WO2001056385A1 - Procedes et compositions permettant d'ameliorer le formyltetrahydropteroylpolyglutamate chez les plantes - Google Patents

Procedes et compositions permettant d'ameliorer le formyltetrahydropteroylpolyglutamate chez les plantes Download PDF

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WO2001056385A1
WO2001056385A1 PCT/US2001/003155 US0103155W WO0156385A1 WO 2001056385 A1 WO2001056385 A1 WO 2001056385A1 US 0103155 W US0103155 W US 0103155W WO 0156385 A1 WO0156385 A1 WO 0156385A1
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acids
percent
group
acid
enhancer
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PCT/US2001/003155
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Arthur M. Nonomura
John N. Nishio
Andrew A. Benson
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Nonomura Arthur M
Nishio John N
Benson Andrew A
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Priority to AU2001234685A priority Critical patent/AU2001234685A1/en
Publication of WO2001056385A1 publication Critical patent/WO2001056385A1/fr

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    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N59/00—Biocides, pest repellants or attractants, or plant growth regulators containing elements or inorganic compounds
    • A01N59/06—Aluminium; Calcium; Magnesium; Compounds thereof
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N37/00—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids
    • A01N37/36—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids containing at least one carboxylic group or a thio analogue, or a derivative thereof, and a singly bound oxygen or sulfur atom attached to the same carbon skeleton, this oxygen or sulfur atom not being a member of a carboxylic group or of a thio analogue, or of a derivative thereof, e.g. hydroxy-carboxylic acids
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N37/00—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids
    • A01N37/44—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids containing at least one carboxylic group or a thio analogue, or a derivative thereof, and a nitrogen atom attached to the same carbon skeleton by a single or double bond, this nitrogen atom not being a member of a derivative or of a thio analogue of a carboxylic group, e.g. amino-carboxylic acids
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N37/00—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids
    • A01N37/44—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids containing at least one carboxylic group or a thio analogue, or a derivative thereof, and a nitrogen atom attached to the same carbon skeleton by a single or double bond, this nitrogen atom not being a member of a derivative or of a thio analogue of a carboxylic group, e.g. amino-carboxylic acids
    • A01N37/46—N-acyl derivatives
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N37/00—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids
    • A01N37/44—Biocides, pest repellants or attractants, or plant growth regulators containing organic compounds containing a carbon atom having three bonds to hetero atoms with at the most two bonds to halogen, e.g. carboxylic acids containing at least one carboxylic group or a thio analogue, or a derivative thereof, and a nitrogen atom attached to the same carbon skeleton by a single or double bond, this nitrogen atom not being a member of a derivative or of a thio analogue of a carboxylic group, e.g. amino-carboxylic acids
    • A01N37/48—Nitro-carboxylic acids; Derivatives thereof
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N43/00—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds
    • A01N43/90—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having two or more relevant hetero rings, condensed among themselves or with a common carbocyclic ring system
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N61/00—Biocides, pest repellants or attractants, or plant growth regulators containing substances of unknown or undetermined composition, e.g. substances characterised only by the mode of action

Definitions

  • the present invention relates generally to methods and compositions for stimulating carbon nutrient uptake that yields enhanced growth in plants with improved water use efficiency.
  • Another event in the C 3 cycle shown in Figure 1 is photorespiration, during which oxygen (02) outcompetes CO 2 and is added to RuBP.
  • oxygen (02) outcompetes CO 2 and is added to RuBP.
  • phosphoglycolate is formed.
  • the phosphoglycolate is dephosphorylated to glycolate which is oxidized to glyoxylate.
  • Glycine is made by attachment of ammonia (NH 3 ).
  • the glycine is deaminated releasing NH 3 and further decarboxylated to CO, plus a single carbon ( ) fragment.
  • one molecule of glutamate can provide NH 3 for animation of glyoxylate to form glycine, while the other is recycled and combines with the NH 3 released when glycine is deaminated, as shown in Figure 2.
  • the serine formed must be recycled back to the RuBP pool for further carboxylations, otherwise photorespiration would drain the RuBP pool. Recycling serine to PGA involves an amino transfer from serine to glyoxylate to form glycine. The resulting 1- hydroxypyruvate is reduced to glycerate and then phosphorylated to form PGA. The PGA can be recycled back to RuBP.
  • the energy intensive process of photorespiration depletes 0 2 and releases CO 2 but salvages 75 percent of the carbons in the glycolate produced.
  • Ci metabolism in higher plants is discussed in Cossins, "One-carbon Metabolism", The Biochemistry of Plants, Vol. 2, Ch. 9, Pp. 365-418, Academic Press, Inc., 1980, and in Cossins, "Folate Biochemistry and the Metabolism of One-Carbon Units," The
  • U.S. Patent No. 3,897,241 describes application of ethanolamine formulations with carboxylic acids of less than 8 carbons, such as, oxalic acid, formic acid, acetic acid, phthalic acid and glutaric acid to fruit-bearing plants 10 to 150 days prior to ripening.
  • European Patent 465 907 Al describes compositions for stimulating the growth and ripening of plants comprised of at least one adduct of menadione bisulfite and a compound chosen from a group including pAB A.
  • British patent application 2 185 472 A describes foliar plant feeding compositions which comprise from 2 percent to 4 percent by weight of protein hydrolysate including amino acids, polypeptides, and oligopeptides. Particular amino acids are not identified.
  • oxamide H N-CO-CO-NH 2
  • the foliar application of radiolabeled proline to wheat is described in Pavlova and Kudrev (1986) Dolk. Bolg. Akad. Nauk. 39: 101-103. Barel and Black (1979) Agron. J.
  • foliar fertilizers incorporating polyphosphate compounds combined with a surfactant (0.1 percent Tween® 80).
  • U.S. Patent No. 4,863,506 describes the incorporation of L-(d)-lactic acid in foliar sprays where the lactic acid is alleged to act as a growth regulator.
  • U. S. Patent No. 4,799,953 describes polymeric condensates of the sulfur- polymers, thiolactic and thioglycolic acids, increasing the rate of growth and chlorophyll specific to tissue and hydroponic culture of Lemna minor.
  • the Ci-THF molecule can be segmented into distinct components including a formylpteroyl glutamide (the example shown in Fig. 6 is folinate carrying multiple glutamates) and a polyglutamate (Glu n ) chain.
  • the formylpteroyl glutamide can be further subdivided into a Ci-fragment, a pteridine and an aminobenzoylglutamic acid.
  • Ci-THF Foliar application of substances that increase the amount of Ci-THF, either by contributing any one of these components to the structure of Ci-THF or otherwise promoting the formation of Ci-THF, increases carbon fixation by the Ci pathway and enhances plant growth.
  • Such substances which increase the amount of Ci-THF in the leaf of a plant are referred to herein as “enhancers” or “enhancer substances”.
  • an enhancement of the carbon pathway focuses on modulating the flow of carbon through Q-THF in a manner that enhances fixation of Ci fragments in plants.
