is the main driving force for metabolic activity.Plant hormones: ideal metabolic marker molecules. In growth. is the signal molecule in plants signalling and promoting cell division. also the nutritional demands have to be fulfilled. a growth solely based on these two signals is insufficient because they only give a rough estimate of the status of the cell. Zeatine. (version 1. They can result in promoting growth of extra cell tissue which cannot be supported later on when photosynthetic activity is low.
A “check list” molecule is a molecule that can only be produced if the advanced status has been reached. Only in this way the check list molecule can represent and signal the presence of the advanced status. The production of gibberellins requires the presence of numerous enzymes. Gibberellins S. reflecting the extra growth potential of the cells. and tetraterpenoids. the phytol side chain of chlorophyll. A molecule signalling an advanced status has to be a “check list” molecule. Gibberellins are derived from geranylgeranyl pyrophosphate. The basic rules phrased at the beginning however stated that no specific hormone synthese enzymes can be present in a cell before the signal molecule. Chlorophyll derivatives cannot fulfil the requirements of a signal molecule stated in the beginning of the article. Building the extra photosynthetic apparatus will require the presence of nutrients and an already present optimal functioning photosynthetic apparatus that can deliver the energy needed. and an optimal functioning cell apparatus. A rice in geranylgeranyl pyrophosphate does not give detailed information on the presence and/or activity of a particular process in the cell. This does not endanger the plant at later time points when unfavourable weather conditions occur. Only when the requirements for extra photosynthetic apparatus rises and the cell possesses enough energy to fulfill these needs. If this is true. the concentration of geranylgeranyl pyrophosphate will rise. one or more of the enzymes only present in the case of the advanced status are used. Chlorophyll itself is far from an ideal candidate to become the precursor of a signal molecule. The synthesis of geranylgeranyl pyrophosphate is the last chemical reaction that these different groups of compounds share. Van Laer 4 www. are the gibberellins. The most expensive building block of chlorophyll is geranylgeranyl pyrophosphate.0) 5/12/2010 only account and signal for a limited growth. including carotenoids. When applying the same logic which was used for auxin and cytokinin. Geranylgeranyl pyrophosphate is the precursor of many diterpenoid compounds. Also.plantevolution.be . It is highly unlikely that chlorophyll derivatives can pass the membrane without a specific transporter. All this ma kes geranylgeranyl pyrophosphate an ideal candidate for a precursor of a signal molecule. geranylgeranyl pyrophosphate will only rise under optimal conditions for the plant. This leads to the logical conclusion that for the production of the check list molecule. The same accounts for gibberellins which are derived from geranylgeranyl pyrophosphate. Logical deduction leads to the conclusion that the enzymes needed for the production of gibberellins are enzymes that are also used for metabolic processes linked to the advanced status for which the gibberellins signal. Most of the time the concentration of geranylgeranyl pyrophosphate will be relatively low because different biosynthetic enzymes compete for geranylgeranyl pyrophosphate. Brassenosteriods are derivatives of cholesterol and therefore ideal candidates for signaling extra growth potential. Chlorophyll itself is build from several other basic building molecules/blocks. A rise in geranylgeranyl pyrophosphate reflects a status of energy and nutrient abundance. including quinines.Plant hormones: ideal metabolic marker molecules. the other most expensive building block of plant cells. The history of gibberellins is less clear. the presence of enzymes able to catalyse drastic changes in the structural properties of chlorophyll to make it a more diffusible molecule is not evident. In some cases however extra growth is possible and an extra signal molecule is necessary. allowing/inducing extra growth. although there is a relation with the synthesis of chlorophyll. (version 1. The two most expensive basic building blocks of the cell are cholesterol and chlorophyll. The “check list” molecule must be the result of a process that checked if all the requirements of the advanced status were fulfilled. The need for extra photosynthetic apparatus will occur when the light intensity is high. The molecules signalling an advanced status in a plant. An advanced status is however more than that. the signal molecule in this case must be a derivative of the most expensive molecule/building block of the cell. Many of the molecules derived from geranylgeranyl pyrophosphate play critical roles in photosynthesis.
