CA2119806A1 - Seed conditioning process providing stress resistance - Google Patents
Seed conditioning process providing stress resistanceInfo
- Publication number
- CA2119806A1 CA2119806A1 CA002119806A CA2119806A CA2119806A1 CA 2119806 A1 CA2119806 A1 CA 2119806A1 CA 002119806 A CA002119806 A CA 002119806A CA 2119806 A CA2119806 A CA 2119806A CA 2119806 A1 CA2119806 A1 CA 2119806A1
- Authority
- CA
- Canada
- Prior art keywords
- seed
- triazole
- plant
- triazol
- dimethyl
- Prior art date
- Legal status (The legal status is an assumption and is not a legal conclusion. Google has not performed a legal analysis and makes no representation as to the accuracy of the status listed.)
- Abandoned
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Classifications
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- A—HUMAN NECESSITIES
- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
- A01C—PLANTING; SOWING; FERTILISING
- A01C1/00—Apparatus, or methods of use thereof, for testing or treating seed, roots, or the like, prior to sowing or planting
- A01C1/06—Coating or dressing seed
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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
- A01N25/00—Biocides, pest repellants or attractants, or plant growth regulators, characterised by their forms, or by their non-active ingredients or by their methods of application, e.g. seed treatment or sequential application; Substances for reducing the noxious effect of the active ingredients to organisms other than pests
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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
- 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
- A01N37/42—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 within the same carbon skeleton a carboxylic group or a thio analogue, or a derivative thereof, and a carbon atom having only two bonds to hetero atoms with at the most one bond to halogen, e.g. keto-carboxylic acids
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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
- A01N43/00—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds
- A01N43/48—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with two nitrogen atoms as the only ring hetero atoms
- A01N43/50—1,3-Diazoles; Hydrogenated 1,3-diazoles
-
- 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
- A01N43/64—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with three nitrogen atoms as the only ring hetero atoms
- A01N43/647—Triazoles; Hydrogenated triazoles
- A01N43/653—1,2,4-Triazoles; Hydrogenated 1,2,4-triazoles
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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
- A01N53/00—Biocides, pest repellants or attractants, or plant growth regulators containing cyclopropane carboxylic acids or derivatives thereof
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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
- 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
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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
- A01N2300/00—Combinations or mixtures of active ingredients covered by classes A01N27/00 - A01N65/48 with other active or formulation relevant ingredients, e.g. specific carrier materials or surfactants, covered by classes A01N25/00 - A01N65/48
Landscapes
- Life Sciences & Earth Sciences (AREA)
- Health & Medical Sciences (AREA)
- Environmental Sciences (AREA)
- General Health & Medical Sciences (AREA)
- Wood Science & Technology (AREA)
- Plant Pathology (AREA)
- Engineering & Computer Science (AREA)
- Dentistry (AREA)
- Pest Control & Pesticides (AREA)
- Zoology (AREA)
- Agronomy & Crop Science (AREA)
- Toxicology (AREA)
- Soil Sciences (AREA)
- Agricultural Chemicals And Associated Chemicals (AREA)
Abstract
Seeds are conditioned by a process which provides protection against environmental stress in the resulting plants. The process involves treatment of the seeds with a solution which comprises a compound or mixture of compounds, such as triazoles, and a source of potassium, which solution is capable of inhibiting ergosterol and gibberrellin biosyntheisis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene.
Description
SEED CONDITIONING PROCESS PROVIDING STRESS RESISTANCE
FIELD OF THE INVENTION
Seeds are conditioned by a process which provides protection against 5 environmental stress in the resulting plants.
BACKGROUND OF THE INVENTION
Triazole compounds are known fungicides used to control fungal diseases in plants and ~nim~l~ and are also known to be plant growth regulating compounds (Fletcher R.A., Hofstra G., Gao, J. (1986) Plant Cell Physiol.
10 27:367). The mechanism of triazole regulation blocks sterol, specifically ergoslerol, and gibberellin biosynthesis (Buchenauer H., Rohner E. (1981) Pesticide Biochem. and Physiol. 15:58), and causes a transient rise in abscisic acid (Mackay C.E., Hall J.C., Hofstra G., Fletcher R.A. (1990) Pesticide Biochem and Physiol. 37:74) and an increase in cytokinins (Fletcher R.A., Arnold V. (1986) Physiologia Plantarum 66:197). Ergosterol is formed by the isoprenoid pathway which is present in many org~ni~m~ and generates many metabolically important compounds (Betyia E.D., Porter J.W. (1976) Annu.
Rev. Biochem. 45:113). The isoprenoid pathway also produces other important plant growth hormones such as cytokinins and abscisic acid. It appeals that the plant growth regulating pr~ellies of the triazoles are mP~ ted by hll~lrerence with the isoprenoid pathway leading to a shift in growth regulator balance of such hormones as gibberellins, abscisic acid and cytokinins (Fletcher R.A., Hofstra G. (1985) Plant and Cell Physiol. 26:775). Plants treated with triazole have increased levels of abscisic acid (Mackay C.E., Hall J.C., Hofstra G., Fletcher R.A. (1990) Pesticide Biochem. and Physiol. 37:74). It has been suggested that triazoles reduce shoot elongation through inhibition of cytochrome P-450-dependent oxidations during gibberellin biosynthesis (Rademacher W. et al (1987) Pestic Sci 21 :241). Potassium has also been used in the treatment of seeds (Fletcher, R.A. et al. (1982) Plant Physiol.
69:675).
Both seeds and plants have been treated with triazoles to provide protection against fungal ~ e~es (European Patent 0 095 242). Seed dressings and plant treatments cont~ining triazole have also been used to regulate growth in plants (US Patent 4,931,082, C~n~ n Patent 1,194,882 and European Patent 0 099 165). Seedling treatment has inhibited heat stress-in~ ce~l ethylene production in wheat and soybean see-lling~ (Kraus, T.E., Murr, D.P., Fletcher R.A. (1991) J. Plant Growth Regul. 10:229). The decrease in total chlorophyll in bean leaves after heat and chilling treatment of plants is prevented by root drench treatment with a triazole (Asare-Boamah N.K., Fletcher R.A. (1986) Physiologia Plantarum 67:353). Root application of a triazole reduces ~l~nspi~ation, protects plants, and increases yield under waterstress conditions (Fletcher R.A., Nath V. (1984) Physiol. Plant 62:422, Asare-Boamah N.K., Hofstra G., Fletcher R.A., Dumbroff E.B. (1986) Plant Cell Physiol. 27:383). Soil drench treatments or foliar sprays of triazole protect plants against ozone and chilling (Fletcher, R.A., Hofstra G. (1985) Plant Cell Physiol. 26:775).
Seed dressing lleaL,llelll with triazole provides protection against ozone treatment (Mackay C.E., Senaratna T., McKersie B.D., Fletcher R.A. (1987) Plant Cell Physiol. 28:1271). The seed triazole application increased the endogenous concentrations of antioxidants in the cellular membranes. Seedling triazole treatment results in an increase in total lipid soluble antioxidants, alpha-tocopherol and ascorbic acid levels in leaves (Senaratna T., Mackay C.E., McKersie B.D., Fletcher R.A. (1988) J. Plant Physiol. 133:56). Seeds treated with triazole had see~llin~s which sere more resistant to heat stress (Booker H.M., Gillespie T.J., Hofstra G., Fletcher R.A. (1991) Physiologia Plantarum 81:335). See~llin~ of seeds treated with a triazole, were more resistant to water-stress and subsequent heat stress. The degree of protection against heat shock was enh~nre~l if the seellin~ were subjected to water-stress (24 days) prior to the heat shock (Fletcher R.A., ~S~nt~h~m~ri M., Murr D.P. (1988) Physiologia Plantarum 74:360).
Thus seed and plant treatments provide stress resistance to various environmental stresses with soil treatment being preferred to foliar spray.
However, in another expe~ lelll seed dressing treatment with a triazole proved mc~ccessful in protecting against cold stress in tomato seeds (Davis, T.D., Ells, J.E., Walser, R.H. (1990) HortScience 25:312).
Plants treated with triazole are shorter and more compact with thicker and darker green leaves with a significantly higher content of chlorophyll, 5 carotenes, xanthophylls and nucleic acids. Generally growth is inhibited in triazole treated plants, however, there are a few reports that triadimefon actually stim~ tes growth of some species (Kolbe W. (1981) Pflanzenschutz Machrichten Bayer 34:213; Fletcher R.A., Nath V. (1984) Physiol. Plant.
62:422).
The present invention is a novel seed treatment procedure which conditions a seed with triazole(s) or other similarly active compound. Plants from seeds treated by the process of the present invention are rendered resistant to environmental stresses.
The present invention provides a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to the plant derived from the conditioned seed.
The plant seed conditioning process comprises:
i) cont~cting the plant seed with a solution cont~ining a compound or a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and a carrier for the compound or ure thereof for delivering the compound mixture through a seed coat of the seed and into the seed, ii) the solution including a source of potassium at a concentration sufficient to enhance the effects of cytokinins in the seed.
According to a preferred aspect of the invention, the source of potassium is selected from the group which includes KNO3, KCl and K2SO4.
According to an aspect of the invention, a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to said plant derived from said conditioned seed, said plant seed conditioning process comprising:
i) cont~cting the plant seed for less than 2 minutes with a solution cont~ining a mixture of compounds capable of inhibiting ergosterol and 5 gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and an organic solvent carrier for the mixture for delivering the compound mixture through a seed coat of the seed and into the seed;
ii) the solution including a source of potassium at a concentration 10 sufficient to enhance the effects of cytokinins in the seed.
According to an aspect of the invention, a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to the plant derived from the conditioned seed, the plant seed conditioning process comprises:
i) cont~cting the plant seed for an extended period in the range of 2 to 18 hours with a solution cont~ining a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants with water for delivering the colnl)ou"d llli~LUle through a seed coat of the seed 0 and into the seed, ii) the solution including a source of pot~sil-m at a concentration sufficient to enhance the effects of cytokinins in the seed, and iii) a hardening step which preferably comprises subjecting the seed to an elevated temp~,~ture in the range of 35C to 45C for at least 1 hour to 25 enhance stress resistance in a plant resulting from the conditioned seed. Thesolution preferably contains a mixture of triadimefon, paclobutrazol and ancymidol. A cereal seed may conditioned in the solution for an extended period in the range of 16 to 18 hours. A soybean or canola seed may be conditioned in the solution for a shorter period in the range of 2 to 4 hours.
2119~0~
BRIEF DESCRIPTION OF THE DRAWINGS
Figure 1 is the isoprenoid pathway in-licating the sites of inhibition and increased production due to seed treatment Figure 2a compares the harvest weights of c~ ll,hels from dirrelellt 5 seed treatments.
Figure 2b compales the cl~m~ tive number of cucumbers harvested from dirre~ seed treatments.
