WO2010079805A1 - イネ科植物の雄性不稔を回復させる方法および雄性不稔回復剤 - Google Patents
イネ科植物の雄性不稔を回復させる方法および雄性不稔回復剤 Download PDFInfo
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- WO2010079805A1 WO2010079805A1 PCT/JP2010/050101 JP2010050101W WO2010079805A1 WO 2010079805 A1 WO2010079805 A1 WO 2010079805A1 JP 2010050101 W JP2010050101 W JP 2010050101W WO 2010079805 A1 WO2010079805 A1 WO 2010079805A1
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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
- A01N43/56—1,2-Diazoles; Hydrogenated 1,2-diazoles
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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
- A01N37/10—Aromatic or araliphatic carboxylic acids, or thio analogues thereof; 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
- A01N39/00—Biocides, pest repellants or attractants, or plant growth regulators containing aryloxy- or arylthio-aliphatic or cycloaliphatic compounds, containing the group or, e.g. phenoxyethylamine, phenylthio-acetonitrile, phenoxyacetone
- A01N39/02—Aryloxy-carboxylic acids; Derivatives thereof
- A01N39/04—Aryloxy-acetic acids; 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
- A01N43/00—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds
- A01N43/34—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one nitrogen atom as the only ring hetero atom
- A01N43/36—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one nitrogen atom as the only ring hetero atom five-membered rings
- A01N43/38—Biocides, pest repellants or attractants, or plant growth regulators containing heterocyclic compounds having rings with one nitrogen atom as the only ring hetero atom five-membered rings condensed with carbocyclic rings
Definitions
- the present invention relates to a method for recovering the sterility of a plant in which male sterility has occurred, for example, a gramineous plant, particularly a wheat plant, and a composition for recovering the sterility of wheat.
- the present invention relates to methods and compositions for recovering the sterility of plants that have become male sterile due to high or low temperature stress, such as gramineous plants, especially wheat plants.
- Acupuncture wall tapet cells are cells that are destined to supply pollen and ultimately be destroyed by cell death.
- wheat plants such as wheat and barley
- the walls are high at 30 ° C night and 25 ° C (especially at night and above 25 ° C).
- pollen mother cell cessation of cell division occurs and this high temperature condition persists for more than 4 days, it is no longer completely recoverable and premature decay of the wall wall tapet cells occurs.
- pollen sterility male sterility
- auxin which is one of the plant hormones and plays an important role in cell division, growth, and developmental differentiation
- auxin which is one of the plant hormones and plays an important role in cell division, growth, and developmental differentiation
- varieties that are resistant to high temperature and low temperature stress have been developed by conventional breeding and creation of recombinants by genetic recombination techniques.
- auxin is a general term for plant hormones that mainly promote plant growth, and natural and synthetic auxins are known.
- natural auxins 3-indoleacetic acid (IAA) and indolebutyric acid (IBA) are known, and as synthetic auxins, 4-chloroindoleacetic acid, phenylacetic acid, 2,4-dichlorophenoxyacetic acid (2,4-D) , ⁇ -naphthaleneacetic acid (NAA), 2,6-dichlorobenzoic acid, indolebutyric acid (IBA), 4-chlorophenoxyacetic acid, ethyl 5-chlorointazole acetate, naphthoxyacetic acid and 2,4,5-trichlorophenoxyacetic acid It has been known.
- These auxins are used as growth regulators. They are used for tissue culture in laboratory environments and 2,4-D etc. are used as herbicides in the field.
- a method for recovering male sterility of grasses, particularly male sterility due to high or low temperature stress, and a composition for recovering male sterility of grasses, particularly male sterility due to high or low temperature stress. provide.
- the present invention produced male sterility comprising spraying the plant with a fertility restoring agent for gramineous plants, particularly wheat plants, which contains auxin as an active ingredient, and auxin.
- a fertility restoring agent for gramineous plants particularly wheat plants, which contains auxin as an active ingredient, and auxin.
- This is a method for restoring fertility of gramineous plants, especially wheat plants. More specifically, the present invention is as follows. (1) A fertility recovery agent for gramineous plants with male sterility, which is applied to gramineous plants so that the auxin concentration is 10 ⁇ 4 M to 10 ⁇ 7 M (M: molar concentration). (2) The fertility-recovering agent according to (1), which is applied at least once before or before the fifth leaf development date in the young panicle stage at the latest.