Osmotic stress leads to a major burst of methylation of a magnitude that only occurs then. Every time a selection process took place during which a gibberellin was selected.0) 5/12/2010 are the “check list” molecules of the plant. The production of gibberellins can even be plant species specific.Plant hormones: ideal metabolic marker molecules. one candidate stands out: Sadenosyl methionine. a plant hormone S. but also serve as osmoprotectants of proteins. When looking at the synthesis of both betaine and methylated amino acids and proteins. This changes when drought stress increases. the plant increases the osmotic potential of the plant cell and this without interfering the activity of the enzymes present in the cells. This does not mean that one specific gibberellin only represents one specific advanced status. This gibberellin had to represented the status in the best way and may not be in conflict with an already coupled function to that specific gibberellin. At that time point a second defence mechanism will become more active. Choline. Other candidates that can not be ruled out are the precursors of these molecule because also the concentration of these molecules will rise in the case of drought stress. the precursor of betaine. Water is needed for the transport of nutrients. Plant growth is highly dependent on the availability of water. Different gibberellins represent different check lists. In this process. The first response in case of water loss will be an increase in the free sugar content of the cells. it will need more than one negative growth marker. By doing so. The plant cells posses several mechanisms to overcome the negative effects of the lack of water.be . Plants tend to cope with water deficit stress by a process known as osmotic adjustment. The positive markers are linked with light and nutrition. Proteins are no candidates due to their size. A limited water supply will therefore also limit the availability of nutrients and concordantly also the growth. (version 1. The negative effects upon the growth are limited at this stage. As plant growth can experience different types of stresses that constrain the growth. The production of specific osmolytes is enhanced and an increase in polyols. This makes S-adenosyl a good candidate for a precursor of the osmotic stress signal molecule. There is no need for the plant cells to signal to other plant cell to alter their growing behaviour. The sugars do not only preserve the osmotic equilibrium. Methylation of amino acid and proteins requires the methyl donor Sadenosyl methionine. Ethylene Until now only the positive markers for plant growth were discussed. methylated amino acids and betaine is observed. derivatives of sugars.plantevolution. During the course of evolution. The question now is: which of these compounds would be the best precursor for the signal molecule for osmotic stress? Polyols. A plant however also needs negative markers signalling unfavourable growth conditions. is the most methylated compound that can be found in a plant cell. Also the synthesis of betaine requires S-adenosyl methionine. Ethylene. new advanced statuses had to be signalled in the plant. Three molecules of S-adenosyl methionine are required to produce choline from serine. Furthermore the growth will be inhibited because the plant has to adept itself to the lack of water. are no candidates for signalling drought stress as they are easily metabolised by the neighbouring cells. methylated proteins. The remaining candidates for osmotic stress signalling are betaine and the methylated amino acids. It is possible that different advanced statuses lead to the production of the same gibberellin in different plant tissues or during different physiological ages of the plant. plants decrease their cellular osmotic potential by the accumulation of solutes. Van Laer 5 www. The most stringent growth restrain is experienced during drought.