DETAILED DESCRIPTIONS OF THE PREFERRED EMBODIMENTS
The present invention is a process for enhancing stress resistance in a 10 plant from the embryonic stage to maturity by conditioning a plant seed by contacting the plant seed with a solution cont~ining compounds capable of inhibiting ergosterol and gibberellin biosynthesis in plants, increasing production of abscisic acid and cytokinins and decreasing production of ethylene. This plant growth regulatory activity is explained below. Such 15 compounds include triazoles such as, paclobutrazol, triadimefon and other similarly active compounds, such as, ancymidol (a pyrmidine). The protection provided by this solution is increased by the addition of potassium which enhances the effects of cytokinins in the seed.
Although triazoles and other compounds have been used in treating 20 seeds, see~llingc and plants, the present invention provides an improved process for the LleaL.l~lll of seeds which leads to greatly enh~ncecl stress ~esis~lce.
The first embodiment of the process involves a short term/pulse treatment of seeds with triazole or a similarly active compound and preferably in combination with pot~sil-m and acetone. The second embodiment of the 25 process involves imbibing the seed with triazole or a similarly active compound in water, followed by a hardening step. Both processes provide protection from a variety of ellvilolllllental stresses, however the second process is moreeffective while the first process is easier to pelÇollll on a large commercial scale. Protection of plants from appa~elllly unrelated stresses leads to the 30 conclusion that the triazoles or other active compounds when used in accordance with the process of this invention either reduce free radical production or increase the antioxidant potential.
In the first process, the triazole compound or a similarly active compound or a mixture of compounds in solution with potassium and acetone is 5 shaken with the seeds for less than 2 minlltes. The amount of solution is measured so that it will be soaked into the seeds in the allotted time and it isnot drained off. The treated seeds are then air dried.
Initial trials with wheat seeds showed that a higher than 15%
concentration of acetone in water inhibited gell"illation, whereas pure acetone 10 had no effect on ge"nillation and growth of treated seeds. Initial trials also demonstrated that 120 mM of potassium given as 60 mM of KCl and 60 mM of KNO3 in 10% aqueous acetone gave faster germination and higher growth rates of treated wheat seeds. Hence, the triazoles were dissolved in 10% aqueous acetone cont~ining 120 mM of Pot~sil-m given as 60 mM of KCl and 60 mM
15 of KNO3, at the desired concentrations for seed treatment. Seeds were treatedfor short duration or given a pulse treatment for one minute and then air dried at room temperature (22-23C) overnight.
Acetone is lipophilic and solubilizes portions of the seed coating allowing the triazole or similarly active compound to penetrate into the seed.
20 This penetration is greater than conventional seed coating treatments.
Other solvents besides acetone may be used to aid the triazole or other similarly active compound in penetrating into the seed. The general group of solvents are ketones and alcohols with acetone and ethanol being prefel,ed.
Pot~c~inm has been found to enhance the stress protection and the source of 25 potassium may be selected from KNO3, KCl and K2SO4. The pulse treatment exposes the seeds to the active solution for less than 2 mimltes and preferably 1 minute. The speed of this technique and its simplicit,v make it particularly applicable to large scale commercial use.
In the second process, seeds are imbibed in triazole or similarly active 30 compound with pot~sinm at room temperature for 2 to 18 hours. The seeds are given a hardening treatment which consists of raising the temperature to 35C to 45C for at least 1 hour. The seeds are then air dried.
The potassium in this imbibing solution is preferably a 60 mM
concentration of both KCI and KNO3. However alternative sources of 5 potassium may be utilized such as K2SO4, and also the concentration may be varied. The hardening step is preferably carried out at a temperature of 40C
for two hours, that is the last two hours of the imbibing process. The seeds arepreferably air dried overnight at 22-23C. Dirrerellt species of plants require alterations in the imbibing method. Cereals like barley, wheat and corn require 10 imbibing for 16 to 18 hours whereas seeds like canola and soybean require imbibing for 2 to 4 hours because they tend to split.
The pulse technique requires higher concentrations of the triazole or similarly active compound than the imbibing technique. The imbibing technique provides enh~nred stress protection compared to the pulse treatment 15 since the imbibing solution penetrates right into the core of the seed. It isbelieved that this affects the seed at the level of transcription. In fact treated seeds can be stored for long periods of time and plants from treated seeds stored for one year have greater stress protection than those from freshly treated seeds. The combination of the triazole or other similarly active 20 compound, pot~sil-m and hardening produces a progl~"~",r~l seed that is able to withstand subsequent stress from the early embryonic stage to maturity.
The triazoles and/or other compounds either individually or in ad~ tule used to treat the seeds with either process may be selected from the following list:
Chemical Name Trade Name Common Name Company 1-(4-chlorophenoxy-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butanone Bayleton Triadimefon Bayer I -(4-chlorophenoxy}3,3-dimethyl- 1-( I H- I ,2,4-triazol- 1 -yl)butan-2-ol Baytan Triadimenol Bayer (E)-l-cyclohexyl-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol Baronet Triapenthenol Bayer S all-rac-l-(biphenyl-4-yloxy)-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol Baycor Bitertanol Bayer 1-(3-trifluoromethyltrityl}l H-1,2,4-triazole Persulon Fluotrimazole Bayer (+} I -[2-(2,4-dichlorophenyl}4-propyl- 1 ,3-dioxalan-2-ylmethyl]- I H- I ,2,4-triazole Tilt Propiconazole Ciba-Geigy (+}1-[2-(2,4-dichlorophenyl}4-ethyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4-triazole Sonax Etaconazole Ciba-Geigy 1-(2,4-dichloro-,B propylphenethyl}l H-1,2,4-triazole Topas Penconazole Ciba-Geigy 2-(2,4-dichlorophenyl}2-hydroxy-3-methoxy-3-(1 H-1,2,4-triazol-1-yl)propane ------ BAS 110 BASF
l-phenoxy-3-(1 H-1,2,4-triazol-1-yl)4-hydroxy-5,5-dimethyl-hexane ------ BAS 111 BASF
I -(4-trifluormethyl)-2-( 1 ,2,4-triazolyl-[ 1]-3 -(S-methyl- 1,3 -dioxan-5-yl-propen-3 -ol ------ LAB 150 978 BASF
(2RS,3RS)-1-(4-chlorophenyl}4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol Clipper Paclobutrazol ICI
(2RS,3RS} 1 -(2,4-Dichlorophenyl}4,4-dimethyl-2-( I H- I ,2,4-triazol- 1 -yl)pentan-3-ol Vigil Diclobutrazol ICI
(E)- I -(p-chlorophenyl)-4,4-dimethyl-2-( 1 ,2,4-triazol- 1 -yl} 1 -penten-3 -ol Sumagic Uniconazole Sumitomo (E}1-(2,4-Dichlorophenyl}4,4-dimethyl-2-(1,2,4-triazol}l-yl-1-penten-3-ol Spotless Diconazole Sumitomo a-butyl-a-(4-chlo~opl ~"lyl}l H-1,2,4-triazole-1-propanenitrile Systhane Myclobutanil Rohn & Haas oo o Additionally, other similarly active compounds include ancymidol (a pyrmidine), tetcyclacis (a norbornano~ 7etine) and Hoe 074784 (an imidazole).
Chemical Name Trade Name Common Name Company a-cylcopropyl-a(4-methoxy-phenyl)5-pyrimidine methanol A-Rest Ancymidol Dow-Elanco 5-(4-chlo,ophc.lyl~3,4,5,9,10-p~-~ -tetra-cylco-4,5,1,026,08 "-dodeca-3,9-diene Kenbyo Tetcyclacis BASF
1-(2,6-diethylphenyl)-imidazole-5-carboxamide ---- Hoe 074784 Hoechst All of the above compounds have plant growth regulatory activity and a structural element in common: the lone pair of electrons on the sp2-hybridized nitrogen atom in the heterocycle. The basic ring structures for Triazole, Ancymidol, Tetcyclacis and Hoe 074784 are as follows:
R
N R
~N~ N~1N~
Triazole Ancymidol N = N~3 N ~3 Tetcyclacis Hoe 074784 20 wherein R is a substituent which, when present on the respective cyclic structure does not int~,rele with the inhibition of ergosterol and gibberellin biosynthesis, increased production of abscisic acid and cytokinins and decreasedproduction of ethylene.
The plant growth regulatory activity demonstrated by tre~trnent with the 25 triazoles or other similarly active compounds is due to the effect of these compounds on the isoprenoid pathway. Compared to the prior art processes which only coat the seed, the process of this invention ensures that the active compound(s) penetrate the seed coating and enter the seed core material to alterthe seed isoprenoid pathway. As demonstrated by the examples, this 30 penetration has not been accomplished by the prior art process.
21I98Q~
The isoprenoid pathway explains the generation of animal, fungal, plant and insect hormones as well as other important metabolites, such as, Vitarnin A, E, and phytoallexins. This pathway is diagramed in Figure 1 and is labelled to indicate the points of inhibition (I) of ergosterol and gibberellin biosynthesis and increased production (P) of cytokinins and abscisic acid in plants due to the action of the triazoles or similarly active compounds. This diagram is simplified, for in~t~n~e, there are more steps involved in converting sqaulene to ergosterol (Siegel, M. (1981) 65 Plant Disease 986) and further steps required to convert geranylgeranyl-PP to gibberellins (Rademachen, W. (1989) Gibberellins: Metabolic pathways and inhibitors of biosynthesis, in Target Sites of Herbicide Action, eds. P. Boges, G. .S~n~lm~n, CRC Press Inc., Boco Raton). The inhibition of ergosterol and gibberellin biosynthesis and the transient rise in abscisic acid and increase in cytokinin levels caused by the invention's treatment methods act to protect the plant from envilomllental stresses, such as, drought, heat and cold. Another effect which is not within the isoprenoid pathway is the decreased production of ethylene. Ethylene inhibits deteriorative processes in the plant and affects its post-harvest physiology. The invention's seed treatment methods m~ximi7e the plant's resistance to envi,omllental stresses and "~ "~ fruit quality.
Uniconazole is the most effective triazole tested. The second most effective triazole for the present invention is paclobutrazol. The efficiency ofpaclobutrazol is usually improved in combination with other triazoles such as triadimefon and propiconazole and with the pyrrnidine, ancymidol. In conl~alison, triadimefon has to be used at a much higher concentration than paclobutrazol and is still not as effective.
The present invention has been particularly successful in providing stress protection for the following species of plants:
Cereal Oil Vegetable Misc.
Wheat Canola Tomato Corn Barley Mustard Potato Tuber Sorghulll Rice Peanut Potato Seed Pearl Millet Castor Cucurbit Cotton Safflower Eggplant Sesame Onion The examples demonstrate that treatment with the triazole or similarly 10 active compounds and potassium not only provide increased resistance to stresses, such as, heat, drought, cold and ~i~e~es, but also provide increased yields of crops grown in the field.
EXAMPLES
Example I
The desired concentration of triazole measured in parts per million (ppm) in a solution of 60 mM KCl and 60 mM KNO3 in 10~ aqueous acetone was used to treat the Katepwa cultivar wheat seeds (cv. Katepwa) for 15 minutes. (Table 1). The control seeds were treated only with water. The seeds were then air dried at room temperature (22-23C overnight). The 20 following abbreviations will be used in the Examples:
K KCL + KNO3 (60 mM each) T - Triadimefon U - Uniconazole P - Paclobutrazol An - Ancymidol Tl - Propiconazole.