- Auxin is 3-indoleacetic acid (IAA), 4-chloroindoleacetic acid, phenylacetic acid, 2,4-dichlorophenoxyacetic acid (2,4-D), ⁇ -naphthaleneacetic acid (NAA), 2,6- Any one of (1) to (4), which is dichlorobenzoic acid, indolebutyric acid (IBA), 4-chlorophenoxyacetic acid, ethyl 5-chlorointazole acetate, naphthoxyacetic acid or 2,4,5-trichlorophenoxyacetic acid The fertility recovery agent as described.
- a method for recovering male sterility of a Gramineae plant that has caused male sterility which comprises spraying wheat to a auxin concentration of 10 ⁇ 4 M to 10 ⁇ 6 M.
- auxin is sprayed at least once before or before the fifth leaf development day in the young panicle stage at the latest.
- auxin will be sprayed at least once a day before the 5th leaf deployment date.
- Auxin is 3-indoleacetic acid (IAA), 4-chloroindoleacetic acid, phenylacetic acid, 2,4-dichlorophenoxyacetic acid (2,4-D), ⁇ -naphthaleneacetic acid (NAA), 2,6- Any one of (6) to (9), which is dichlorobenzoic acid, indolebutyric acid (IBA), 4-chlorophenoxyacetic acid, ethyl 5-chlorointazole acetate, naphthoxyacetic acid or 2,4,5-trichlorophenoxyacetic acid
- Gramineae plants include rice and wheat plants.
- Wheat plants include barley, wheat, rye and oat plants.
- the present invention also provides a fertility-recovering agent for rice that has caused male sterility, comprising a substance that inhibits the action of auxin as an active ingredient, and a male fertility comprising a substance that inhibits the action of auxin as an active ingredient. It is also a method to recover the fertility of rice that has caused drought.
- Substances that inhibit the action of auxin include aminooxyacetic acid (AOA), L- ⁇ - (2-aminoethoxyvinyl) glycine (AVG), p-chlorophenoxyisobutyric acid (PCIB), triiodoacetic acid (TIBA), and naphthyl Salamic acid (NPA) is included.
- AOA aminooxyacetic acid
- AVG L- ⁇ - (2-aminoethoxyvinyl) glycine
- PCIB p-chlorophenoxyisobutyric acid
- TIBA triiodoacetic acid
- NPA naphthyl Salamic acid
- fertility of gramineous plants, particularly wheat plants and rice that have become male sterile due to high or low temperatures can be easily recovered.
- damage to seed production due to catastrophic male sterility (insufficient pollen formation) due to high-temperature injury can be completely avoided by appropriate application of an auxin aqueous solution.
- FIG. 1 shows the expression of the auxin-responsive fusion gene DR5-GUS in Arabidopsis thaliana during flower development under suitable temperature cultivation conditions. The expression was most intense at the time of epilepsy.
- FIG. 2 shows changes in the expression of the auxin-responsive fusion gene DR5-GUS in Arabidopsis due to high temperatures. With time, a marked decrease in GUS activity and an induction of GUS activity in pistils and petals are observed in the anther.
- A Control (suitable temperature cultivation),
- C Day 5 of high temperature treatment,
- D Day 7 of high temperature treatment
- FIG. 3 shows the development of young panicles observed during the reproduction and growth process of barley.
- FIG. 4 represents the high temperature injury observed in the barley stamen development process. The high temperature treatment was performed from the fifth leaf development stage when the panicle was 2-3 mm until the panicle was about 10 mm.
- A The correspondence between the time of high temperature treatment and the length of young panicles is shown.
- B Pollen mother cell meiosis and tapete cell disruption in the control and high temperature treated groups (causing male sterility).
- C10, C15, and C20 are stamen micrographs at the time of the young panicle length of 10 mm, 15 mm, and 20 mm in the control group, respectively.
- H10, H15 and H20 are stamen micrographs at the time of 10 mm, 15 mm and 20 mm, respectively, in the high temperature experimental group.