  • Q-THF is a catalyst for Q metabolism, meaning that Q metabolism is dependent upon Ci-THF.
  • the Ci-THF content of a leaf catalysis of Q fragments is enhanced, i.e., the plant's capacity to metabolize Ci-fragments by increasing the flow of carbon through the Ci-THF pool and thereby fix carbon into cellular constituents is increased proportionately. If the level of Ci fragments is then increased, this then results in increased plant growth according to this newly afforded capacity to metabolize what otherwise would have remained an under-utilized carbon source.
  • Ci-THF i.e., by increasing the formyltetrahydropteroylpolyglutamate pool, improved carbon fixation, water use efficiency and plant growth results.
  • the level of Ci fragments may be increased by exposure to appropriate environmental conditions or by the addition of substances which are capable of being metabolized to Ci fragments. All such substances are referred to as "C ⁇ input substances".
  • Ci-THF Organic compounds can find passage through Ci-THF as a consequence of the metabolism of much larger molecules from which Q fragments arise.
  • the incorporation of formimino-amino acids is illustrated in Figs. 4 and 5, wherein, a Ci fragment is transported as formiminotetrahydrofolate.
  • Ci-THF is involved in manners illustrated in Figs. 4 and 5, representing modifications from Besson et al (1993) and Cossins (1980) and (1987), infra.
  • a supramolecular complex in this scheme is thought to be a storage product, serine hydroxymetllyltransferase ⁇ glycine'folyl-polyglutamate.
  • such ternary complexes may be the natural selection for storage because of stability. Modulation of release permits control of the flow of Q-THF, glycine and SHMT. Based on the storage of the ternary complex, treatments are directed towards stimulating its conversion to maintain a high flow of carbon.
  • Q sources such as glycolate or carbon inputs leading to formate and formaldehyde
  • Q-acceptor compounds substances which serve as acceptors for Q fragments which would otherwise be converted to toxic metabolites.
  • These substances can be provided either as part of tile enhancer substance or as a separate substance.
  • these Q-acceptor compounds are sources of polyglutamate.
  • Q-acceptor compounds may also be glycine or substances that may be metabolized to glycine.
  • Ci-acceptor compound sources may include exposure of the plant to environmental conditions which increase polyglutamate levels using existing plant constituents, but this reliance on natural sources lends itself to inconsistency. Most direct Glu n chain-lengthening sources, such as glycine, will decrease plant growth when applied alone; but when applied in combination with enhancer substance(s), they increase plant growth.
  • Highest potency is achieved by foliar application of formulations which provide all the components of Ci-THF or readily metabolized precursors thereto, for example, a Ci fragment, a pteridine, an aminobenzoic acid and a glutamate, or metabolic precursors thereto.
  • Ci-THF Ci-THF
  • readily metabolized precursors thereto for example, a Ci fragment, a pteridine, an aminobenzoic acid and a glutamate, or metabolic precursors thereto.
  • ⁇ M micromolar
  • M molar
  • compositions disclosed herein be preassembled in this fashion.
  • Combinations of substances that contribute to one or more of the previously identified components of the structure of Q-THF also promote plant growth.
  • pteridines formulated with aminobenzoates and Glu n sources would provide the components for the plant to assemble an entire Ci-THF.
  • p- aminobenzoylglutamate may be applied as a single active component, leaving the plant to produce and attach a pteridine.
  • the Ci-THF structure may be segmented even further into additional components.
  • the formulation may comprise two compounds such as pAB A and potassium glutamate.
  • Ci-THF refers to tetrahydrofolates carrying a one-carbon unit.
  • the one- carbon unit can be carried in various oxidation states. In the most reduced form, it is carried as a methyl group. In more oxidized forms, it is carried as a formyl, formimino, or methenyl group.
  • Q-THF Increasing the level of Q-THF as described above gives the plant a greater capacity to fix carbon by the Q pathway. This capacity is then exploited by treatment of plants with substances which can be utilized by Ci-THF as Q fragments or by exposure of the plants to environmental conditions which increase the flow of Ci fragments into and through the Q- THF pool.
  • substances utilizable by Q-THF as Q fragments may be Q fragments themselves, e.g., formate, or may be substances which are metabolized to a Q fragment. All such substances are collectively referred to as "Q-input substances".
  • Environmental conditions which promote the flow of Q fragments into the Q pathway are generally those which promote oxidative metabolism of photorespiration, i.e, the photorespiratory cycle initiated by the oxygenase activity of RuBP carboxylase/oxygenase (Rubisco). These conditions are generally referred to as O -Uptake conditions. Exposure of plants to high levels of Q-input substances and/or O 2 -Uptake levels alone can be toxic because of buildup of intermediates to toxic levels and because of consumption of products of photosynthesis. Therefore, as described earlier, enhancers and Q-acceptor compounds are supplied to route consumption of photosynthetic products back into synthesis of sugar.
  • Enhancer substances are generally Q-THF, pteridines, pterins, pteroic acids, pteroyi derivatives, folinates, substituted benzoic acids, substituted benzoates and derivatives thereof; and salts, hydrates and surfactant-linked derivatives thereof.
  • the enhancer is applied as an aqueous solution at a concentration in the range from 0.0001 percent to 0.5 percent, preferably from 0.0001 to 0.1 percent.
  • Suitable pteridine compounds contribute to the structure of Ci-THF and are represented by the formula below wherein:
  • R° is independently selected from the group consisting of: methylene-aminobenzoate, optionally substituted on the benzoate ring; methylene-aminobenzoyl(Glu) n , wherein n is an integer from 0 to 10, optionally substituted on the benzoyl ring; and its corresponding dihydro- and tetrahydro-reduction products at positions 5, 6, 7, and/or 8 of the pteridine rings; and salts, hydrates and surfactant-linked derivatives thereof.
  • acidic enhancers may have very low solubility in cool water (25°C) and, therefore, require solubilization.
  • pteroic acid, pteroyl(Glu) n and substituted benzoates do not dissolve at sufficiently high concentrations in water to promote plant growth.
  • acids such as formic acid and acetic acid, alcohols such as methanol, alkali metal and alkaline earth metal bases such as potassium hydroxide and calcium hydroxide and carbonates to be formulated in aqueous solution with these agents.
  • Such formulations permit dissolution of the enhancer and facilitate penetration of the enhancer into the leaf of the plant. This then allows enhancement of Q-THF levels in the leaf and promotion of plant growth.
  • Activators are potassium hydroxide for aminobenzoic acids; normal potassium bicarbonate for folates and phthalic anhydrides; hexamethylenetetramine for pteroic acid; methanol for nitrobenzoic acids: and dimethylsulfoxide (DMSO) for terephthalates.
  • the Q-input substance for foliar application is usually selected from the group consisting of components that contribute Q fragments to the Q-tetrahydrofolate pool.
  • any carbon-containing substance which can be metabolized by any of the metabolic pathways in the leaf to generate a Q fragment which can be utilized by Ci-THF in the Q pathway can serve as a Q-input substance.
  • Q-input substances include formimino — , methyl — , methenyl — , methylene — and formyl-fragment sources.