The effect on coleoptile (Col) and root lengths was studied, after 30germin~ting the seeds in petri dishes at 22-23C in the dark for 4 days. The data on see-lling length, fresh and dry weights per unit of see~lling length as well as the effects from heat (48C for 3 hrs) and drought (14 days of drought) following recovery after 24 hrs of treatment or watering, were recorded on 10-14 days old see-lling~ grown in pot-mix in a glasshouse at 22-23C and 16 hrs photoperiod. Heat and drought damage to the see-lling~ were scored on a 0-10 S scale, 0 showing no damage and 10 for dead see~lling~. Data were statistically analyzed using COSTAT software and subjected to Duncan's Multiple Range Test. Means within in the columns which are followed by the same letter are not significantly dirrelellt at P=0.05 according to Duncan's Multiple Range Test. Uniconazole was most effective at 100 ppm given for 15 minutes followed by 100 ppm of P and least effective with 1000 ppm of T.
Example II
The Katepwa cultivar wheat seeds were treated with the in~irate~l concentration of triazole singly or combined with 60 mM KCL, 60 mM KNO3 and 10% acetone by a 1 minute pulse treatment. The seeds were shaken in 1 ml of solution per gram of seeds and then dried at room temperature (22-23C) overnight. The control seeds were treated with water only. The seeds were germin~ted and treated as in Example I. The results are shown in Table 2.
Treatment #2 in Table 2 is the tleal,llellt with only the 60 mM KCl, 60 mM
KNO3 and 10% acetone.
All the three chemicals combined gave high protection against heat with the see-1lin~s ~urr~ lg very little damage (<8%). There is a high positive correlation (0.913 _ 0.062) between fresh weight (F.W.) mg/cm of see~lling and heat tolerance.
Example III
Four cultivars of wheat, two Spring-Katepwa, Celtic and two winter-Karena, Ruby were pulse treated with a combination of 1000 ppm T, 100 ppm U and 100 ppm P. Combined treatment gave protection against heat and drought in all the four wheat varieties. (Table 3).
Example IV
Katepwa wheat cultivars were pulse treated with various concentrations of P and P plus T. A concentration of 200 ppm of P in 120 mM K (KCl +
KNO3) in 10% acetone given as pulse gave the highest fresh wt/cm and drought protection, whereas 1000 T plus 300 P gave the overall best combined fresh weight/cm and protection from heat and drought (Table 4). Both treatments gave a height reduction of about 50% in 14 days growth in a glass house.
S Example V
Katepwa wheat cultivars were pulse treated with A in combination with P+T (Table 5). Ancymidol alone did note provide any protection, but in combination with T+P gave 70% protection against heat and 50% against drought.
A concentration of 200, 300 and 400 ppm of P provided 50% protection against drought and 300 ppm provided maximum protection against heat with less than 40% damage.
Example VI
Katepwa wheat cultivars were pulse treated singly and in combination with A, T and P in 60 mM K(KNO3 and KCl) and 10% aceone (Table 6).
1000 T + 300 P provided the highest protection against heat and drought with the highest fresh weight (mg/cm). This treatment however, gave curving and distortion to the first leaf. The addition of 100 An elimin~te~ this distortion and provided similar protection against heat and drought. Comparison of Treatment 8 and Treatment 9 will show that addition of 1000 T provides protection against drought whereas lleaL~llellt with only P and An does not.
Example VII
Katepwa wheat cultivars were pulse treated with combinations of T, A, P and Tl in 60 mM K(KN03 and KCl) and 10% acetone (Table 7). Combined treatment widl 1000 T + 100 An + 100 P + 100 Tl provided maximum protection against heat and drought. Degree of damage from heat and drought showed the highest correlation to FW (mg/cm) and dry weight (mgxlO/cm), in-lic~ting that fresh and dry weight of see~lin~ are in~ir~tive of their ability to withstand heat and drought stresses.
Example VIII
Katewpa wheat cultivars were pulse treated with a combination of T, A
and P (TAP) or Tl, A and P (PAP) and the results compared (Table 8).
Both combinations, TAP and PAP, gave protection against heat and 5 drought, but PAP was slightly superior to TAP in protection as well as in increasing the fresh and dry weight of the see~llin~. The height of the see-lling was reduced to 1/3 by PAP whereas TAP gave reduction of about half of that of the control.
Example IX
Katepwa wheat cultivars were pulse treated with P and the see~lings immediately grown (Freshly treated) or first stored for 1 month.
Paclobutrazol (P) alone at 300 ppm provides protection to see~lling~ from heat showing damage <40% when freshly treated seeds are sown, but see-lling~ raised from one month stored treated seeds showed < 17% damage from the heat stress. However, P along does not provide much protection from drought when two week old seel1ling~ are subjected to stress (Table 9).
Example X
Leger cultivar barley seeds were pulse treated with various concentrations of T, A, P and Tl (Tables 10 and 11).
An initial trial with 1000 T + 100 An + 100 P showed that higher dose can inhibit ge~ ion. Hence, half and 1/4 doses were tried. Although gelll~ ation and emergence were not affected, the 1~2 dose (500 T + 50 An +
50 P) slightly delayed emergence (by 2 days) and the first leaf was curved and distorted. However, at lt2 dose, the protection to heat and drought stresses washighest showing damage to heat less than 7% and the drought damage less than 11%. Whereas at lh dose, the heat damage was less than 22% and drought damage was 68% (Table 10).
The treatment combination of 100 Tl + 25 An + 50 P gave the best protection against heat and drought, although all the three treatment combinations gave good protection to heat compared to control (Table 11).
. .
Example XI
Katepwa cultivar wheat seeds were imbibed for 18 hours in a combination of paclobutrazol, ancymidol and triadimefon (PAT)(PAT at 50:50:100 mg/L) with or without KCl (120 mM). The control was imbibed in S water for 18 hours. The ten day old wheat see~lling~ were exposed to a heat stress of 50C for 4 hours. ~.e~kin~ss was measured 1 day and survival two weeks after exposure to heat stress (Table 12). The higher percent le~kin~ss in the controls indic~tes loss of membrane integrity leading to lower survival rates. The treatment of PAT in the presence of KCl provided maximum protection. The appearance of the treated (P = 50 mg/L, P:T = 50:100 mg/L, PAT = 50:50:100 mg/L) versus untreated plants (water) demonstrates that the treated plants sustained minim~l damage, especially when treated with PAT, whereas the unllea~ed plant leaves curled and wilted.
The ten day old wheat see~lling~ were placed in a progr~mmed freezer and cooled at the rate of 2C/hr and survival was assessed one week after exposure to the cold (Table 13). The treatment of PAT and KCl provided m~ximllm protection similar to the results obtained with heat stress.
Water was witheld for 3, 6, or 9 days from 7 day old wheat see~linp~
and the amount of water llallspiled per plant measured. The injury was assess on day 9 on a visual rating of 0 - 10 with 10 being the most damaged and dessic~ted (Table 14).
Example XII
Tomato seeds were imbibed with PAT (25:25:100 mg/L) as in Example XI. The see~ling were exposed to heat stress of 50C for three hours. Again, the triazole treatment protected the tomato seeAling whereas the control was almost completely destroyed.
Example XIII
Soybean seeds were imbibed for 2 hours in water (control) and increasing concentrations of paclobutrazol of 5, 15 and 30 mg/L plus KCl (120mM). The three week old see~llings were subjected to drought by withholding water for 12 days. The control see~llin~ was totally dessicated -whereas the see~llinp~ treated with various concentrations of paclobutrazol and KCL were shorter and protected.
Example XIV
Katepwa cultivar wheat seeds were imbibed with triazole - 1 (PAT =
25:25:100 mg/L) and Triazole - 2 (PAT = 50:50:100 mg/L). The seeds were placed in moistened filter paper in a cold (4C) room for 10 days and then returned to room temperature. Germination was assessed after seven days (Table 15). The cold exposure reduced viability by approximately 50% and this effect was prevented by triazole treatement.
Example XV
Katepwa cultivar wheat seeds were imbibed in water or paclobutrazol (50 mg/L) for 18 hours at room temperature. They were transferred to petri dishes on moistened filter paper and exposed to 50C for between 0 to 2 hours and returned to room tem~)el~ture. Germination percentage was recorded after 3 days. After 1.5 hours of exposure to 50C, the gelll~-nation was 40% and 72% in the control and triazole treatment respectively (Table 16).
Example XVI
Loose smut infestation was compared in wheat seeds infested with loose smut and treated by imbibing with paclobutrazol (P = 25 mg/L), pulse treatment with paclobutrazol (P = 300 mg/L) or control. Treatment by imbibition completely eradicated loose smut and pulse treatment greatly decreased loose smut (Table 17).
Example XVII
Broccoli (Table 18, PAT Imbibition = 10:10:100 mg/L, pulse =
33:33:300), beans (Table 19, PAT imbibition = 10:10:100, pulse =
50:50:500); and corn seeds (Table 20, PAT imbibition = 10:10:100, pulse =
33:33:300) were treated with PAT by imbibition and by pulse, ge~ te~l in a greenhouse and then transplanted in the field at the two-leaf stage. In every case the yield was increased with PAT treatment. Generally the imbibition greatly increased the yield over the pulse and control. Although broccoli is cold tolerant, the control see~llingc tended to drop in the afternoon sun whereas the imbibed see-llingc were turgid.
Example XVIII
Katepwa and Celtic wheat cultivar seeds were treated with PAT by imbibition (PAT = 50:50:100) and pulse (PAT = 100:100:1000). The seeds were sown in the field and the height, lodging resistance, stalk breakage, and diseases of mildew, rust and septoria were measured (Table 21). The ranking scale was 0-9 with 9 being highest. Both triazole treatments initially reduced the height of plants, but just prior to harvest they were taller than controls.
The imbibition treated plants were more resistant to lodging and more effective than pulse in reducing diseases.
Spring barley Leger, and AC Burman cultivar seeds treated with PAT
(25:25:250 mg/L) by pulse treatment reduced powdery mildew, leaf rust and blotch diseases signifi~ntly (Table 22).
Example XIX
Soybeans were treated with PAT (Imbibition = 10:10:100, pulse =
33:33:300) by pulse and imbibition and sown in the field. Those sown in the summer (Table 23) demonstrated that the triazole treatment, especially imbibition, increased the height, number of pods, seeds per pod and seed weight of soybean plants.
Soybean seeds sown in early October, demonstrated that when the control leaves had started to turn yellow and senesce, the triazole-treated leaves from both pulse and imbibition treatments were still green, and the pods in these plants were larger than the controls.
Example XX
Cucumber seeds treated with PAT (10:10:100 mg/L) by imbibition treatment resulted in denser growth of the treated plants in the field due to protection from fost which inhibited the growth of the controls.
Cucumber seeds treated with PAT (Imbibition = 10:10:100, pulse =
33:33:300 mg/L) by imbibition and pulse were grown in the field and the harvests compared. The weights of cucumbers harvested per plot consisting of 2119~06 4 rows (each row planted with 10 sets of 3 seeds each) was significantly higher for the imbibition treated seeds whereas pulse was equal to control (Figure 2 a,b).