- FIG. 5 shows the unobserved results observed when the barley is exposed to a high temperature of 30 ° C./night and 25 ° C. for 5 days starting from the fifth leaf development stage of barley and then continued to grow back to normal 20 ° C./night/15° C. Shows developmental wrinkles and complete pollen failure.
- FIG. 6 shows the results of quantifying the relative ratio of gene expression levels in each juvenile length of auxin-repressed protein gene of Contig 7516_at on DNA microarray 22K Barley1leyGeneChip (Affymetrix) by real-time RT-PCR.
- the vertical axis represents the relative expression level of auxin-repressed protein-gene, and the horizontal axis represents the panicle length.
- a white bar indicates a control group (suitable temperature cultivation), and a black bar indicates a high temperature treatment group.
- “Sprouting” indicates the change in expression due to high temperature treatment of the same gene in the above-ground part subjected to the same high temperature treatment for 5 days using an individual 5 days after sowing.
- FIG. 7 shows the high temperature treatment period of the fifth barley leaf development stage. Each pot solution (4 plants) was sprayed with 25 ml of auxin aqueous solution of each concentration. Spraying was performed on the 18th, 19th, 21st and 23rd days after sowing.
- FIG. 8 shows the recovery of the head length at the heading stage by spraying with an auxin aqueous solution.
- FIG. 9 shows an example of the recovery of the morphology and pollen fertility of stamens and pistils at the heading stage by spraying with an auxin aqueous solution.
- A Individual in control group (suitable temperature cultivation),
- B Individual in high temperature treatment group,
- C Individual in experimental group sprayed with high temperature treatment and auxin at each concentration.
- An example of recovery of seed fertility by auxin spraying was shown.
- a 10 -4 M auxin (NAA) spray was applied to each of the control group (suitable temperature cultivation) and the high temperature treatment group.
- the two photographs on the left show the recovery of seed fertility in the high-temperature treated individuals with auxin.
- Plants suitable for applying the present invention are plants in which the auxin level changes greatly due to high or low temperature stress, and the auxin level greatly affects the pollen formation process as a result.
- Such plants include gramineous plants, in particular barley plants (barley etc.), wheat plants (bread wheat, duran wheat, crab wheat, spell wheat, emma wheat etc.) rye plants (rye etc.) and oat plants
- Wheat plants including rice (oat, oats, oats, etc.) and rice are included.
- monocotyledonous plants and dicotyledonous plants, including wheat plants showed high auxin activity during the early development of cocoons, but it was found that the activity was significantly reduced at high temperatures.
- auxin is caused by high temperature, and cell division and early disintegration of pollen mother cells and cocoon wall cells occur, which ultimately leads to pollen sterility and reduced seed fertility.
- the auxin signal that should be reduced with normal maturation of mushrooms is not lowered by the low temperature, and as a result, the decay of wall wall tapet cells is inhibited and male sterility occurs. Therefore, when high-temperature stress occurs during the development and differentiation of wheat plants such as wheat and barley, artificially spraying a substance having an auxin action, it is possible to identify the moth-specific effects expected from high-temperature stress.
- auxin substances having the same action as the natural auxin 3-indoleacetic acid are collectively referred to as “auxin” or “substance having auxin action”.
- the auxin used in the present invention may be a natural auxin or a synthetic auxin.
- natural auxin includes 3-indoleacetic acid (IAA)
- synthetic auxin includes 4-chloroindoleacetic acid, phenylacetic acid, 2,4-dichlorophenoxyacetic acid (2,4-D), ⁇ - Naphthalene acetic acid (NAA), 2,6-dichlorobenzoic acid, indole butyric acid (IBA), 4-chlorophenoxyacetic acid, ethyl 5-chlorointazole acetate, naphthoxyacetic acid and 2,4,5-trichlorophenoxyacetic acid.
- the auxin that can be used in the present invention is not particularly limited, and all natural and synthetic auxins can be used.
- substances that inhibit the auxin action used in the present invention include aminooxyacetic acid (AOA), L- ⁇ - (2-aminoethoxyvinyl) glycine (AVG), p-chlorophenoxyisobutyric acid (PCIB), triiodoacetic acid. (TIBA) and naphthyl salamic acid (NPA) are included.
- the plants that can be used in the present invention are plants that are likely to become male sterilized by high temperature or low temperature, particularly plants that can cause male sterility by high temperature stress.