  • Suitable single component enhancer substance formulations in which the Q-acceptor compound is inherent in the molecular make up of the enhancer include, but are not limited to, folinate, pteroyl(Glu) n , phthlaloyl(Glu) n , aminobenzoyl(Glu) n , nitrobenzoyl(Glu) n , and the like.
  • Ci-acceptor compound substances include compounds such as but not limited to glycine, glutamate, nitrates and formamidines.
  • elevated concentrations of CO can be applied to plants after application of an enhancer and a Ci-acceptor compound.
  • the plant is exposed to the elevated CO during daylight hours with continuous exposure to the Ci- acceptor compound substance and high light intensity.
  • the plant is moved back to air at night.
  • Methods of the present invention may comprise two steps, where a formulation of enhancer(s) is applied to the leaves of the plant to initiate enhancement of the Ci-THF pool. After passage of a time sufficient to allow for Ci-THF accumulation, the plant is exposed to conditions which increase the flow of carbon, such as by foliar application of Q-inputs and Ci-acceptor compound or exposure to O 2 -Uptake condition and a Q-acceptor compound.
  • the foliar application of Q -input substance(s) or the exposure to an O 2 -Uptake condition will be done at least twice following each application of the enhancer substance. Often it will be more than twice over a period from 1 day to 15 days following each enhancer substance application. It will be recognized that even though it is preferable to apply the enhancer substance before the Q -input substance, the order of application may be reversed.
  • Preferred Q-inputs for such a one-step application include, but are not limited to, formamidine # glycolate and formamidine* formate.
  • Preferred Ci-acceptor compounds for such a one-step application include, but are not limited to, glycine, glutamate, glutamine, and formamidine nitrate.
  • Compositions according to the present invention include plant growth promoting systems comprised of a first aqueous solution and a second aqueous solution.
  • the first aqueous solution contains an amount of an enhancer substance selected to increase Q-THF when applied to a plant.
  • the second aqueous solution contains an amount of Q-input + Q-acccptor compound substance selected to increase the flow of carbon in a leaf.
  • Preferred enhancer substances include but are not limited to folinate, pteroic acid, -nitrobenzoic acid and substituted benzoates, such as terephthalic acid.
  • Preferred Q-inputs include but are limited to carbon dioxide, glycolate, formamidine glycolate; formamidine . formate; formiminoamino acids; and other formimino — , methyl — , methenyl — , methylene — and formyl — sources.
  • Preferred Q-acceptor compounds include but are not limited to glutamate, glutamine, glycine and formamidines.
  • the first aqueous solution can contain a Q- acceptor compound substance and the second aqueous solution can contain an enhancer and/or a Q-input.
  • Another plant growth promoting composition for two-step application according to the present invention is a first aqueous solution comprising all enhancer and a second aqueous solution comprising a Q-acceptor compound.
  • Application of the enhancer is followed by exposure to 0 2 -Uptake conditions and application of the Q-acceptor compound.
  • plant growth is promoted by enhancement of Ci-THF in the leaf and increased flow of Q fragments in the Ci pathway by foliar application of Q-acccptor compounds with enhancers, Q-inputs, and/or with exposure to O 2 - Uptake conditions.
  • Figure 1 is a simplified depiction of the Q photosynthetic pathway in plants.
  • Figure 2 is an enlarged depiction of tile relation between the GOGAT (glutamine: 2- oxoglutarate amino transferase) cycle and oxygen uptake in the C 3 photosynthetic pathway in plants.
  • GOGAT glycol transferase
  • Figure 3 is a simplified depiction of an alternate pathway for glyoxylate, bypassing the GOGAT cycle.
  • Figure 4 is a detailed depiction of tile Q pathway, further illustrating paths for enhancing Ci-THF and the flow of Q fragments.
  • Figure 5 is a simplified depiction of a Q pathway, further illustrating paths for enhancers, Ci -inputs, Q -acceptor compounds and O 2 -Uptake.
  • Figure 6 is a structure of Q-THF with bars corresponding to the various components that can be applied to foliage for enhancement. Attachment of the single carbon fragment at the 5-position of the pteridine structure is shown with regard to availability of the 10- and 5,10-positions, also.
  • Figure 7 is a graphic depiction showing the effects of folinate or methanol on long- term CO 2 gas exchange. Soy foliage was sprayed with 20 ⁇ M folinate, 40 mM glycine, 1 percent dimethylsulfoxide and 0.05 percent HamposylTMC (T); or 5 M methanol, 40 mM glycine and 0. 1 percent Triton ® X-100 (V). Each point is the mean of 3 to 8 measurements:
  • Figure 8 is a graphic depiction of results of experimentation showing the long-term effects of Uvinul ® P-25 or formiminoglycine on CO 2 gas exchange. Leaves of sugar beets were sprayed with 0.5 mM Uvinul ® P-25, 13 mM glycine and 0.05 percent HainposyiTMC (o); or 2 mM formiminogiycine, 13 mM glycine and 0.05 percent HamposylTMC(v). Each point is the mean of 3 to 6 measurements of foliar CO 2 gas exchange.
  • Figure 9 is a graphic depiction of results of experimentation showing the long-term effects of formamidine acetate or potassium glycolate on CO 2 Assimilation (A). Soy foliage was sprayed with 30 mM formamidine acetate (FAM) plus 0.05 percent Hamposyl ml C (left frame) or 30 mM potassium glycolate plus 0.05 percent HamposylTMC (right frame). Arrows indicate when plants were treated. Each point is the mean of 3 separate measurements + SD. No SD shown is less than symbol size.
  • Figure 10 is a graphic depiction of results of experimentation showing the long-term effects of formamidine acetate on the Assimilation/Transpiration (A/T) ratio.
  • the present invention provides methods and compositions for promoting the growth of green higher plants, that is, all plants which are actively photosynthetic.
  • green higher plants is intended to include virtually all species with active light-gathering surfaces capable of receiving foliar sprays, particularly higher plants that fix carbon dioxide.
  • Higher plants include all plant species having true stems, roots, and leaves, thus excluding lower plants, yeasts and molds.
  • Q plants refers to all plants capable of fixing carbon via tile C 3 photosynthetic pathway.
  • Suitable plants which may benefit from Ci pathway carbon fertilization include crop plants, such as cranberry, cotton, tea, onions, garlic, leek, bach ciao, coffee, cassava, mustard, melon, rice, peanut, barley, broccoli, cauliflower, mint, grape, potato, eggplant, zucchini, squash, cucumber, legume, lettuce, kale, sugar beet, radish, kale, tobacco, alfalfa, oat, soy, turnip, parsnip, spinach, parsley, corn, sugar cane, Stevia, sorghum and the like; flowering plants, such as New Guinea Impatiens, geranium, passion fruit, breadfruit, poinsettia, Dusty Miller, mimulus, snapdragon, pansy, fuchsia, lobelia, carnation, impatiens, rose, coleus, chrysanthemum, poppy, gesneriads, bromeliads, bougainvillea, oleander,
  • crop plants
  • the methods and compositions of the present invention may be used to promote growth in photosynthetic parts of either juvenile or mature plants.