Example XXI
Tomato seeds were treated with PAT (25:25:100 mg/L)`by imbibition.
See-llin~ were gel",i"~ted in the growth room and transplanted at the 3-leaf stage. After a mild frost, control see~ling~ were damaged with necrosis of the leaf tips whereas the treated see~ling~ were protected.
Tomato seeds treated with PAT imbibition were grown in the field. The tomatoes harvested from control and treated plants were stored at 4C for 14 days. The nulllber of fruits that rotted was far greater in the control plants than the treated plants. Tomatoes harvested from control and treated plants were stored at 15C for 14 days. Again there were far more rotted tomatoes for the control plants.
Tomato seeds imbibed in water or PAT, Triazole - 1 (25:25:100 mg/L) and Triazole - 2 (50:50:100 mg/L),were gel",il-~te~ in the greenhouse and tomato see-lling~ were transplanted in the field. Tomatoes were harvested when they were mature green. The tomatoes were stored at 4C or 15C for 21 days.
Sucrose equivalent based on specific gravity showed that the treated fruits had signific~ntly higher levels than the controls (P = 0.001). The triazole treatment also signifi~ntly protected the fruits from damage during storage at both temperatures (Table 25).
Tomato seeds were treated with PAT (Imbibition 25:25:100, pulse 100:100:1000) by imbibition and pulse and disease inle~lsily measured in leafs of the tomato plants grown in the field. Disease intensity was significantly reduced in the triazole treated plants, more so with the imbibition (Table 26).
Harvests of the tomatoes demonstrated that triazole treatment promotes the yield of tomatoes. The yield was increased the most with the pulse treated seeds (Table 27).
~y~m~le XXII
Green pepper seeds were imbibed in water (control) or paclobutrazol (P
25 and 50 mg/l) in 40 mM KCI for 18 h and hardened at 40C for 2 h and air dried. Seetllinge were gel",in~ted in the greenhouse and one set of 5-week-old see~linge were transplanted early on May 10, 1993. A second set of 5-week-old see-llinge were planted on June 5, 1993.
The control see~lling~e from the first planting were damaged with necrosis of the leaf tips after a mild frost, whereas the treated see~llin~e were protected.
Thereafter there was no stress. The results reflect that the yield was increasedin the treated plants of the early planting due to protection from stress whereas in the late planting with no stress, the treatment had no effect.
Example XXIII
Potato (tubers) were dipped for 30 seconds in 10% acetone alone (control), or cont~ining paclobutrazol (P). They were allowed to air dry for 30 minutes and planted at Cambridge on May 5, 1993.
The increase in yield obtained with paclobutrazol at 0.15 mg/l is negated at the higher concentration tested. Although, the lllargi,lal 9% increase in yield for potatoes treated with 0.15 mg/L Paclobutrazol appears marginal, the growing season for summer of 1993 was particularly unstressful. Even in such conditions, the treatment provided a noticeable increase in yield. This example therefore demonstrates that potato tubers treated by this invention's method provides protection against stress.
211980i~
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Protecticn of 10-day Old Wheat See-lling~
from Ex~o~ to a Heat Stress of 50C for 4 h T e~kin~ss Survival Treatment (% Conductivity) (%) Control 71 45 KCl 64 57 PAT + KCl 30 100 Percent Survival of Wheat Seedlings After F,~ ..re to Low T~ ures % Survival Treatrnent (C) -6 -8 -10 Control 49 14 0 KCl 51 38 0 PAT + KCl 100 100 68 T~BL E 14 ~e~ ction of Tr~iralion and Protection of Wheat See~lling~ Against Drought Water Transpired g/plant Injury (0-10) Trea~nent (Days) 3 6 9 8 Control 4.8 11.1 19.1 6 K Cl 4.7 8.4 14.2 6 P A T 3.2 5.2 8.3 2 P A T + K Cl 3.2 4.1 8.2 Wheat Seeds (cv. Katepwa) Were Placed in Moi~..ed Filter Paper in Cold (4C) Room for 10 Days Then Returned to Room Tcm~.~ re. Ge~ ion was ~ ss~l After 7 days.
Treatment Germination %
Control 49 Triazole-1 94 Triazole-2 96 21I~806 -Control of Loose Smut in Wheat by Paclobutrazol (P) Treatment % Loose Smut Control 9 a Imbibition 0 b Pulse 0.9 b Field Data (Summer 1992) for Broccoli Grown in Cambridge, Ontario, Canada Treatment Yield/Plant (g) % Increase Control 183 a ---Imbibition 233 b 27 Pulse 178 a ---Field Data (Summer 1992) for Beans Grown in Arkell (1) and Cambridge - 1st Planting (2) and 2nd ~ h~r (3), Ontario, Csm~
(1) Treatment Yield/Plant (g) % Increase Control 167 a ---Imbibition 299 c 37 Pulse 198 b 19 (2) Treatment Yield/Plant (g) % Increase Control 45 a ---Imbibition 93 b 107 Pulse 40 a (-)12 (3) Treatment Yield/Plant (g) % Increase Control 44 a ---Imbibition 64 c 44 Pulse 55 b 25 Field Data (Summer 1992) for Corn Grown in Arkell, Ontario, C~(lq Treatment Yield/Plant (g) % Increase Control 159 a ---Imbibition 229 c 35 Pulse 178 b 24 2119~6 . ~
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c~ x ~oo oo ~ oo t - x oo . -~ ~ c y c 211g806 ~_' 37 Table 22 Triazole reduce diseases (powdery mildew, leaf rust, and blotch) in spring barley grown in Elora, Ontario (June 24, 1992) cv. Leger a) o O
cv. AC Burman ~ , .~ .
I I
Control Pulse T~BL E 23 Field Data (Summer 1992) for Soybeans Grown in Cambridge, Ontario, Canada Treatment Height (cm) Pod/Plant (No.) Seeds/Pod (No.) Seed wt. (g) Control 52.9 30.2 2.62 0.69 Imbibition 71.6 33.4 2.78 0.79 Pulse 66.1 37.6 2.78 0.79 oo o Table 24 ~__ 1~
~_ Shelf life of tomatoes halvested ~om control and treated (imbibed-PAT) plants. The tomatoes that were spoiled in each case are shown in blacl;. Atboth storage temperatures the triazole significantly protects spoilage.
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MIDDLE LEAVES
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WHOLE PLANT
Control Imbibition Pulse Table 2 7 Triazoles promote yield of tomato plants grown in Arkell, Ontario (Summer 1 992) v~ O Control V
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O -No. of Harvests 211~gO6 T~UBLE 28 FIELD DATA (SUl\Il\IER 1993) FOR GREEN ~ ;KS
GROWN AT CAMBRIDGE, ON TWO PLANTING DATES
s Treatment Early Planting Late Planting 10 mg/l yield/plant yield/plant (g) ~ Increase (g) Control 218 a - 212 a P-25 Imbibition 276 b 27 205 a P-50 Imbibition 290 b 33 201 a FIELD DATA (SUMMER 1993) FOR POTATOES
(VAR. KENEBEC) GROWN AT CAMBRIDGE
s Treatment Yield per 30 ft. row mg/l (lbs.) Control 45 a P0.15 49a P 0.25 41 c Although preferred embodiments of the invention are described herein in detail, it will be understood by those skilled in the art that variations may bemade thereto without departing from the spirit of the invention or the scope of 25 the appended claims.
FIELD OF THE INVENTION
Seeds are conditioned by a process which provides protection against 5 environmental stress in the resulting plants.
BACKGROUND OF THE INVENTION
Triazole compounds are known fungicides used to control fungal diseases in plants and ~nim~l~ and are also known to be plant growth regulating compounds (Fletcher R.A., Hofstra G., Gao, J. (1986) Plant Cell Physiol.
10 27:367). The mechanism of triazole regulation blocks sterol, specifically ergoslerol, and gibberellin biosynthesis (Buchenauer H., Rohner E. (1981) Pesticide Biochem. and Physiol. 15:58), and causes a transient rise in abscisic acid (Mackay C.E., Hall J.C., Hofstra G., Fletcher R.A. (1990) Pesticide Biochem and Physiol. 37:74) and an increase in cytokinins (Fletcher R.A., Arnold V. (1986) Physiologia Plantarum 66:197). Ergosterol is formed by the isoprenoid pathway which is present in many org~ni~m~ and generates many metabolically important compounds (Betyia E.D., Porter J.W. (1976) Annu.
Rev. Biochem. 45:113). The isoprenoid pathway also produces other important plant growth hormones such as cytokinins and abscisic acid. It appeals that the plant growth regulating pr~ellies of the triazoles are mP~ ted by hll~lrerence with the isoprenoid pathway leading to a shift in growth regulator balance of such hormones as gibberellins, abscisic acid and cytokinins (Fletcher R.A., Hofstra G. (1985) Plant and Cell Physiol. 26:775). Plants treated with triazole have increased levels of abscisic acid (Mackay C.E., Hall J.C., Hofstra G., Fletcher R.A. (1990) Pesticide Biochem. and Physiol. 37:74). It has been suggested that triazoles reduce shoot elongation through inhibition of cytochrome P-450-dependent oxidations during gibberellin biosynthesis (Rademacher W. et al (1987) Pestic Sci 21 :241). Potassium has also been used in the treatment of seeds (Fletcher, R.A. et al. (1982) Plant Physiol.
69:675).
Both seeds and plants have been treated with triazoles to provide protection against fungal ~ e~es (European Patent 0 095 242). Seed dressings and plant treatments cont~ining triazole have also been used to regulate growth in plants (US Patent 4,931,082, C~n~ n Patent 1,194,882 and European Patent 0 099 165). Seedling treatment has inhibited heat stress-in~ ce~l ethylene production in wheat and soybean see-lling~ (Kraus, T.E., Murr, D.P., Fletcher R.A. (1991) J. Plant Growth Regul. 10:229). The decrease in total chlorophyll in bean leaves after heat and chilling treatment of plants is prevented by root drench treatment with a triazole (Asare-Boamah N.K., Fletcher R.A. (1986) Physiologia Plantarum 67:353). Root application of a triazole reduces ~l~nspi~ation, protects plants, and increases yield under waterstress conditions (Fletcher R.A., Nath V. (1984) Physiol. Plant 62:422, Asare-Boamah N.K., Hofstra G., Fletcher R.A., Dumbroff E.B. (1986) Plant Cell Physiol. 27:383). Soil drench treatments or foliar sprays of triazole protect plants against ozone and chilling (Fletcher, R.A., Hofstra G. (1985) Plant Cell Physiol. 26:775).