Therefore the opening of stomata is regulated in a different way. for example heavy metals. an increase in osmotic stress will occur. Ethylene is linked with situations of extreme stress in which plant cells experience problems with sustaining their activity. The production of ethylene for ripening fruits probably evolved from the ethylene production during stress. Stomata are only found on higher plants and the movement of stomata do not act upon drought stress. A more volatile molecule will move quicker and this is especially important in a plant where sap flow ceased due to drought stress. However. It is there that the carotenoids are present.Plant hormones: ideal metabolic marker molecules. Ethylene signals the plant that drastic measures must be undertaken. The level of oxidative stress a plant experiences varies during the day and depends largely on the level of sunlight radiation. This does not account for glutathione and carotenoids. To much sunlight however can damage the plant. Also the carotenoids can not be ruled out as they protect the photosynthetic apparatus of the plant cell. Because oxidative stress can be live threatening. These mechanism capture and/or convert the oxygen radicals. (version 1. Van Laer 6 www. but close before the plant experience drought stress. During the course of evolution. such as the abscission of leaves. Oxidative stress specifically rises in the neighbourhood of the photosynthetic apparatus. antioxidants themselves can not serve as a signal molecule because they would not be recognized as such by neighbouring cells. As soon as the vascular tissue does not function optimal and/or the vascular tissue can not fulfil the needs of a plant organ. the carotenoids take the upper hand. High UV irradiation can cause oxidative stress in the plant. Carotenoids in the chloroplast help S.0) 5/12/2010 signalling stress in plants. Similar to auxin and cytokinin. Absisic acid Plants need sunlight to grow. Elevated oxidative stress levels enhance the production of antioxidants in a cell. The role of ethylene in ripening is only partly linked to stress.plantevolution. Oxidative stress can also be induced by toxic substances.be . The action upon the hormone ethylene results on the long term in the restoration of the osmotic equilibrium in the plant. Also the role of ethylene in fruit ripening was not discussed. the ethylene production capacity and the response of tissue upon exposure of ethylene was selected to govern the ripening of the fruit so that animals could be attracted that feed the fruits and disperse the seeds in the fruits. however the most important are ascorbic acid and glutathione. is produced from S-adenosyl methionine. The next question to answer is: the derivative of which antioxidant is the best representative of an increase in oxidative stress? There are numerous antioxidants. it is important that plants posses a plant hormone that signals an increase of oxidative stress. During the course of evolution plants obtained several mechanism to concur oxidative stress. the most likely candidate derivatives are oxidative breakdown products of the antioxidants. the better potential candidates are the derivatives of the antioxidants. The opening and closing of stomata were not discussed in the paragraphs above. Ascorbic acids and derivatives of ascorbic acid are easily metabolised in the neighbouring cells and are therefore not suited as a signal molecule. The fact that a volatile molecule became a plant hormone for osmotic stress probably is linked with the properties of the molecule and the status of the plant. The rice in antioxidants would be the ideal signal for oxidative stress. Because we are dealing with oxidative radicals reacting with the antioxidants. When comparing the potential of glutathione and carotenoids.
The fittest plant will be the plant that is able to react to the loss of expensive building blocks and avoid further losses of those building blocks. This links the production of absisic acid in the roots with water shortage.be . Absisic acid however is almost never linked to oxidative stress by scientists. gives a better idea of the severity of the oxidative stress a plant cell experiences. Oxidative stress due to irradiation is the highest in chloroplast which makes the oxidative breakdown products of carotenoids the best candidate for signalling oxidative stress. It is only logical to assume that the level of the signal molecule for oxidative stress will also rise in the roots. Water shortage also induces root elongation. Although derivatives of glutathione would give a good estimate of the overall oxidative stress in the cell. The reactions of a plant to the osmotic stress signalling molecule ethylene illustrate this assumption. Water shortage induces absisic acid synthesis in roots. Being a carotenoid and playing a role in the xanthophyll cycle made violaxanthin the ideal candidate precursor for the oxidative stress signal. The strength of the plant is determined by the thickness and composition of the cell walls and the architecture of the plant tissue. Root elongation requires cell wall loosening to enable cell expansion. The building cost of carotenoids is the second reason why carotenoids are better candidates then glutathione. Wounding a plant causes mechanical stress in the form of structural instability. For a plant to react to an increase of absisic acid in the roots has evolutionary advantages. (version 1. The protection is achieved by the xanthophyll cycle. A regulation of oxidative defence based on a signal coming from the breakdown of carotenoids is therefore likely to be more effective and efficient. for example hail and high wind velocity. The fittest plant is also the plant that only switches its extra defence mechanisms when necessary.Plant hormones: ideal metabolic marker molecules.plantevolution. starting at the onset of elongation. a signal molecule is found which is derived from the oxidative breakdown of carotenoids. These extremely reactive molecules attack cell wall polysaccharides. Salicylic acid A plant subjected to extreme weather conditions. Absisic acids best known function is to signal drought stress in plants. Cell wall loosening in roots is achieved by enzymes and by the production of hydrogen peroxide and superoxide. The reactive oxygen species do not only react with the polysaccharides but also other cell components. A rice in oxidative stress only occurs when the level of irradiation exceeds the xanthophyll cycle protection potential. will experience mechanical stress. The oxidative burst starts at the onset of elongation. or to grazing by animals. Evolution as a driving force of the building plan of the cell favours energy efficient architecture and operation. Absisic acid is produced from the oxidative breakdown of violaxanthin and neoxanthin. they would not estimate the effect of the oxidative stress on photosynthetic apparatus as good as the derivatives of carotenoids. The same accounts for pathogenic micro-organisms that breakdown the cell wall. Wounds S. A plant which is able to react to water shortage before an osmotic imbalance occurs is fitter then the plant that only reacts when the osmotic imbalance occurred. namely absisic acid. for example cows or caterpillars. which is transported to the upper part of the plant where it induces stomatal closure.0) 5/12/2010 capturing the sunlight and also protect the photosynthetic apparatus from oxidative stress. In plants. The loss of an expensive component which is not recycled that easily as in the case of glutathione. breaking the bearing structure of the cell wall. Excess light energy is dissipated through the xanthophyll cycle. The loss of high energy cost building blocks must be avoided. Van Laer 7 www. with the formation of zeaxantin from violaxanthin.