  • the plants include at least the sprouted cotyledon (that is the "seed leaves") or other substantial light-gathering surfaces, including, of course, the true leaves.
  • Improved growth occurs as a result of enhancement of Ci-THF as in Fig. 6 or via the Q pathway of Figs. 4 and 5.
  • High foliar content of Q-THF maintains high rates of carbon fixation even under detrimental conditions and plant growth is improved.
  • the aqueous solution of the Q- THF enhancer substance will be applied to the leaves of the plant, usually as a foliar spray, but also including dipping, brushing, wicking, misting, and the like of liquids, foams, gels and other formulations.
  • Foliar sprays will comprise atomized or other dispersed droplets of the aqueous solution which are directed at the plant leaves in such a way as to substantially wet the surface of the leaf with the minimum amount of the aqueous solution being lost on the soil, it will be desirable to minimize the amount of aqueous solution which is lost in the soil, with typically at least 75 percent of the aqueous solution being directed at the leaves, preferably at least 90 percent by weight, and more preferably substantially all of the solution being directed at the leaves.
  • Such foliar sprays can be applied to the leaves of the plant using commercially available spray systems, such as those intended for the application of foliar fertilizers, pesticides, and the like, and available from commercial vendors such as FMC Corporation, John Deere, Valmont and Spraying Systems (TeeJetTM).
  • spray systems such as those intended for the application of foliar fertilizers, pesticides, and the like, and available from commercial vendors such as FMC Corporation, John Deere, Valmont and Spraying Systems (TeeJetTM).
  • Suitable light and temperature conditions may be achieved by prolonged exposure of the plant to direct sunlight or other suitable high light intensity illumination sources while maintaining optimal to hot temperatures, usually above 20° to 35°C.
  • the plants should remain exposed to the sunlight or light intensity illumination for a period of time sufficient to allow for incorporation of treatments. Usually, the plants should remain exposed to sunlight or other illumination during daylight photoperiods for at least two hours before and for two weeks following fertilizer application. Sufficient nutrients must also be supplied to support healthy growth.
  • a Q-acceptor compound source is supplied with application of enhancer or Ci-input.
  • Enhancer compounds include those which increase Ci-THF.
  • Preferred contributors to Q-THE include its fragments such as folinate, pteridines, and substituted benzoates.
  • Q-input substances include those which increase the flow of Q fragments.
  • Preferred Q-input substances include organic compounds such as formamidine* glycolate and formamidine* formate that yield Ci fragments from Q- acceptor compound .
  • Preferred Ci-acceptor compounds include those such as formamidine nitrate, glycine and glutamate which add to the Glu n chain.
  • Activator compounds are those in which enhancers will dissolve prior to mixture with an aqueous solution. Preferred Activators also benefit plant growth by contributing nutrients and include methanol, potassium carbonate, potassium hydroxide, calcium hydroxide and acetic acid.
  • Plant illumination either sunlight or artificial, should have an intensity and duration sufficient to enhance photorespiration.
  • a minimum suitable illumination intensity is 100 ⁇ mol photosynthetically active quanta (400-700 nm) mV 1 , with direct sunlight normally providing much higher illumination.
  • Leaf temperature should be sufficiently high for optimal growth or hotter, usually above 25°to 35°C. It is preferable that the plant be exposed to at least two and preferably twelve hours of intense illumination following application of formulations.
  • Elevated carbon dioxide levels will be above normal atmospheric levels, that is above 0.03 percent, typically being above 1 percent, and preferably being above 10 percent. Such elevated carbon dioxide levels may be provided in controlled high light intensity environments, such as greenhouses, treatment chambers, protective crop and bedding covers, phytotrons, and other controlled-environment sealed enclosures for plant culture, and the like. Plants are initially treated with an enhancer to stimulate tile rate of carbon dioxide fixation. Enhancement of the Glu n portion of Q-THF by application of a Q- acceptor compound according to the present invention is necessary with exposure of plants to elevated carbon dioxide levels which, in the absence of such, would be toxic to many or all treated plants. Enhancer, Ci-input and Q -acceptor compound substances suitable for use in the methods and compositions of the present invention may be selected with reference to
  • FIG. 2 Figures 2, 3, 4, and 5 which are depictions of the Q pathway and Fig. 6 which depicts a representative Q-THF molecule.
  • Enhancer, Q -input and Ci-acceptor compound formulations may be applied to plants in sequence or in combination for improved plant growth.
  • formulations generally include a
  • Ci-acceptor compound substance Ci-acceptor compound substance.
  • Suitable enhancer substances include those compounds which are whole molecules, fragments or precursors of Q-THF in the pathway, as well as all salts, hydrates, aldehydes, esters, amines, and other biologically or chemically equivalent compounds and substances which can be metabolized in the leaf to contribute a component to Q-THF.
  • Preferred enhancer substances include Q-THF compounds, pteridine compounds, and substituted benzoate compounds.
  • Ci-THF compounds include folinates and compounds which may be converted to N-formyltetrahydropteroyl(Glu) n when applied to the treated plant.
  • Ci-THF compounds include folinic acid; anhydroleucovorin; 5- formyltetrahydropteroyl(Glu) n (depicted in Figure 5); 10-fo ⁇ nyltetrahydropteroyl(Glu) n ; 10- formyl-THF; 5-methenyl-THF; 5, 10-methenyl-THF; 5, 1 O-methylene-THF; 5,6,7,8-THF;
  • Ci-THF 5-formimino-THF; and ethoxylates, salts and hydrates thereof.
  • Ci-THF compounds will be applied to the plant as an aqueous solution having a concentration in the range from
  • pteridines are expected to promote plant growth. These include, but are not limited to neopterin, biopterin, leucopterin and the like. Suitable pteridine compounds contribute to the structure of Q-THF and are represented by the formula below, wherein:
  • R ,01 i hydrogen or is a hydrocarbyl group capable of being metabolized to a one carbon substituent having the oxidation state of a methyl, hydroxymethyl, formyl or formic acid residue; and R° is independently selected from the group consisting of: methylene-aminobenzoate , optionally substituted on the benzoate ring; methylene- aminobenzoyl(Glu) n , wherein n is an integer from 0 to 10, optionally substituted on the benzoyl ring; its corresponding dihydro- and tetrahydro-reduction products at positions 5, 6, 7, and/or 8 of the pteridine rings; and salts, hydrates and surfactant-linked derivatives thereof.
  • hydrocarbyl shall refer to an organic radical comprised of carbon chains to which hydrogen and other elements are attached.
  • the term includes alkyl, alkenyl, alkynyl and aryl groups, groups which have a mixture of saturated and unsaturated bonds, carbocyclic rings and includes combinations of such groups. It may refer to straight chain, branched-chain, cyclic structures or combinations thereof.