Seed dressing lleaL,llelll with triazole provides protection against ozone treatment (Mackay C.E., Senaratna T., McKersie B.D., Fletcher R.A. (1987) Plant Cell Physiol. 28:1271). The seed triazole application increased the endogenous concentrations of antioxidants in the cellular membranes. Seedling triazole treatment results in an increase in total lipid soluble antioxidants, alpha-tocopherol and ascorbic acid levels in leaves (Senaratna T., Mackay C.E., McKersie B.D., Fletcher R.A. (1988) J. Plant Physiol. 133:56). Seeds treated with triazole had see~llin~s which sere more resistant to heat stress (Booker H.M., Gillespie T.J., Hofstra G., Fletcher R.A. (1991) Physiologia Plantarum 81:335). See~llin~ of seeds treated with a triazole, were more resistant to water-stress and subsequent heat stress. The degree of protection against heat shock was enh~nre~l if the seellin~ were subjected to water-stress (24 days) prior to the heat shock (Fletcher R.A., ~S~nt~h~m~ri M., Murr D.P. (1988) Physiologia Plantarum 74:360).
Thus seed and plant treatments provide stress resistance to various environmental stresses with soil treatment being preferred to foliar spray.
However, in another expe~ lelll seed dressing treatment with a triazole proved mc~ccessful in protecting against cold stress in tomato seeds (Davis, T.D., Ells, J.E., Walser, R.H. (1990) HortScience 25:312).
Plants treated with triazole are shorter and more compact with thicker and darker green leaves with a significantly higher content of chlorophyll, 5 carotenes, xanthophylls and nucleic acids. Generally growth is inhibited in triazole treated plants, however, there are a few reports that triadimefon actually stim~ tes growth of some species (Kolbe W. (1981) Pflanzenschutz Machrichten Bayer 34:213; Fletcher R.A., Nath V. (1984) Physiol. Plant.
62:422).
The present invention is a novel seed treatment procedure which conditions a seed with triazole(s) or other similarly active compound. Plants from seeds treated by the process of the present invention are rendered resistant to environmental stresses.
The present invention provides a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to the plant derived from the conditioned seed.
The plant seed conditioning process comprises:
i) cont~cting the plant seed with a solution cont~ining a compound or a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and a carrier for the compound or ure thereof for delivering the compound mixture through a seed coat of the seed and into the seed, ii) the solution including a source of potassium at a concentration sufficient to enhance the effects of cytokinins in the seed.
According to a preferred aspect of the invention, the source of potassium is selected from the group which includes KNO3, KCl and K2SO4.
According to an aspect of the invention, a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to said plant derived from said conditioned seed, said plant seed conditioning process comprising:
i) cont~cting the plant seed for less than 2 minutes with a solution cont~ining a mixture of compounds capable of inhibiting ergosterol and 5 gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and an organic solvent carrier for the mixture for delivering the compound mixture through a seed coat of the seed and into the seed;
ii) the solution including a source of potassium at a concentration 10 sufficient to enhance the effects of cytokinins in the seed.
According to an aspect of the invention, a process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to the plant derived from the conditioned seed, the plant seed conditioning process comprises:
i) cont~cting the plant seed for an extended period in the range of 2 to 18 hours with a solution cont~ining a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants with water for delivering the colnl)ou"d llli~LUle through a seed coat of the seed 0 and into the seed, ii) the solution including a source of pot~sil-m at a concentration sufficient to enhance the effects of cytokinins in the seed, and iii) a hardening step which preferably comprises subjecting the seed to an elevated temp~,~ture in the range of 35C to 45C for at least 1 hour to 25 enhance stress resistance in a plant resulting from the conditioned seed. Thesolution preferably contains a mixture of triadimefon, paclobutrazol and ancymidol. A cereal seed may conditioned in the solution for an extended period in the range of 16 to 18 hours. A soybean or canola seed may be conditioned in the solution for a shorter period in the range of 2 to 4 hours.
2119~0~
BRIEF DESCRIPTION OF THE DRAWINGS
Figure 1 is the isoprenoid pathway in-licating the sites of inhibition and increased production due to seed treatment Figure 2a compares the harvest weights of c~ ll,hels from dirrelellt 5 seed treatments.
Figure 2b compales the cl~m~ tive number of cucumbers harvested from dirre~ seed treatments.
DETAILED DESCRIPTIONS OF THE PREFERRED EMBODIMENTS
The present invention is a process for enhancing stress resistance in a 10 plant from the embryonic stage to maturity by conditioning a plant seed by contacting the plant seed with a solution cont~ining compounds capable of inhibiting ergosterol and gibberellin biosynthesis in plants, increasing production of abscisic acid and cytokinins and decreasing production of ethylene. This plant growth regulatory activity is explained below. Such 15 compounds include triazoles such as, paclobutrazol, triadimefon and other similarly active compounds, such as, ancymidol (a pyrmidine). The protection provided by this solution is increased by the addition of potassium which enhances the effects of cytokinins in the seed.
Although triazoles and other compounds have been used in treating 20 seeds, see~llingc and plants, the present invention provides an improved process for the LleaL.l~lll of seeds which leads to greatly enh~ncecl stress ~esis~lce.
The first embodiment of the process involves a short term/pulse treatment of seeds with triazole or a similarly active compound and preferably in combination with pot~sil-m and acetone. The second embodiment of the 25 process involves imbibing the seed with triazole or a similarly active compound in water, followed by a hardening step. Both processes provide protection from a variety of ellvilolllllental stresses, however the second process is moreeffective while the first process is easier to pelÇollll on a large commercial scale. Protection of plants from appa~elllly unrelated stresses leads to the 30 conclusion that the triazoles or other active compounds when used in accordance with the process of this invention either reduce free radical production or increase the antioxidant potential.
In the first process, the triazole compound or a similarly active compound or a mixture of compounds in solution with potassium and acetone is 5 shaken with the seeds for less than 2 minlltes. The amount of solution is measured so that it will be soaked into the seeds in the allotted time and it isnot drained off. The treated seeds are then air dried.
Initial trials with wheat seeds showed that a higher than 15%
concentration of acetone in water inhibited gell"illation, whereas pure acetone 10 had no effect on ge"nillation and growth of treated seeds. Initial trials also demonstrated that 120 mM of potassium given as 60 mM of KCl and 60 mM of KNO3 in 10% aqueous acetone gave faster germination and higher growth rates of treated wheat seeds. Hence, the triazoles were dissolved in 10% aqueous acetone cont~ining 120 mM of Pot~sil-m given as 60 mM of KCl and 60 mM
15 of KNO3, at the desired concentrations for seed treatment. Seeds were treatedfor short duration or given a pulse treatment for one minute and then air dried at room temperature (22-23C) overnight.
Acetone is lipophilic and solubilizes portions of the seed coating allowing the triazole or similarly active compound to penetrate into the seed.
20 This penetration is greater than conventional seed coating treatments.
Other solvents besides acetone may be used to aid the triazole or other similarly active compound in penetrating into the seed. The general group of solvents are ketones and alcohols with acetone and ethanol being prefel,ed.
Pot~c~inm has been found to enhance the stress protection and the source of 25 potassium may be selected from KNO3, KCl and K2SO4. The pulse treatment exposes the seeds to the active solution for less than 2 mimltes and preferably 1 minute. The speed of this technique and its simplicit,v make it particularly applicable to large scale commercial use.
In the second process, seeds are imbibed in triazole or similarly active 30 compound with pot~sinm at room temperature for 2 to 18 hours. The seeds are given a hardening treatment which consists of raising the temperature to 35C to 45C for at least 1 hour. The seeds are then air dried.
The potassium in this imbibing solution is preferably a 60 mM
concentration of both KCI and KNO3. However alternative sources of 5 potassium may be utilized such as K2SO4, and also the concentration may be varied. The hardening step is preferably carried out at a temperature of 40C
for two hours, that is the last two hours of the imbibing process. The seeds arepreferably air dried overnight at 22-23C. Dirrerellt species of plants require alterations in the imbibing method. Cereals like barley, wheat and corn require 10 imbibing for 16 to 18 hours whereas seeds like canola and soybean require imbibing for 2 to 4 hours because they tend to split.
The pulse technique requires higher concentrations of the triazole or similarly active compound than the imbibing technique. The imbibing technique provides enh~nred stress protection compared to the pulse treatment 15 since the imbibing solution penetrates right into the core of the seed. It isbelieved that this affects the seed at the level of transcription. In fact treated seeds can be stored for long periods of time and plants from treated seeds stored for one year have greater stress protection than those from freshly treated seeds. The combination of the triazole or other similarly active 20 compound, pot~sil-m and hardening produces a progl~"~",r~l seed that is able to withstand subsequent stress from the early embryonic stage to maturity.
The triazoles and/or other compounds either individually or in ad~ tule used to treat the seeds with either process may be selected from the following list:
Chemical Name Trade Name Common Name Company 1-(4-chlorophenoxy-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butanone Bayleton Triadimefon Bayer I -(4-chlorophenoxy}3,3-dimethyl- 1-( I H- I ,2,4-triazol- 1 -yl)butan-2-ol Baytan Triadimenol Bayer (E)-l-cyclohexyl-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol Baronet Triapenthenol Bayer S all-rac-l-(biphenyl-4-yloxy)-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol Baycor Bitertanol Bayer 1-(3-trifluoromethyltrityl}l H-1,2,4-triazole Persulon Fluotrimazole Bayer (+} I -[2-(2,4-dichlorophenyl}4-propyl- 1 ,3-dioxalan-2-ylmethyl]- I H- I ,2,4-triazole Tilt Propiconazole Ciba-Geigy (+}1-[2-(2,4-dichlorophenyl}4-ethyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4-triazole Sonax Etaconazole Ciba-Geigy 1-(2,4-dichloro-,B propylphenethyl}l H-1,2,4-triazole Topas Penconazole Ciba-Geigy 2-(2,4-dichlorophenyl}2-hydroxy-3-methoxy-3-(1 H-1,2,4-triazol-1-yl)propane ------ BAS 110 BASF
l-phenoxy-3-(1 H-1,2,4-triazol-1-yl)4-hydroxy-5,5-dimethyl-hexane ------ BAS 111 BASF
I -(4-trifluormethyl)-2-( 1 ,2,4-triazolyl-[ 1]-3 -(S-methyl- 1,3 -dioxan-5-yl-propen-3 -ol ------ LAB 150 978 BASF
(2RS,3RS)-1-(4-chlorophenyl}4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol Clipper Paclobutrazol ICI
(2RS,3RS} 1 -(2,4-Dichlorophenyl}4,4-dimethyl-2-( I H- I ,2,4-triazol- 1 -yl)pentan-3-ol Vigil Diclobutrazol ICI
(E)- I -(p-chlorophenyl)-4,4-dimethyl-2-( 1 ,2,4-triazol- 1 -yl} 1 -penten-3 -ol Sumagic Uniconazole Sumitomo (E}1-(2,4-Dichlorophenyl}4,4-dimethyl-2-(1,2,4-triazol}l-yl-1-penten-3-ol Spotless Diconazole Sumitomo a-butyl-a-(4-chlo~opl ~"lyl}l H-1,2,4-triazole-1-propanenitrile Systhane Myclobutanil Rohn & Haas oo o Additionally, other similarly active compounds include ancymidol (a pyrmidine), tetcyclacis (a norbornano~ 7etine) and Hoe 074784 (an imidazole).