and will not be recognized as a signal molecules by other cells.plantevolution. As plant cells have to compete. The precursors of cellullose. do not diffuse true the cell membrane and cell wall. By doing so. pectin and lignine. Salicilyc acid is produced from alanine. From this it can be concluded that salicilyc acid is the result of a rice of alanine that is produced for the production of lignine to repair the mechanical injury.Plant hormones: ideal metabolic marker molecules. driving out water and strengthening the cell wall. The injury itself does not lead to an increase of a certain molecule. often cork tissue. During the course of evolution. The cells will increase the fluidity of the membrane. Lignin is build by alanine. The molecule that became the signal molecule for mechanical stress had to be a molecule that is produced more abundantly after a mechanical injury. the cell membrane becomes less permeable. Also oxidative stress can lead to membrane leakage when oxygen radicals react with the membrane. hemicellulose and pectin are sugars. Other candidates for becoming the precursor of the signal molecule are the precursors of cellulose. However they are still to large and it is highly unlikely that they can diffuse true the membrane without a specific transporter. The fluidity of the membrane will be increased by the production of cholesterol. A cell reacts upon membrane leakage by adjusting the permeability of the membrane. A derivative of alanine would be an ideal signal molecule for signalling mechanical stress to the neighbouring cells. Cellulose and hemicellulose form a network which is embedded in the pectin matrix. the concentration of different molecules rises. they will normally produce a membrane that fulfills the functions for which it is designed and they build it in the most economical way. The reason why membranes are not always more fluid has to do with the energy cost making the membrane more fluid. namely cellulose. All this changes when membrane leakage occurs. Reinforcement of the cell wall comprises a thicker cell wall and more cross links in the network. Sugars and their derivatives will never be signal molecules. Cell walls wil only contain lignin or beter stated just enough lignin to ensure rigidity of the cell wall. The large increase of lignine production after cell wall injury is there fore exceptional. pectin and lignine. The stability is reinstated by reinforcing the cell walls in the surrounding tissue. especially the molecules that are produced to fix the injury. However. The cell wall consist out of cellulose. This is reflected in the pectine composition of plant cell walls of different plants. Derivatives of these molecule could have unique characteristics so that they potentially could be recognized by other cells as a signal. hemicellulose. The lignin production of a plant is highly regulated because building lignin costs a lot of energy. Sometimes wound tissue is formed. natural selection favoured those plants that could signal mechanical stress and react upon that signal avoiding further damage. In the modern plant salicilyc acid is linked with signalling mechanical stress. Van Laer 8 www. Jasmonic acid The last type of stress discussed in this theory is membrane leakage.be . pectin and lignin. membrane associated proteins and the conversion of the lipids of the S. The same accounts for pathogens that produce toxins that disturb the membrane permeability. Non of these molecule poses the characteristics for becoming a signal molecule as they are all to large. hemicellulose. Membrane leakage can be inflicted by chemicals that react or interact with the membrane. during the reaction after the injury of the plant cells. Furthermore extra lignin is produced that penetrates the spaces in the network. hemicellose. (version 1. The increase of lignin will inflict also a large increase of alanine.0) 5/12/2010 inflict structural instability in the surrounding tissue.