  • hydrocarbyl refers to a hydrocarbyl group which can optionally be mono-, di-, or tri-substituted, independently, with hydroxylower-alkyl, aminolower-alkyl, hydroxyl, thiol, amino, halo, nitro, lower-alkylthio, lower-alkoxy, mono- lower-alkylamino, di-lower-alkylamino, acyl, hydroxycarbonyl, lower-alkoxycarbonyl, hydroxysulfonyl, lower-alkoxysulfonyl lower-alkylsulfonyl, lower-alkylsulfinyl, trifluoromethyl, cyano, tetrazoyl, carbamoyl, lower-alkylcarbamoyl, and di-lower- alkylcarbamoyl.
  • lower as used herein in connection with organic radicals or compounds respectively defines such with up to
  • surfactant refers to surface-active agents, i.e., which modify the nature of surfaces, often by reducing the surface tension of water. They act as wetting agents, dispersants or penetrants. Typical classes include cationic, anionic (for example., alkylsulfates), nonionic (for example., polyethylene oxides) and ampholytic. Soaps, alcohols and fatty acids are other examples.
  • surfactant-linked derivative refers to a derivative of the parent compound, the derivative having a surfactant covalently attached to the parent compound.
  • a representative example of a parent compound and a surfactant-linked derivative thereof is p- aminobenzoic acid and the corresponding polyethoxylated -aminobenzoic acid (Uvinul ® P- 25).
  • R 01 Suitable non-limiting examples of R 01 include lower-alkyl, alkyl, hydroxymethyl, hydroxyalkyl, 1 -hydroxy alkyl, hydroxylower-alkyl, 1 -hydroxylower-alkyl, alkoxyalkyl, 1 ⁇ alkoxyalkyl, alkoxylower-alkyl, 1 -alkoxylower-alkyl, haloalkyl, 1-haloalkyl, 1 -halolower- alkyl, aminoalkyl, 1 -aminoalkyl, 1 -aminolower-alkyl, thioalkyl, 1 -thioalkyl and 1 - thiolower-alkyl.
  • Exemplary pteridine compounds are pterin; pteroic acid; pteroyl(Glu) n such as folic acid, pteropterin and pteroylhexaglutamylglutamic acid (PHGA); dihydrofolate; Rhizopterin; xanthopterin, isoxanthopterin, ieucopterin; and ethoxylates, salts and hydrates thereof.
  • Such pteridine compounds will be applied to the plant as an aqueous solution in a concentration in the range of about 0.0001 percent to 0.5 percent by weight, preferably in the range of about 0.0001 percent to about 0.1 percent.
  • Suitable substituted benzoate compounds contribute to the structure of Q-THF and are represented by the formula below, wherein:
  • R is H, hydrocarbyl, halogen; -OH; -SH, NH 2 , N-linked amino acid, N-linked polypeptide, -OR 3 , -SR 3 , NHR 3 , wherein R 3 is selected from the group consisting of optionally substituted hydrocarbyl, alkyl, acyl, amino acids or polypeptide chains, -NR 4 R 5 wherein R 4 and R 5 which may be the same or different and are independently selected from the group consisting of H, optionally substituted hydrocarbyl, alkyl, aryl, acyl, C-terminal linked amino acids, C-terminal linked polypeptide chains, or R 4 and R 5 together with the nitrogen atoms to which they are linked form a heterocyclic ring;
  • R 1 and R 2 are independently selected from the group consisting of: optionally substituted hydrocarbyl groups, alkyl, aryl, acyl, aroyl, halo, cyano, thio, hydroxy, alkoxy, aryloxy, amino, alkylamino, aminoalkyl, arylamino, aminoaryl, acylamino, ureido, alkylureido, arylureido, hydrazino, hydroxamino, alkoxycarbonylamino, aryloxycarbonylamino, nitro, nitroso, carboxy, alkoxycarbonyl, aryloxycarbonyl, aminocarbonyl, carboxamido, monoalkylaminocarbonyl, dialkylaminocarbonyl, formyl, sulfo, sulfamoyl, sulfoamino, alkylsulfonyl, arylsulfonyl, s
  • Cationic salts of the benzoates include cations selected from the group consisting of cations of alkali metals, alkaline earth metals, ammonium, organic ammonium (amine), quaternary ammonium, and mixtures of such salts.
  • aminobenzoic acids such as m-aminobenzoic acid and /7-aminobenzoic acid
  • aminobenzoic acids such as m-aminobenzoic acid and /7-aminobenzoic acid
  • derivatives such as N-benzoyl amino acids, N-acyl- aminobenzoic acids, aliphatic aminobenzoate esters, aliphatic N-acyl-aminobenzoate esters, N-acyl-N '-aminobenzoyl-amino acids, N - formylaminobenzoic acids, 2-chloro-4- aminobeuzoic acid, and ethoxylates such as Uvinul ® P-25; nitrobenzoic acids and derivatives such as m-nitrobenzoic acid p — nitrobenzoic acid, nitrobenzoyl amino acids, polyethyleneglycol nitrobenzoate, 4-chloro-2-nitrobenzoic acid, 2-chloro-4-nitrobenzoic acid;
  • Activator solution dissolve with enhancer substances in water and include organic acids, particularly hydrocarbyl acids and aliphatic alkyl acids such as, for example, formic acid, acetic acid, propionic acid and the like.
  • Activators include alkali and alkaline earth hydroxides (KOH, NaOH, ammonium hydroxide, Ca(OH) 2 and the like); alcohols such as methanol, isopropanol, and ethanol; alkali and alkaline earth carbonates and organic bases such as pyridine, diethylamine; surfactants such as, but not limited to phthalic glycerol alkyd resin (e.g. Latron B-1956TM; and penetrants such as organic solvents, particularly dipolar aprotic solvents such as DMSO.
  • Preferred Activators which are also Q-inputs include methanol, trimethylorthoformate, hexamethylenetetramine and DMSO.
  • Penetrants are typically organic solvent-based carriers which enable the applied substance to penetrate into the plant leaf.
  • the sustained flow of Q-carbon required for rapid growth is preferably accomplished by application of Q -input with Ci-acceptor compound to the plant being treated.
  • These Ci-input with Q-acceptor compound compositions can be applied as distinctly separate formulations or in combination with enhancers.
  • Q-inputs withQ- acceptor compounds are preferably applied separately after enhancer substances usually at least about 6 hours after enhancer substances had been applied, and preferably at least one day to one week after the enhancer substance had been applied.
  • the Q-inputs with Q-acceptor compounds will be applied at least twice between successive applications of an enhancer substance, frequently being applied from 2 to 10 times between such successive applications. It would be possible in some cases to apply the Q-inputs with Ci-acceptor compounds continuously between successive applications of another enhancer substance.
  • the aqueous solution containing an enhancer substance such as folinate may be applied periodically with successive applications being spaced apart by a period in the range from 7 days to 20 days, with aqueous solutions comprising the Q-input such as formate with a Ci-acceptor compound such as glutamine being applied from 1 time to 50 times between such successive applications.