Chemical Name Trade Name Common Name Company a-cylcopropyl-a(4-methoxy-phenyl)5-pyrimidine methanol A-Rest Ancymidol Dow-Elanco 5-(4-chlo,ophc.lyl~3,4,5,9,10-p~-~ -tetra-cylco-4,5,1,026,08 "-dodeca-3,9-diene Kenbyo Tetcyclacis BASF
1-(2,6-diethylphenyl)-imidazole-5-carboxamide ---- Hoe 074784 Hoechst All of the above compounds have plant growth regulatory activity and a structural element in common: the lone pair of electrons on the sp2-hybridized nitrogen atom in the heterocycle. The basic ring structures for Triazole, Ancymidol, Tetcyclacis and Hoe 074784 are as follows:
R
N R
~N~ N~1N~
Triazole Ancymidol N = N~3 N ~3 Tetcyclacis Hoe 074784 20 wherein R is a substituent which, when present on the respective cyclic structure does not int~,rele with the inhibition of ergosterol and gibberellin biosynthesis, increased production of abscisic acid and cytokinins and decreasedproduction of ethylene.
The plant growth regulatory activity demonstrated by tre~trnent with the 25 triazoles or other similarly active compounds is due to the effect of these compounds on the isoprenoid pathway. Compared to the prior art processes which only coat the seed, the process of this invention ensures that the active compound(s) penetrate the seed coating and enter the seed core material to alterthe seed isoprenoid pathway. As demonstrated by the examples, this 30 penetration has not been accomplished by the prior art process.
21I98Q~
The isoprenoid pathway explains the generation of animal, fungal, plant and insect hormones as well as other important metabolites, such as, Vitarnin A, E, and phytoallexins. This pathway is diagramed in Figure 1 and is labelled to indicate the points of inhibition (I) of ergosterol and gibberellin biosynthesis and increased production (P) of cytokinins and abscisic acid in plants due to the action of the triazoles or similarly active compounds. This diagram is simplified, for in~t~n~e, there are more steps involved in converting sqaulene to ergosterol (Siegel, M. (1981) 65 Plant Disease 986) and further steps required to convert geranylgeranyl-PP to gibberellins (Rademachen, W. (1989) Gibberellins: Metabolic pathways and inhibitors of biosynthesis, in Target Sites of Herbicide Action, eds. P. Boges, G. .S~n~lm~n, CRC Press Inc., Boco Raton). The inhibition of ergosterol and gibberellin biosynthesis and the transient rise in abscisic acid and increase in cytokinin levels caused by the invention's treatment methods act to protect the plant from envilomllental stresses, such as, drought, heat and cold. Another effect which is not within the isoprenoid pathway is the decreased production of ethylene. Ethylene inhibits deteriorative processes in the plant and affects its post-harvest physiology. The invention's seed treatment methods m~ximi7e the plant's resistance to envi,omllental stresses and "~ "~ fruit quality.
Uniconazole is the most effective triazole tested. The second most effective triazole for the present invention is paclobutrazol. The efficiency ofpaclobutrazol is usually improved in combination with other triazoles such as triadimefon and propiconazole and with the pyrrnidine, ancymidol. In conl~alison, triadimefon has to be used at a much higher concentration than paclobutrazol and is still not as effective.
The present invention has been particularly successful in providing stress protection for the following species of plants:
Cereal Oil Vegetable Misc.
Wheat Canola Tomato Corn Barley Mustard Potato Tuber Sorghulll Rice Peanut Potato Seed Pearl Millet Castor Cucurbit Cotton Safflower Eggplant Sesame Onion The examples demonstrate that treatment with the triazole or similarly 10 active compounds and potassium not only provide increased resistance to stresses, such as, heat, drought, cold and ~i~e~es, but also provide increased yields of crops grown in the field.
EXAMPLES
Example I
The desired concentration of triazole measured in parts per million (ppm) in a solution of 60 mM KCl and 60 mM KNO3 in 10~ aqueous acetone was used to treat the Katepwa cultivar wheat seeds (cv. Katepwa) for 15 minutes. (Table 1). The control seeds were treated only with water. The seeds were then air dried at room temperature (22-23C overnight). The 20 following abbreviations will be used in the Examples:
K KCL + KNO3 (60 mM each) T - Triadimefon U - Uniconazole P - Paclobutrazol An - Ancymidol Tl - Propiconazole.
The effect on coleoptile (Col) and root lengths was studied, after 30germin~ting the seeds in petri dishes at 22-23C in the dark for 4 days. The data on see-lling length, fresh and dry weights per unit of see~lling length as well as the effects from heat (48C for 3 hrs) and drought (14 days of drought) following recovery after 24 hrs of treatment or watering, were recorded on 10-14 days old see-lling~ grown in pot-mix in a glasshouse at 22-23C and 16 hrs photoperiod. Heat and drought damage to the see-lling~ were scored on a 0-10 S scale, 0 showing no damage and 10 for dead see~lling~. Data were statistically analyzed using COSTAT software and subjected to Duncan's Multiple Range Test. Means within in the columns which are followed by the same letter are not significantly dirrelellt at P=0.05 according to Duncan's Multiple Range Test. Uniconazole was most effective at 100 ppm given for 15 minutes followed by 100 ppm of P and least effective with 1000 ppm of T.
Example II
The Katepwa cultivar wheat seeds were treated with the in~irate~l concentration of triazole singly or combined with 60 mM KCL, 60 mM KNO3 and 10% acetone by a 1 minute pulse treatment. The seeds were shaken in 1 ml of solution per gram of seeds and then dried at room temperature (22-23C) overnight. The control seeds were treated with water only. The seeds were germin~ted and treated as in Example I. The results are shown in Table 2.
Treatment #2 in Table 2 is the tleal,llellt with only the 60 mM KCl, 60 mM
KNO3 and 10% acetone.
All the three chemicals combined gave high protection against heat with the see-1lin~s ~urr~ lg very little damage (<8%). There is a high positive correlation (0.913 _ 0.062) between fresh weight (F.W.) mg/cm of see~lling and heat tolerance.
Example III
Four cultivars of wheat, two Spring-Katepwa, Celtic and two winter-Karena, Ruby were pulse treated with a combination of 1000 ppm T, 100 ppm U and 100 ppm P. Combined treatment gave protection against heat and drought in all the four wheat varieties. (Table 3).
Example IV
Katepwa wheat cultivars were pulse treated with various concentrations of P and P plus T. A concentration of 200 ppm of P in 120 mM K (KCl +
KNO3) in 10% acetone given as pulse gave the highest fresh wt/cm and drought protection, whereas 1000 T plus 300 P gave the overall best combined fresh weight/cm and protection from heat and drought (Table 4). Both treatments gave a height reduction of about 50% in 14 days growth in a glass house.
S Example V
Katepwa wheat cultivars were pulse treated with A in combination with P+T (Table 5). Ancymidol alone did note provide any protection, but in combination with T+P gave 70% protection against heat and 50% against drought.
A concentration of 200, 300 and 400 ppm of P provided 50% protection against drought and 300 ppm provided maximum protection against heat with less than 40% damage.
Example VI
Katepwa wheat cultivars were pulse treated singly and in combination with A, T and P in 60 mM K(KNO3 and KCl) and 10% aceone (Table 6).
1000 T + 300 P provided the highest protection against heat and drought with the highest fresh weight (mg/cm). This treatment however, gave curving and distortion to the first leaf. The addition of 100 An elimin~te~ this distortion and provided similar protection against heat and drought. Comparison of Treatment 8 and Treatment 9 will show that addition of 1000 T provides protection against drought whereas lleaL~llellt with only P and An does not.
Example VII
Katepwa wheat cultivars were pulse treated with combinations of T, A, P and Tl in 60 mM K(KN03 and KCl) and 10% acetone (Table 7). Combined treatment widl 1000 T + 100 An + 100 P + 100 Tl provided maximum protection against heat and drought. Degree of damage from heat and drought showed the highest correlation to FW (mg/cm) and dry weight (mgxlO/cm), in-lic~ting that fresh and dry weight of see~lin~ are in~ir~tive of their ability to withstand heat and drought stresses.
Example VIII
Katewpa wheat cultivars were pulse treated with a combination of T, A
and P (TAP) or Tl, A and P (PAP) and the results compared (Table 8).
Both combinations, TAP and PAP, gave protection against heat and 5 drought, but PAP was slightly superior to TAP in protection as well as in increasing the fresh and dry weight of the see~llin~. The height of the see-lling was reduced to 1/3 by PAP whereas TAP gave reduction of about half of that of the control.
Example IX
Katepwa wheat cultivars were pulse treated with P and the see~lings immediately grown (Freshly treated) or first stored for 1 month.
Paclobutrazol (P) alone at 300 ppm provides protection to see~lling~ from heat showing damage <40% when freshly treated seeds are sown, but see-lling~ raised from one month stored treated seeds showed < 17% damage from the heat stress. However, P along does not provide much protection from drought when two week old seel1ling~ are subjected to stress (Table 9).
Example X
Leger cultivar barley seeds were pulse treated with various concentrations of T, A, P and Tl (Tables 10 and 11).
An initial trial with 1000 T + 100 An + 100 P showed that higher dose can inhibit ge~ ion. Hence, half and 1/4 doses were tried. Although gelll~ ation and emergence were not affected, the 1~2 dose (500 T + 50 An +
50 P) slightly delayed emergence (by 2 days) and the first leaf was curved and distorted. However, at lt2 dose, the protection to heat and drought stresses washighest showing damage to heat less than 7% and the drought damage less than 11%. Whereas at lh dose, the heat damage was less than 22% and drought damage was 68% (Table 10).
The treatment combination of 100 Tl + 25 An + 50 P gave the best protection against heat and drought, although all the three treatment combinations gave good protection to heat compared to control (Table 11).
. .
Example XI
Katepwa cultivar wheat seeds were imbibed for 18 hours in a combination of paclobutrazol, ancymidol and triadimefon (PAT)(PAT at 50:50:100 mg/L) with or without KCl (120 mM). The control was imbibed in S water for 18 hours. The ten day old wheat see~lling~ were exposed to a heat stress of 50C for 4 hours. ~.e~kin~ss was measured 1 day and survival two weeks after exposure to heat stress (Table 12). The higher percent le~kin~ss in the controls indic~tes loss of membrane integrity leading to lower survival rates. The treatment of PAT in the presence of KCl provided maximum protection. The appearance of the treated (P = 50 mg/L, P:T = 50:100 mg/L, PAT = 50:50:100 mg/L) versus untreated plants (water) demonstrates that the treated plants sustained minim~l damage, especially when treated with PAT, whereas the unllea~ed plant leaves curled and wilted.
The ten day old wheat see~lling~ were placed in a progr~mmed freezer and cooled at the rate of 2C/hr and survival was assessed one week after exposure to the cold (Table 13). The treatment of PAT and KCl provided m~ximllm protection similar to the results obtained with heat stress.
Water was witheld for 3, 6, or 9 days from 7 day old wheat see~linp~
and the amount of water llallspiled per plant measured. The injury was assess on day 9 on a visual rating of 0 - 10 with 10 being the most damaged and dessic~ted (Table 14).