- Such plants include gramineous plants, in particular wheat plants, gramineous maize (eg maize), solanaceae eggplants (eg tomato, eggplant, pepper, peppers and paprika), legume cowpea (eg Azuki bean and cowpea), a plant of the Brassicaceae genus Brassica (for example, rapeseed).
- Plants that are particularly suitable for restoring male sterility according to the present invention are grasses.
- Gramineae plants include rice and wheat plants.
- Wheat plants include barley, wheat, rye and oat plants. Barley plants include barley, wheat plants include bread wheat, duran wheat, crab wheat, spelled wheat and emma wheat, rye plants include rye, oat plants include oat, oat and Oats are included.
- auxin or a substance that inhibits auxin action to a wheat plant application of these solutions to a plant using a solvent that is less harmful to the plant, such as leaf and stem Dispersion over the entire aerial part or part thereof is preferred.
- Spraying is preferably by spraying.
- Auxin can also be applied to the petiole or stem portion where the young panicle is located. In the case of barley and rice, young ears are not picked at the right time to spread auxin, so they cannot be sprayed directly on young ears, but when young ears are exposed at the right time, Auxin can be sprayed or applied directly.
- auxin can be sprayed or applied directly to the inflorescences or flower buds exposed to the outside.
- the concentration of auxin is preferably 10 ⁇ 4 M to 10 ⁇ 8 M, and 10 ⁇ 4 M to 10 ⁇ 7 M. More preferably, 10 ⁇ 5 M to 10 ⁇ 6 M is particularly preferable.
- 10 ⁇ 4 M to 10 ⁇ 7 M is more preferable, and 10 ⁇ 5 M to 10 ⁇ 6 M is particularly preferable.
- a surfactant may be added to the fertility recovery agent of the present invention in order to improve the spreadability of the fertility recovery agent of the present invention to the plant body.
- a surfactant such as Tween 20 can be added as necessary when auxin is given to a grass family according to the present invention.
- the surfactant concentration is preferably about 0.1% (v / v).
- auxin When auxin is given to gramineous plants according to the present invention, it is preferably performed at a time when the auxin level can affect pollen formation.
- the preferred time to give auxin is generally from the start of stamen differentiation or the time when the shoot apex begins to differentiate into young panicles (around the 4th leaf development stage) to the end of meiosis of the pollen mother cells (around the 6th leaf development stage). Until the microspore stage.
- a recombinant plant in which a ⁇ -glucuronidase gene (GUS) is fused to an auxin-responsive transcription activation sequence DR5 for each plant species or plant variety is prepared, and the male plant of the recombinant plant,
- the preferred period of giving auxin can be clarified by examining the time course of GUS activity. The time when GUS activity in sputum significantly increases or decreases would be an appropriate time to give auxin. For plants in which the stamen differentiation start time is not known, the preferred time to give auxin can be determined as described above. Production of a recombinant plant can be performed according to a known method for each plant species. It is also possible to directly measure auxin levels in sputum histologically and biochemically using anti-auxin antibodies. The optimum auxin concentration for each plant species or plant variety can be determined in the same manner.
- At least one fertility recovery agent of the present invention or auxin is given according to the present invention during or before the fifth leaf development date in the young panicle stage. It is preferable. In particular, higher temperatures at night than during daytime tend to cause pollen sterility in wheat plants. Therefore, when it is predicted that the day when the night temperature will be 25 ° C. or higher continuously from the fifth leaf deployment date will continue for three days or more, the inertia of the present invention will be performed at least once a day before the fifth leaf deployment. It is particularly preferred to provide a restoring agent or auxin according to the present invention.
- the fertility-recovering agent of the present invention or the auxin according to the present invention at least once during the period from the fifth leaf development date to 5 days later, for example, at least once before or after the fifth leaf deployment date.
- a biodegradable auxin for example, IAA
- IAA biodegradable auxin
- each plant can be treated as described above when high temperature damage is expected under the conditions where each plant species or plant variety is cultivated. It is preferable to use the optimal concentration determined in advance for the seed or plant variety.
- Low temperature damage in rice is caused by exposure to low temperature stress at a maximum temperature of less than 20 ° C during the booting period (immediately after meiosis of the pollen mother cells) for several days. If a low temperature is expected immediately after division, spraying with a substance that inhibits auxin activity is effective.