Fatty acids with 3 double bindings have the greatest effect upon the liquidity. upon which the plant will react and defend itself. Because the biosynthesis of plant hormones uses enzymes of other metabolic processes. Extra cytokinin produced by such an enzyme would change the nutrient flow in the plant by directing more phloem vessels towards the fruit tissue. Jasmonic acid is a derivative of linolenic acid. namely linolenic acid will only be increased if it is really necessary. The production of the most costly saturated acid. When membrane leakage occurs. In my opinion it is illogical to assume that later on specific biosynthetic enzymes arose for the production of the signal molecules. However. Fruit produce cytokinin to induce extra phloem to redirect the sugars coming from photosynthetic active organs. The most common unsaturated fatty acid with 3 double bindings found in plants is linolenic acid. Signals for signalling the production of biosynthetic enzymes to produce a signal molecule for signalling is like contacting a person by telephoning another person and asking him if he would phone to the person you would like to contact. the development and growth of fruit asked for adjustment of the nutrient flow to fulfil the nutritional needs for growing fruit. There is however also a second possibility. Evolution selected signal molecules based upon existing metabolic processes in the plant that needed signalling for efficient coordinated growth. Furthermore. if specific biosynthetic enzymes would arise. it is linked to these processes and signals also the activity of these processes. This special characteristic of linolenic acid production was the reason why in plants jasmonic acid became the signal molecule for membrane leakage. the production of unsaturated fatty acids will increase. This could be a case where the plant gained the capacity to produce cytokinins during evolution in a plant organ that normally does not produce large amounts of cytokinin.Plant hormones: ideal metabolic marker molecules. new signals would be needed to activate these enzymes. Normally cytokinin production is linked to nutrient availability according to the theory presented above.0) 5/12/2010 membranes from saturated to unsaturated types. namely the production of cytokinin by enzymes from another metabolic process that is specifically active in fertilised seeds. Furthermore. Different types of unsaturated fatty acid are formed and these can have up to 3 double bindings.plantevolution. There could however be one exception to this rule. A boost in the production of linolenic acid is therefore probably exceptional. The influence of the unsaturated fatty acids upon the liquidity of the membrane is linked to the amount of double bindings. The expression of a specific cytokinin biosynthetic enzyme in the tissue that is destined to become fruit would solve this problem. Van Laer 9 www. The precursors of the signal molecules had to be key molecules in these metabolic processes in order to give a good reflection of the activity of these processes. namely in the case where during evolution plants benefited from drastic morphological changes that were only possible by redirecting normal growth. Overviewing the evolution of the metabolic markers From one cell to multicellular asked for collaboration. Especially the modification of the lipids increases the fluidity of the membrane.be . these signal molecules produced from these precursors had to be products of existing enzymes. This possibility could be an explanation why for many plant hormones there S. For example. It is unlikely that this extra value was lost later on during the evolution. in this case the plant needs extra nutrient for the growth of the fruits. They are also the most costly to build. (version 1. Jasmonic acid signals neighbouring cells membrane leakage stress.