Example XII
Tomato seeds were imbibed with PAT (25:25:100 mg/L) as in Example XI. The see~ling were exposed to heat stress of 50C for three hours. Again, the triazole treatment protected the tomato seeAling whereas the control was almost completely destroyed.
Example XIII
Soybean seeds were imbibed for 2 hours in water (control) and increasing concentrations of paclobutrazol of 5, 15 and 30 mg/L plus KCl (120mM). The three week old see~llings were subjected to drought by withholding water for 12 days. The control see~llin~ was totally dessicated -whereas the see~llinp~ treated with various concentrations of paclobutrazol and KCL were shorter and protected.
Example XIV
Katepwa cultivar wheat seeds were imbibed with triazole - 1 (PAT =
25:25:100 mg/L) and Triazole - 2 (PAT = 50:50:100 mg/L). The seeds were placed in moistened filter paper in a cold (4C) room for 10 days and then returned to room temperature. Germination was assessed after seven days (Table 15). The cold exposure reduced viability by approximately 50% and this effect was prevented by triazole treatement.
Example XV
Katepwa cultivar wheat seeds were imbibed in water or paclobutrazol (50 mg/L) for 18 hours at room temperature. They were transferred to petri dishes on moistened filter paper and exposed to 50C for between 0 to 2 hours and returned to room tem~)el~ture. Germination percentage was recorded after 3 days. After 1.5 hours of exposure to 50C, the gelll~-nation was 40% and 72% in the control and triazole treatment respectively (Table 16).
Example XVI
Loose smut infestation was compared in wheat seeds infested with loose smut and treated by imbibing with paclobutrazol (P = 25 mg/L), pulse treatment with paclobutrazol (P = 300 mg/L) or control. Treatment by imbibition completely eradicated loose smut and pulse treatment greatly decreased loose smut (Table 17).
Example XVII
Broccoli (Table 18, PAT Imbibition = 10:10:100 mg/L, pulse =
33:33:300), beans (Table 19, PAT imbibition = 10:10:100, pulse =
50:50:500); and corn seeds (Table 20, PAT imbibition = 10:10:100, pulse =
33:33:300) were treated with PAT by imbibition and by pulse, ge~ te~l in a greenhouse and then transplanted in the field at the two-leaf stage. In every case the yield was increased with PAT treatment. Generally the imbibition greatly increased the yield over the pulse and control. Although broccoli is cold tolerant, the control see~llingc tended to drop in the afternoon sun whereas the imbibed see-llingc were turgid.
Example XVIII
Katepwa and Celtic wheat cultivar seeds were treated with PAT by imbibition (PAT = 50:50:100) and pulse (PAT = 100:100:1000). The seeds were sown in the field and the height, lodging resistance, stalk breakage, and diseases of mildew, rust and septoria were measured (Table 21). The ranking scale was 0-9 with 9 being highest. Both triazole treatments initially reduced the height of plants, but just prior to harvest they were taller than controls.
The imbibition treated plants were more resistant to lodging and more effective than pulse in reducing diseases.
Spring barley Leger, and AC Burman cultivar seeds treated with PAT
(25:25:250 mg/L) by pulse treatment reduced powdery mildew, leaf rust and blotch diseases signifi~ntly (Table 22).
Example XIX
Soybeans were treated with PAT (Imbibition = 10:10:100, pulse =
33:33:300) by pulse and imbibition and sown in the field. Those sown in the summer (Table 23) demonstrated that the triazole treatment, especially imbibition, increased the height, number of pods, seeds per pod and seed weight of soybean plants.
Soybean seeds sown in early October, demonstrated that when the control leaves had started to turn yellow and senesce, the triazole-treated leaves from both pulse and imbibition treatments were still green, and the pods in these plants were larger than the controls.
Example XX
Cucumber seeds treated with PAT (10:10:100 mg/L) by imbibition treatment resulted in denser growth of the treated plants in the field due to protection from fost which inhibited the growth of the controls.
Cucumber seeds treated with PAT (Imbibition = 10:10:100, pulse =
33:33:300 mg/L) by imbibition and pulse were grown in the field and the harvests compared. The weights of cucumbers harvested per plot consisting of 2119~06 4 rows (each row planted with 10 sets of 3 seeds each) was significantly higher for the imbibition treated seeds whereas pulse was equal to control (Figure 2 a,b).
Example XXI
Tomato seeds were treated with PAT (25:25:100 mg/L)`by imbibition.
See-llin~ were gel",i"~ted in the growth room and transplanted at the 3-leaf stage. After a mild frost, control see~ling~ were damaged with necrosis of the leaf tips whereas the treated see~ling~ were protected.
Tomato seeds treated with PAT imbibition were grown in the field. The tomatoes harvested from control and treated plants were stored at 4C for 14 days. The nulllber of fruits that rotted was far greater in the control plants than the treated plants. Tomatoes harvested from control and treated plants were stored at 15C for 14 days. Again there were far more rotted tomatoes for the control plants.
Tomato seeds imbibed in water or PAT, Triazole - 1 (25:25:100 mg/L) and Triazole - 2 (50:50:100 mg/L),were gel",il-~te~ in the greenhouse and tomato see-lling~ were transplanted in the field. Tomatoes were harvested when they were mature green. The tomatoes were stored at 4C or 15C for 21 days.
Sucrose equivalent based on specific gravity showed that the treated fruits had signific~ntly higher levels than the controls (P = 0.001). The triazole treatment also signifi~ntly protected the fruits from damage during storage at both temperatures (Table 25).
Tomato seeds were treated with PAT (Imbibition 25:25:100, pulse 100:100:1000) by imbibition and pulse and disease inle~lsily measured in leafs of the tomato plants grown in the field. Disease intensity was significantly reduced in the triazole treated plants, more so with the imbibition (Table 26).
Harvests of the tomatoes demonstrated that triazole treatment promotes the yield of tomatoes. The yield was increased the most with the pulse treated seeds (Table 27).
~y~m~le XXII
Green pepper seeds were imbibed in water (control) or paclobutrazol (P
25 and 50 mg/l) in 40 mM KCI for 18 h and hardened at 40C for 2 h and air dried. Seetllinge were gel",in~ted in the greenhouse and one set of 5-week-old see~linge were transplanted early on May 10, 1993. A second set of 5-week-old see-llinge were planted on June 5, 1993.
The control see~lling~e from the first planting were damaged with necrosis of the leaf tips after a mild frost, whereas the treated see~llin~e were protected.
Thereafter there was no stress. The results reflect that the yield was increasedin the treated plants of the early planting due to protection from stress whereas in the late planting with no stress, the treatment had no effect.
Example XXIII
Potato (tubers) were dipped for 30 seconds in 10% acetone alone (control), or cont~ining paclobutrazol (P). They were allowed to air dry for 30 minutes and planted at Cambridge on May 5, 1993.
The increase in yield obtained with paclobutrazol at 0.15 mg/l is negated at the higher concentration tested. Although, the lllargi,lal 9% increase in yield for potatoes treated with 0.15 mg/L Paclobutrazol appears marginal, the growing season for summer of 1993 was particularly unstressful. Even in such conditions, the treatment provided a noticeable increase in yield. This example therefore demonstrates that potato tubers treated by this invention's method provides protection against stress.
211980i~
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Protecticn of 10-day Old Wheat See-lling~
from Ex~o~ to a Heat Stress of 50C for 4 h T e~kin~ss Survival Treatment (% Conductivity) (%) Control 71 45 KCl 64 57 PAT + KCl 30 100 Percent Survival of Wheat Seedlings After F,~ ..re to Low T~ ures % Survival Treatrnent (C) -6 -8 -10 Control 49 14 0 KCl 51 38 0 PAT + KCl 100 100 68 T~BL E 14 ~e~ ction of Tr~iralion and Protection of Wheat See~lling~ Against Drought Water Transpired g/plant Injury (0-10) Trea~nent (Days) 3 6 9 8 Control 4.8 11.1 19.1 6 K Cl 4.7 8.4 14.2 6 P A T 3.2 5.2 8.3 2 P A T + K Cl 3.2 4.1 8.2 Wheat Seeds (cv. Katepwa) Were Placed in Moi~..ed Filter Paper in Cold (4C) Room for 10 Days Then Returned to Room Tcm~.~ re. Ge~ ion was ~ ss~l After 7 days.
Treatment Germination %
Control 49 Triazole-1 94 Triazole-2 96 21I~806 -Control of Loose Smut in Wheat by Paclobutrazol (P) Treatment % Loose Smut Control 9 a Imbibition 0 b Pulse 0.9 b Field Data (Summer 1992) for Broccoli Grown in Cambridge, Ontario, Canada Treatment Yield/Plant (g) % Increase Control 183 a ---Imbibition 233 b 27 Pulse 178 a ---Field Data (Summer 1992) for Beans Grown in Arkell (1) and Cambridge - 1st Planting (2) and 2nd ~ h~r (3), Ontario, Csm~
(1) Treatment Yield/Plant (g) % Increase Control 167 a ---Imbibition 299 c 37 Pulse 198 b 19 (2) Treatment Yield/Plant (g) % Increase Control 45 a ---Imbibition 93 b 107 Pulse 40 a (-)12 (3) Treatment Yield/Plant (g) % Increase Control 44 a ---Imbibition 64 c 44 Pulse 55 b 25 Field Data (Summer 1992) for Corn Grown in Arkell, Ontario, C~(lq Treatment Yield/Plant (g) % Increase Control 159 a ---Imbibition 229 c 35 Pulse 178 b 24 2119~6 . ~
C~
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c~ x ~oo oo ~ oo t - x oo . -~ ~ c y c 211g806 ~_' 37 Table 22 Triazole reduce diseases (powdery mildew, leaf rust, and blotch) in spring barley grown in Elora, Ontario (June 24, 1992) cv. Leger a) o O
cv. AC Burman ~ , .~ .
I I
Control Pulse T~BL E 23 Field Data (Summer 1992) for Soybeans Grown in Cambridge, Ontario, Canada Treatment Height (cm) Pod/Plant (No.) Seeds/Pod (No.) Seed wt. (g) Control 52.9 30.2 2.62 0.69 Imbibition 71.6 33.4 2.78 0.79 Pulse 66.1 37.6 2.78 0.79 oo o Table 24 ~__ 1~
~_ Shelf life of tomatoes halvested ~om control and treated (imbibed-PAT) plants. The tomatoes that were spoiled in each case are shown in blacl;. Atboth storage temperatures the triazole significantly protects spoilage.