  • Ci-input compounds that pass Q fragments include, but are not limited to, formamidine carboxylate salts selected from the group consisting of formamidine glycolate, formamidine acetate and formamidine formate; a formimino amino acid selected from the group consisting of formiminoglycine, foimiminoglutamate, formiminoalanine, and formiminoaspartate; a carboxylic acid selected from the group consisting of glycolate, oxalate and formate; an aldehyde selected from the group consisting of formaldehyde and acetaldehyde; a trialkylorthoester selected from the group consisting of trimethylorthoformate, triethylorthoformate; an N-formylated organic compound selected from the group consisting of diformylhydrazine, formamide, methyl formamide and dimethyl formamide; an acetamide selected from the group consisting of acetamide, methyl acetamide and dimethyl acet
  • Ci-acceptor compounds act as Ci sinks and/or enhance Glu n as a consequence of Q-THF metabolism.
  • Formamidines such as, formamidine nitrate
  • Other suitable Q-acceptor compounds include, but are not limited to, glycine, glutamine, glutamate, serine, sarcosine, homocysteine, cystathionine, methionine, hexamethylenetetramines, and formamide.
  • Suitable Ci-acceptor compounds are also available as conventional nitrogen fertilizers. They include nitrates, ureas, and the like.
  • Ci-acceptor compounds are utilized differently from conventional fertilizers in the present invention because they are used in combination with enhancers and Q-inputs to enhance carbon fixation. They directly target increases of the polyglutamate component of Ci-THF in the leaf. This has a safening effect that eliminates toxicity of Q- inputs. Thereby, application of high concentrations of Q -inputs results in carbon-based growth rather than retarding or killing the plant.
  • the more closely the Q-acceptor compound resembles Glu n the lower the energy requirement for metabolism; therefore, the simpler compounds of conventional fertilizers generally do not contribute to Q-THF as efficiently as do the preferred Ci-acceptor compounds.
  • the Q-acceptor compound may come as an inherent part of the enhancer and Ci-input molecules to more fully enhance Q- THF rather than being utilized strictly for nitrogen.
  • Suitable Ci-acceptor compounds may also be linked to surfactants to enhance penetration into the leaf, such as, for example, with PEG glutamate or with HamposylTMC.
  • compositions used in this invention are mixtures for promoting growth of a plant comprising an aqueous solution or a nonaqueous material which when combined with an aqueous carrier contains an enhancer such as p-nitrobenzoic acid or folinic acid; a formamidine salt of a carboxylic acid (for example., formamidine formate or formamidine glycolate) and agronomically suitable additives.
  • PEG threadPNBA polyethoxylated -nitrobenzoic acid
  • Ci-input with Ci-acceptor compound will typically be applied at a concentration ranging from 0.001 percent by weight to 5 percent by weight.
  • Preferred Ci - input with Q -acceptor compound formulations include trimethylorthoformate with formamidine nitrate applied as an aqueous solution at a concentration in the range of 0.01 percent by weight to 1 percent by weight; formate and potassium glutamate applied as an aqueous solution at a concentration in the range from 0.001 percent by weight to 5 percent by weight; and glycolate and formamidine nitrate applied as an aqueous solution with glycolate at a concentration in the range from 0.01 percent by weight to 0.5 percent by weight and formamidine nitrate in the range from 0.001 percent to 1 percent by weight.
  • Q-input:Q-acceptor compound ratio will be in the broad range from 1 ,000: 1 to 1 : 100 to a narrow range of 5: 1 to 1:1.
  • compositions of the present invention may consist essentially of the aqueous solutions of Ci-acceptor compound substances with enhancers and Q-inputs, they will usually contain other ingredients and components which improve performance in various ways.
  • compositions will usually contain a surfactant present in an amount sufficient to promote leaf wetting and penetration of the active substances, and optionally other components.
  • Suitable surfactants include anionic, cationic, nonionic, and zwitterionic detergents, such as ethoxylated alkylamines, quaternary amines, LED3ATM, TeepolTM, Tween ® , Triton ® , LatronTM, DawnTM dish detergent, and the like.
  • penetrants such as, dimethylsulf oxide (DMSO), sodium dodecylsulfate (SDS), formamides, and lower aliphatic alcohols
  • DMSO dimethylsulf oxide
  • SDS sodium dodecylsulfate
  • formamides formamides
  • lower aliphatic alcohols lower aliphatic alcohols
  • Ethoxylation of an active component or otherwise chemically modifying the active components by incorporating a penetrant substance is preferred because, as exemplified by Uvinul ® P-25, formulation without additional surfactant is achieved.
  • formulations will often include one or more conventional fertilizer constituents such as nitrogen, phosphorus, potassium, and the like.
  • Compositions may further comprise secondary nutrients, such as sources of sulfur, calcium, and magnesium, as well as micronutrients, such as chelated iron, boron, cobalt, copper, manganese, molybdenum, zinc, nickel and the like. Incorporation of such plant nutrients into foliar fertilizer formulations is well described in the patent and technical literature.
  • Other conventional fertilizer constituents which may be added to the compositions of the present invention include pesticides, fungicides, antibiotics, plant growth regulators, and the like.
  • Compositions according to the present invention may be tailored for specific uses, including water use efficiency; enhanced performance under environmental stress; aftermarket caretaking; floral or fruit optimization, and in all areas of agriculture in which heightened carbon fixation is beneficial. Compositions may also be formulated at very low concentrations for liquid suspension culture media.
  • Formiminoglycine 0.001 percent tol percent 0.05 percent to 0.3 percent
  • Calcium folinate 1 ppm to 500ppm 5 ppm to 50 ppm Surfactant (Tween ® 80) 0.01 percent to 0.5 percent 0.05 percent to 0.1 percent DMSO 0.5 percent to 3 percent 0.8 percent to 1 percent
  • KOH Activator 0.01 percent to 1 percent 0.1 percent to 0.3 percent Surfactant (TWEEN ® 80) 0.1 percent to 1 percent 0.2 percent to 0.5 percent FeHEEDTA 0.1 ppm to 5 ppm 1 ppm to 3 ppm Potassium glutamate 0.01 percent to 1 percent 0.1 percent to 0.5 percent Phosphate buffer pH 5 to pH 7 pH 5.5 to pH 6.5
  • compositions For use on plants subjected to O 2 -Uptake, one-part compositions may be formulated.
  • the one-part composition is typically composed of low concentrations of substances iii combination with a surfactant in a single solution. Exemplary one-part compositions follow.
  • Chemicals (abbreviations) and sources sodium glutamate, Ajinomoto; formamidine formate (FAF), glycine (Gly), HAMPOSYLTMC, potassium glycolate (GO), potassium phosphate (KOH), and purified water, Hampshire Chemical Corporation; adenosine triphosphate (ATP), m-aminobenzoic acid (MABA),/?-aminobenzoic acid
  • Radioisotopic 14 C0 2 was applied to plants to determine the fate of active substances and changes in the path of carbon fixation.
  • Cabbage plants were sprayed with one of the following three solutions: 1) 90 ⁇ M folinate, 0.2 percent glycine, 1 percent DMSO, and 0.1 percent Triton® X-100; 2) 40 percent MeOH, 0.2 percent glycine, and 0.05 percent
  • HamposylTM C 3) 0.5 mM pABG, 0.2 percent glycine, and 0.05 percent HamposylTMC.