This theory is published on the web. If the theory proofs to be wrong. The theory stated above can be found at www. (version 1.plantevolution. so that readers can react when finding evidence that sustains or refutes the hypotheses. The indole pyruvate pathways is more common in plant beneficial bacteria that operate in a less rich environment with more competition for nutrients. Researchers have also noticed differential biosynthetic activity in different parts of the plant and during different time point in the live time of the plant. but many different pathways occur.plantevolution. The latter is only the most economical in an environment with high nitrogen availablility. References are also normally published to sustain a theory. the theory can be put to the test by looking at the modern plants as sources of archaeological evidence. one can state that the evolutionary selection process of the theory starts now.be . In a way. The response upon the signal molecule however can not differ every time in the plant. In this case the sustainability is put to the test by the readers. Other theories on plant evolution and plant design will follow. Sometimes. This theory waits upon your reaction and your view. whereas others are limited to certain plant species. However. This assumption stated above also means that different metabolic states can lead to an increase of the same signal molecule. the theory itself can evolve. but many different biosynthetic pathways. Some of the pathways occur in all plants. but also articles that refuted alternative hypotheses. Almost all possible biosynthetic pathways seem to occur in plants. Not only articles that sustain this theory were important. Auxin biosynthesis in bacteria that specifically produce IAA to influence plant growth is done through the indole pyruvate and the indole acetamide biosynthetic pathway. Also the next theory “Plant hormones: ideal metabolic markers” can be found on this website. In this hypothesis also the relation between plant growth regulators and basic plant architecture will be addressed. References The theory was build by reading many articles. According to me. The situation in plants is totally different. The hypothesis presented in this article is the foundation on which the plant will be build in the next article. All researchers on plants must be thanked. the latter were even more important. Normally one would expect that evolution would only benefit and select the most energy efficient biosynthetic pathway. S.plantevolution.be. From one cell to a multicellular plant with coordinated growth governed by derivatives of metabolic processes. this can only be explained if the enzymes responsible for the production of the signal molecules play a role in the process that needed signalling. Furthermore the list of articles is still growing as this is not a static theory. As more evidence comes available.0) 5/12/2010 does not exist one biosynthetic pathway. The acetamide biosynthetic pathway is therefore more found in plant pathogenic bacteria that operate in and on the surface of the plant where there is enough nitrogen present for the bacteria. According to me this is the only possible explanation why evolution did not opted only for the most energy efficient pathways. Van Laer 10 www. The question now is: Is this view a correct view of what happened? Non of the above can be proven. This is the case for example in plant associated bacteria that produce auxin. The enzymes that govern the production of signal molecules can be looked at as sensors for certain metabolic processes that the signal molecules signal. Making a choice/selection which articles to refer to is therefore difficult.Plant hormones: ideal metabolic marker molecules.be. The theory presented above is my view on the origin of plant hormones. This problem will be addressed by the next hypothesis: “Building a plant: plant hormone functions revised”. adjustments will be made and published on www.
both signals have been linked with advanced growth regulation. Annex: comparing animal hormones with plant hormones Although only plant hormones were discussed in this article. Steroids are advanced status marker molecules.Plant hormones: ideal metabolic marker molecules. Animals do not produce there own tryptophane. namely signalling extra growth potential. other ideas were adjusted and new ideas were added. Cytokinins reflects nutrient availability in plants according to the theory presented above. The arguments given when explaining the origin of gibberellins and brasinosteroides also account for steroids of animals. the role of cytokinins in the growth has to be looked at in relation to auxin that reflects the activity of photosynthesis. The explanation for cytokinin is somewhat different as animals do produce adenine the precursor of cytokinin. The theory on the non existence of specific growth hormone biosynthetic enzymes is an adjusted idea based upon findings done by and ideas of other scientists. In plants. The crosstalk between nutrient availability and photosynthetic capacity is something that does not take place in animals. growth based on nutrients alone is limited to ensure that growth can be sustained on the long term. Furthermore. the explanation given for auxin and cytokinin also explain why these plant hormones do not fulfil the same function in animals. The originality of the ideas in this article is protected by www. Furthermore. two other growth signals appeared during evolution that signalled the possibility for extra sustainable growth. tryptophane itself performs the role of growth hormone in animals as the availability of amino acids determines if a cell can grow or not. (version 1. Signalling the concentration of tryptophane is not linked to a metabolic process that reflects the growth state potential of an animal cell. In the case of plants. In a way. a similar theory building can be done for animal hormones. Animals cells however do not need such a signal.be .boip. Van Laer 11 www. These hormones are produced from cholesterol. The new ideas in this theory are the explanation of the origin of the signal molecules. There are however some similarities between growth regulated by nutrients in animals and growth regulation by auxin and cytokinin in plants. A similar thing happened in animal cells. It is the crosstalk between those two hormones that determines the growth. Steroids have a similar function as gibberellines and brassenosteroids in plant cells. No specific growth signal is needed. others are not. The best example are the steroids. namely gibberellines and brassenosteroids.plantevolution. It is the nutrient availability itself that determines the growth.int S. Good ideas were taken over. Also in the case of animals cells.0) 5/12/2010 What is original and what is not? Some of the ideas in this theory are new.
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