~, oo ~o o V
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~ ~ o o Table 26 Triazoles reduce Septoria leaf spots in tomato plants grown in Arkell, Ontario (Summer 1 992) I
MIDDLE LEAVES
~Q
Z
C
~ 50 -I
WHOLE PLANT
Control Imbibition Pulse Table 2 7 Triazoles promote yield of tomato plants grown in Arkell, Ontario (Summer 1 992) v~ O Control V
Imbibition 6 - v Pulse y 4-V ~
a) 2 - /~
_~
c" V
80- /~
/v//
40- ~//
O -No. of Harvests 211~gO6 T~UBLE 28 FIELD DATA (SUl\Il\IER 1993) FOR GREEN ~ ;KS
GROWN AT CAMBRIDGE, ON TWO PLANTING DATES
s Treatment Early Planting Late Planting 10 mg/l yield/plant yield/plant (g) ~ Increase (g) Control 218 a - 212 a P-25 Imbibition 276 b 27 205 a P-50 Imbibition 290 b 33 201 a FIELD DATA (SUMMER 1993) FOR POTATOES
(VAR. KENEBEC) GROWN AT CAMBRIDGE
s Treatment Yield per 30 ft. row mg/l (lbs.) Control 45 a P0.15 49a P 0.25 41 c Although preferred embodiments of the invention are described herein in detail, it will be understood by those skilled in the art that variations may bemade thereto without departing from the spirit of the invention or the scope of 25 the appended claims.
Claims (15)
1. A process for enhancing stress resistance in a plant from embryonic stage through to maturity by conditioning a plant seed to transfer stress resistance to said plant derived from said conditioned seed, said plant seed conditioning process comprising:
i) contacting said plant seed with a solution containing a compound or a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and a carrier for the compound or mixture thereof, for delivering said compound mixture through a seed coat of said seed and into said seed, ii) said solution including a source of potassium at a concentration sufficient to enhance the effects of cytokinins in said seed.
i) contacting said plant seed with a solution containing a compound or a mixture of compounds capable of inhibiting ergosterol and gibberellin biosynthesis, increasing production of abscisic acid and cytokinins and decreasing production of ethylene in plants and a carrier for the compound or mixture thereof, for delivering said compound mixture through a seed coat of said seed and into said seed, ii) said solution including a source of potassium at a concentration sufficient to enhance the effects of cytokinins in said seed.
2. A process of claim 1, wherein the compound or mixture of compounds are selected from the group of compounds consisting of:
1 -(4-chlorophenoxy-3 ,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butanone;
1-(4-chlorophenoxy)3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol;
(E)-1-cyclohexyl-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol;
all-rac-1-(biphenyl-4-yloxy)-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol;
1-(3-trifluoromethyltrityl)-1 H-1,2,4-triazole;
(?)-1-[2-(2,4-dichlorophenyl)-4-propyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4 triazole;
(?)-1-[2-(2,4-dichlorophenyl)-4-ethyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4-triazole;
1-(2,4-dichloro-.beta.-propylphenethyl)-1 H-1,2,4-triazole;
2-(2,4-dichlorophenyl)-2-hydroxy-3-methoxy-3-(1 H-1,2,4-triazol-1-yl)propane;
1-phenoxy-3-(1 H-1,2,4-triazol-1-yl)4-hydroxy-5,5-dimethyl-hexane;
1-(4-trifluormethyl)-2-(1,2,4-triazolyl-[1]-3-(5-methyl-1,3-dioxan-5-yl-propen-3-ol;
(2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol;(2RS,3RS)-1-(2,4-Dichlorophenyl)-4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol;
(E)-1-(p-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol;
(E)-1-(2,4-Dichlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol)-1-yl-1-penten-3-ol;
.alpha.-butyl-.alpha.-(4-chlorophenyl)-1 H-1,2,4-triazole-1-propanenitrile;
.alpha.-cylcopropyl-.alpha.(4-methoxy-phenyl)5-pyrimidine methanol;
5-(4-chlorophenyl)-3,4,5,9,10-pentaaza-tetra-cylco-4,5,1,02,6,08,11-dodeca-3,9-diene;
1-(2,6-diethylphenyl)-imidazole-5-carboxamide;
1 -(4-chlorophenoxy-3 ,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butanone;
1-(4-chlorophenoxy)3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol;
(E)-1-cyclohexyl-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol;
all-rac-1-(biphenyl-4-yloxy)-3,3-dimethyl-1-(1 H-1,2,4-triazol-1-yl)butan-2-ol;
1-(3-trifluoromethyltrityl)-1 H-1,2,4-triazole;
(?)-1-[2-(2,4-dichlorophenyl)-4-propyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4 triazole;
(?)-1-[2-(2,4-dichlorophenyl)-4-ethyl-1,3-dioxalan-2-ylmethyl]-1 H-1,2,4-triazole;
1-(2,4-dichloro-.beta.-propylphenethyl)-1 H-1,2,4-triazole;
2-(2,4-dichlorophenyl)-2-hydroxy-3-methoxy-3-(1 H-1,2,4-triazol-1-yl)propane;
1-phenoxy-3-(1 H-1,2,4-triazol-1-yl)4-hydroxy-5,5-dimethyl-hexane;
1-(4-trifluormethyl)-2-(1,2,4-triazolyl-[1]-3-(5-methyl-1,3-dioxan-5-yl-propen-3-ol;
(2RS,3RS)-1-(4-chlorophenyl)-4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol;(2RS,3RS)-1-(2,4-Dichlorophenyl)-4,4-dimethyl-2-(1 H-1,2,4-triazol-1-yl)pentan-3-ol;
(E)-1-(p-chlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol-1-yl)-1-penten-3-ol;
(E)-1-(2,4-Dichlorophenyl)-4,4-dimethyl-2-(1,2,4-triazol)-1-yl-1-penten-3-ol;
.alpha.-butyl-.alpha.-(4-chlorophenyl)-1 H-1,2,4-triazole-1-propanenitrile;
.alpha.-cylcopropyl-.alpha.(4-methoxy-phenyl)5-pyrimidine methanol;
5-(4-chlorophenyl)-3,4,5,9,10-pentaaza-tetra-cylco-4,5,1,02,6,08,11-dodeca-3,9-diene;
1-(2,6-diethylphenyl)-imidazole-5-carboxamide;
3. A process of claim 2 wherein said source of potassium is selected from the group consisting of KNO3, KCl and K2SO4.
4. A process of claim 2 wherein said carrier is an organic solvent which expedites delivery of said compound mixture through said seed coat, said solvent being selected from the group of solvents consisting of ketones and alcohols.
5. A process of claim 4, wherein said selected solvent is acetone.
6. A process of claim 4, wherein said selected solvent is ethanol.
7. A process of claim 4, 5, or 6 wherein said seed is contacted with said compound mixture in said solvent for less than 2 minutes.
8. A process of claim 2 wherein said carrier is water, said seed being contacted with said compound or mixture of compounds in water for an extended period in the range of 2 to 18 hours.
9. A process of claim 8 wherein said seed after said extended period of contact with water and compound or mixture of compounds is subjected to a hardening step.
10. A process of claim 9 wherein said hardening step comprises subjecting said seed to an elevated temperature in the range of 35°C to 45°C for at least 1 hour to enhance stress resistance in a plant resulting from said conditioned seed.
11. A process of claim 10 wherein said hardening step is carried out at 40°C for approximately 2 hours.
12. A process of claim 11 wherein said compound mixture comprises triadimefon, paclobutrazol and ancymidol.
13. A process of claim 11 wherein said compound comprises paclobutrazol.
14. A process of claims 12 and 13 wherein said seed is a cereal seed, said seed being conditioned in said solution for said extended period in the range of16 to 18 hours.
15. A process of claims 12 and 13 wherein said seed is a soybean or canola seed, said seed being conditioned in said solution for said extended period in the range of 2 to 4 hours
Priority Applications (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CA002119806A CA2119806A1 (en) | 1994-03-24 | 1994-03-24 | Seed conditioning process providing stress resistance |
Applications Claiming Priority (1)
| Application Number | Priority Date | Filing Date | Title |
|---|---|---|---|
| CA002119806A CA2119806A1 (en) | 1994-03-24 | 1994-03-24 | Seed conditioning process providing stress resistance |
Publications (1)
| Publication Number | Publication Date |
|---|---|
| CA2119806A1 true CA2119806A1 (en) | 1995-09-25 |
Family
ID=4153232
Family Applications (1)
| Application Number | Title | Priority Date | Filing Date |
|---|---|---|---|
| CA002119806A Abandoned CA2119806A1 (en) | 1994-03-24 | 1994-03-24 | Seed conditioning process providing stress resistance |
Country Status (1)
| Country | Link |
|---|---|
| CA (1) | CA2119806A1 (en) |
Cited By (5)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| EP1922928A1 (en) * | 2006-09-29 | 2008-05-21 | Syngeta Participations AG | A method for enhancing intrinsic productivity of a plant |
| EP2168434A1 (en) | 2008-08-02 | 2010-03-31 | Bayer CropScience AG | Use of azols to increase resistance of plants of parts of plants to abiotic stress |
| EP2255626A1 (en) | 2009-05-27 | 2010-12-01 | Bayer CropScience AG | Use of succinate dehydrogenase inhibitors to increase resistance of plants or parts of plants to abiotic stress |
| WO2014009322A1 (en) | 2012-07-11 | 2014-01-16 | Bayer Cropscience Ag | Use of fungicidal combinations for increasing the tolerance of a plant towards abiotic stress |
| CN111406469A (en) * | 2020-05-19 | 2020-07-14 | 中国农业科学院麻类研究所 | Folding retractable profile modeling seeder |
-
1994
- 1994-03-24 CA CA002119806A patent/CA2119806A1/en not_active Abandoned
Cited By (10)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| EP1922928A1 (en) * | 2006-09-29 | 2008-05-21 | Syngeta Participations AG | A method for enhancing intrinsic productivity of a plant |
| WO2008037489A3 (en) * | 2006-09-29 | 2008-05-22 | Syngenta Participations Ag | A method for enhancing intrinsic productivity of a plant |
| EP2168434A1 (en) | 2008-08-02 | 2010-03-31 | Bayer CropScience AG | Use of azols to increase resistance of plants of parts of plants to abiotic stress |
| WO2010015337A3 (en) * | 2008-08-02 | 2010-07-08 | Bayer Cropscience Ag | Use of azoles for increasing the abiotic stress resistance of plants or plant parts |
| JP2011529863A (en) * | 2008-08-02 | 2011-12-15 | バイエル・クロツプサイエンス・アクチエンゲゼルシヤフト | Use of azoles to increase abiotic stress resistance of plants or plant parts |
| US8614168B2 (en) | 2008-08-02 | 2013-12-24 | Monheim | Use of azoles for increasing the abiotic stress resistance of plants or plant parts |
| EA019605B1 (en) * | 2008-08-02 | 2014-04-30 | Байер Кропсайенс Аг | Use of azoles for increasing resistance of plants to the abiotic stress factors, a spray solution for treatment of plants and use thereof for increasing resistance of plants to the abiotic stress factors |
| EP2255626A1 (en) | 2009-05-27 | 2010-12-01 | Bayer CropScience AG | Use of succinate dehydrogenase inhibitors to increase resistance of plants or parts of plants to abiotic stress |
| WO2014009322A1 (en) | 2012-07-11 | 2014-01-16 | Bayer Cropscience Ag | Use of fungicidal combinations for increasing the tolerance of a plant towards abiotic stress |
| CN111406469A (en) * | 2020-05-19 | 2020-07-14 | 中国农业科学院麻类研究所 | Folding retractable profile modeling seeder |
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