  • a quartz halogen light type EKE, 21 V, 150 watt
  • a leaf that was to be used for experiments was placed in an open chamber that was constantly flushed with pure O 2 during the acclimation period.
  • a leaf plug, 3.67 cm 2 was then removed, and placed in a hermetically sealed PlexiglassTM leaf chamber containing pure O 2 being pumped at a rate of 2-3 L min _1 .
  • the chamber was illuminated with 1 ,000 ⁇ mol photosynthetically active quanta m ⁇ s " directed through a fiber optic cable connected to a quartz halogen light similar to the one used for preillumination.
  • 5 mL CO 2 containing 0.8 ⁇ Ci Na 14 CO 2 (specific activity of 5 Ci mol "1 ) was injected with a syringe to a final concentration of about 700 ppm CO 2 .
  • the leaf plugs were allowed to incorporate 14 C0 2 for 15, 60 or 180 s, and then fixation was immediately stopped. In other experiments the leaf plugs were pulsed for 15 s, then chased for 1 min or 3 min. The chase was carried out under ambient air.
  • Enzymes related to Ci-THF were tested for enhancement following treatments.
  • the three enzymes that comprise Q-THF-synthase, 10-formyl-THF-synthetase (EC 6.3.4.3), 5,1 0-methenyl-TH F-cyclohydrolase (EC 3.5.4.9), and 5, 10-methylene-THF-dehydrogenase (EC 1 .5. 1.5) were assayed according to methods of Edwin A. Cossins (University of Alberta, Edmonton, Canada).
  • Leaf tissue was extracted in 25 mM n- ⁇ 2-hydroxyethyl] piperazine-N'-[ethanesulfonic acid], 0.3 ⁇ M aprotinin, 10 mM KC1, and 10 mM 2- mercaptoethanol, adjusted to pH 7.5 with KOH.
  • a test tube containing 100 ⁇ L 1 M triethanolamine buffer (pH 7.5), 100 ⁇ L 1 M ammonium formate (pH 7.5), 100 ⁇ L 25 mM MgCl 2 , 100 ⁇ L 2 M KC1, 100 ⁇ L 10 mM THF, freshly made 100 ⁇ L 20 mM ATP, and 390 ⁇ L water was pre-incubated for 2 min at 30°C.
  • a 10 ⁇ L aliquot of the extracted sample containing enzyme was added and incubated 15 min at 30°C.
  • the reaction was terminated by adding 2 mL 0.36 N HCl .
  • the 10-formyl-THF formed during the incubation period was converted to 5, 10-methenyl-THF by the acid.
  • the absorbency was measured at 350 nm against a water blank and the 5, 10 — methenyl — THF formed was determined by using an extinction coefficient of 24,900 M "1 cm "1 .
  • Photosynthetic CO Gas Exchange was determined by taking measures with CIRAS-1 (PP Systems, Bedford, MA)or LI-6200 (LI-CORTM, Inc., Lincoln, NE) portable gas exchange system. The quantity of the Ci-THF pool was determined in two steps. Culture, treatment, collection and preparation of samples for shipment was undertaken at the University of Wyoming. Microbial assays were undertaken at the University of Alberta. Sugar beets ((Beta vulgaris L.) cv Monohikari (Seedex, Longmont, Colorado)) were grown under standard greenhouse conditions described above.
  • tissue extracts for Ci-THF pool quantification 0.5 g fresh live leaf tissue from different plants was placed in 61 mM ascorbate buffer (pH 6 adjusted by KOH). The samples were then immediately placed in a boiling water bath for 15 min. After cooling, the tissues were homogenized and centrifuged at 18,000 g for 10 min. The supernatant was collected. The pellet was extracted two more times and the supernatants were combined. The samples were sealed, frozen and packed on dry ice for express shipment to the University of Alberta (Department of Biological Sciences, Faculty of Science, Edmonton, Canada). Upon receipt, E.A. Cossins and S.Y.
  • the formulations included 50 ⁇ M folinate, 40 mM glycolate and/or 40 mM glycine each with Triton ® X-100 surfactant sprayed on tomato plants grown identically in movable containers. Four hours following treatment, the plants were moved from direct sunlight to shade or the reverse. In experiments to measure growth yields, plants were cultured in greenhouses or PercivalTM growth chambers at the University of Massachusetts Cranberry Experimental Station, East Wareham, MA. The plants were grown in 8 to 16 cm diameter pots containing PerliteTM with VermiculiteTM and topsoil. In greenhouses, no special control of physical conditions was attempted, but all comparable treatments were made simultaneously and were subjected to normal greenhouse conditions. Each sample was tested on 4 or more replicate containers of plants.
  • foliar treatments contained 0.5 ml/L of surfactant selected from HAMPOS YLTM C, Tween ® 80, or Triton ® X- 100. Hoagland solution nutrients in water were routinely supplied as needed. Seven to ten days after the last treatment, the plants were removed from pots and the roots were cleaned. Shoot and root lengths and fresh and dry weights were determined. Changes in shoot and root growth were recorded in all cases, but data on roots is given only for radish tests while data on shoot growth is given for other plants. All experiments used 4 replicates and results were subjected to analysis of variance and mean separation by LSD test and showed significance within narrow standard deviations in all cases. The concentrations of compounds were bracketed around optimal concentrations for growth. The plants tested for growth response included radish (cv Cherry Bell), wheat (cv Stoa), rice (cv M-202) and corn (cv Butter Sugar).
  • Ci-THF enzymes but glycine, HamposylTM C, FAM and oAB A did not. Folinate increased enzyme activity in com. The addition of glycine to NB A did not depress the overall stimulation of Q-THF enzymes as compared against similar glycine levels that, alone, depressed enzyme activity. This result is indicative of the synergistic involvement of the metabolism of enhancers with Ci-acceptor compounds.
  • gas exchange was enhanced significantly in soy, sunflower, cabbage, kale and sugar beet by foliar treatments with folate, folinate, pAB A, pNBA, pFBA, Uvinul ® P-25, FIGly, FAM, FAM with GO, BenzocaineTM and MeOH.
  • Gas exchange was significantly higher after Uvinul ® P-25 treatments as compared to pAB A treatments.
  • ⁇ AH stimulated gas exchange in soy. Neither oABA nor MecABA showed increases on gas exchange.
  • Figures 7, 8, 9 and 10 summarize results of the long-term effects of various treatments on gas exchange.
  • Figure 7 compares the effects of foliar sprays of 20 ⁇ M folinate to 5 M MeOH and showing increased CO 2 gas exchange over soy controls lasting three weeks. Plants were treated at the start and at 120 h. Peak activity was observed during the first and second weeks, the response to folinate far exceeding responses to MeOH.
  • Figure 8 shows that 0.5 mM Uvinul ® or 2 mM FIGly applications to sugar beet increased CO 2 gas exchange for ten days following treatments.
  • Figure 9 shows that 3 foliar sprays of 30 mM FAM or 30 mM GO formulations increased gas exchange over controls for approximately a week between treatments.
  • Figure 10 shows the relationship between assimilation and transpiration. Measurements of the assimilation/transpiration (A/T) ratio in cabbage leaves treated with a formulation containing 30 mM FAM showed CO 2 assimilation maintained with reduced transpiration for the duration of the experiment. Generally, gas exchange increases were observable several hours after treatment.
  • Active components were formulated in the following control solutions: (A) 0.05 percent HAMPOSYLTM C surfactant, (B) 0.2 percent glycine + 0.05 percent HAMPOSYLTM C, (C) 200 ppm potassium formate + 0.2 percent glycine + 0.05 percent HAMPOSYLTM C, and (D) water. Folinate was followed by GO lx once at 2 d or G0 2 X twice at 2d and 4d. **Bean yeild was measured at University of Wyoming. The treatments given in Table 8 were selected from a broad range of concentrations of each formulation that was applied and represent optimal doses for growth improvement in each case. The most effective concentration is directly related to the volume of solution applied, therefore, the actual concentration must be adjusted according to the quantity absorbed into foliage.
  • the Q-THF pool was substantially enhanced by our plant treatments. Foliar applications of ⁇ M to mM quantities of substances such as folinate, substituted benzoates and pteridines that appear to contribute to the Ci-THF structure increased rates of CO 2 fixation, sugar, photosynthetic gas exchange, Ci-THF enzymes, Q-THF and water use efficiency. The improved physiology and biochemistry we observed was directly correlated to improved growth of green plants that we tested. Furthermore, enhancement of Ci-THF followed by Ci-input pulses or by continuous exposure to O 2 -Uptake, plant growth improved for long durations. We conclude that enhancement of Ci-THF increases the efficiency of carbon fixation into plant growth.
  • Glycolate accumulation was assayed in leaf discs floated on aqueous solutions at 0.39 mmol photosynthetically active quanta ⁇ V 1 and 30°C. These conditions are not necessarily photoinhibitory, but glycolate synthesis was apparently decreased in the leaf discs treated with 30 mM L-glutamate (Oliver & Zelitch 1977b). Inhibition of photosynthesis caused by accumulation of glycolate has been reported (Ogren 1984; Wendler et al. 1992; Zelitch 1978) and it is generally accepted that glycolate must be further metabolized, otherwise, it is detrimental to photosynthesis. Formate feeding to plants has been used to determine how formate is metabolized (Tolbert 1955; Tolbert 1981).
  • Ethoxylates of other active compounds such as pteroic acid, phthalates or nitrobenzoates would be useful for single component product formulations.
  • Ethoxylating pAB A has the additional benefit of improving the solubility of p ABA in water thus characterizing ethoxylation as an Activator.
  • Salts, such as, calcium folinate, are water soluble; furthermore, in view of its reduced state, folinate has long been considered the activated form of vitamin M (Merck Index, monograph #4111).
  • Contributors to the Ci-THF pool that we now know to increase Ci metabolism may modulate the flow of carbon through the C 3 cycle.
  • QT-HF is present in fungal spores and can originate nonenzymatically from 5, 10-methenyl-THF.
  • a futile cycle in the fungus Neurospora crassa exists, in which, rather than rapidly contributing to the flow of carbon, C THF binds tightly to serine hydroxymethyltransf erase, glycine which further inhibits methenyl-THF-synthetase and other enzymes downstream (Stover et al. 1993; Kmschwitz et al. 1994) limiting the supramolecular ternary complex to a slow release of its contents.
  • Q-THF-dependent enzymes bind Ci-THF in pea foliage (Besson et al.
  • the Glu n stored in the Ci-THF pool may be released over several days or weeks depending on demands to maintain equilibrium during photorespiration. Regulation of photorespiration may be determined according to maintenance of high flows of Ci fragments, an equilibrium that can benefit growth with maintenance of high glycine: serine ratios. In our studies, it is possible that appreciable quantities of glycine were separated from the serine hydroxymethyltransf erase, glycine* pteroylpolyglutamate ternary complex during processing of our samples for chromatographic separations, our data thus fitting this pathway. In our studies of the Ci-THF pool, when FIGly was used the greatest increase was observed in the portion of the pteroylGlu n pool having the long chain Glu n residues.
  • the ⁇ M quantities of folinate or substituted benzoates applied to plants in our studies are far greater than the nanomolar quantities normally found in foliage and should provide adequate stores for cell cycles even in the face of limited penetration.
  • serine hydroxymethyltransf erase may he activated to bind the input, thus, necessitating inputs aimed at lengthening the Glu n chain.
  • the C 3 cycle might be diverted from the GOGAT cycle. Specifically, decarboxylation of glyoxylate has been shown (for example Grodzinski 1978; Zelitch 1972) where glycolate from the C 3 cycle can he broken down to carbon dioxide and formate.
  • Ci-THF methyltetrahydrofolate
  • Cossins E. A. 1980. One-Carbon Metabolism. In The Biochemistry of Plants, v. 2, P. K. Stumph & E. E. Conn, Eds., Academic Press, NY, Pp.365-4 18. Cossins, E. A. 1987. Folate Biochemistry and the Metabolism of One-Carbon Units.

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Abstract

La présente invention concerne des procédés et des compositions permettant d'améliorer la croissance des plantes par application foliaire d'une substance qui améliore l'accumulation du formyltetrahydroptéroyle polyglutamate (C1-THF) dans une plante traitée. Le traitement avec des substances qui contribuent à la structure du C1-THF accroissent le taux et la quantité de fixation de carbone par la plante. Subséquemment, la croissance des plantes est davantage améliorée par l'exposition de la plante à un degré élevé d'oxygène, d'éclairement et de chaleur ou par apport foliaire de sources de fragments de carbone simple. On obtient des résultats optimaux par la combinaison du traitement avec une substance servant de puits aux fragments de C1 produits dans la feuille.
PCT/US2001/003155 2000-02-02 2001-01-31 Procedes et compositions permettant d'ameliorer le formyltetrahydropteroylpolyglutamate chez les plantes WO2001056385A1 (fr)

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WO2002064881A1 (fr) * 2001-02-13 2002-08-22 Clariant International Ltd Protection de colorants sensibles a la reduction
EP1854348A1 (fr) * 2005-03-04 2007-11-14 Suntory Limited Plante transformee capable de produire de l'acide polyglutamique
CN107047028A (zh) * 2017-05-10 2017-08-18 蚌埠王恒亮盆景艺术有限公司 一种利用酸性溶液胁迫提高盆景三角枫光合生理特性的方法
RU2685477C1 (ru) * 2018-04-28 2019-04-18 Федеральное государственное бюджетное образовательное учреждение высшего образования "Горский государственный аграрный университет" Способ подкормки озимой пшеницы

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WO2002064881A1 (fr) * 2001-02-13 2002-08-22 Clariant International Ltd Protection de colorants sensibles a la reduction
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