- the treatment concentration is in accordance with the method of use within the range of known physiological influence using each substance.
- the auxin level in cocoons is the strongest at the time of cocoon development, and shows that the auxin level changes dynamically during the development period, gradually decreasing as the pod wall tapet cells disappear and pollen matures. It was.
- BIOTRON LH300RDS under conditions of a day temperature of 20 ° C, a night temperature of 15 ° C, and a day length of 16 hours.
- Twenty fluorescent lamps for plant growth (16; National FL40S FR-P, 4; National FL20S FR-P) were used as light sources.
- the seeding day is day 0, and from day 18 to 19 after seeding when the fifth leaf appears, the plants are grown under high temperature conditions of 30 ° C day temperature, 25 ° C night temperature, and 16 hours day length for 5 days (120 hours).
- the high temperature treatment was performed.
- the cultivation conditions were returned to the day temperature of 20 ° C. and the night temperature of 15 ° C., the high temperature treatment was stopped, and the cultivation was continued until the heading and seed ripening thereafter.
- the young panicle length becomes 2 to 3 mm, and in each spikelet, the primary process of the stamen and pistil develops and differentiates, and then the reproductive growth proceeds continuously (Fig. 3, 4).
- the young panicle reaches about 10 mm, and the four-layered wing wall cells containing tapet cells are completed, and the pollen mother cells begin to shift to meiosis.
- the pollen mother cells begin to undergo meiosis at 15 mm panicle length, and pollen microspores are formed at the 20 mm panicle length, and the decay of the wall wall tapetum cells proceeds. It takes 5 days to reach the 10 mm to 20 mm panicles.
- HYPONEX 0.2% (v / v) (Hyponex Japan Co., Ltd.) was placed in the lower box, and used as a plant water source. Seeds stored at 4 ° C. in the dark were sown at 25 ° C. for 48 hours in the dark at 4 corners of germinated barley Haruna Nijo in 4 corners per pot. They were grown using an artificial meteorological instrument (Japan Medical Instrument Manufacturing Co., Ltd. BIOTRON LH300RDS) under conditions of a day temperature of 20 ° C, a night temperature of 15 ° C, and a day length of 16 hours. Twenty fluorescent lamps for plant growth (16; National FL40S FR-P, 4; National FL20S FR-P) were used as light sources.
- auxin aqueous solution to plants is 18 days after seeding (the day before high temperature treatment), 19 days (just before high temperature treatment), 21 days (high temperature treatment period), 23 days (high temperature treatment period) A total of 4 times was performed (FIG. 7).
- the auxin aqueous solution is sprayed almost uniformly on the entire leaves and stems within 10 minutes after leaving the plant once outside the artificial weather device. Returned promptly.
- the amount of auxin solution sprayed is 25 ml per pot (about 6 ml per barley).
- Three spikelets before flowering pollination were collected from barley ears that had emerged from day 33 to day 40 of sowing.
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Abstract
Description
東北地方におけるイネの低温障害(冷害)は、穂ばらみ期(花粉母細胞の減数分裂直後)に、最高気温が20℃に満たない低温ストレスに数日間曝されると、葯壁タペート細胞の肥大が生じ、花粉形成が阻害される。葯壁タペート細胞は、花粉に影響を供給し、最終的には細胞死により崩壊するべく運命決定された細胞である。
一方、コムギやオオムギなどのムギ類植物は、イネとは逆に、高温ストレスに対して感受性が高く、昼30℃夜25℃(特に夜間25℃以上の条件では)の高温条件では、葯壁細胞ならびに花粉母細胞の細胞分裂の停止が生じ、この高温条件が4日間以上継続すると、もはや完全に回復不能な状況に陥り、葯壁タペート細胞の早期崩壊が生じる。その結果、正常な花粉が形成できず花粉不稔を生じ(雄性不稔)、最終的に種子稔性の低下につながる。これら細胞分裂の停止や早期崩壊は、葯においてのみ観察されるものであり、雌蕊や葉、茎などの生育には影響なく、雄性の花粉形成においてのみ観察される(Sakataら、Journal of Plant Research (2000) 113,395-402、Abikoら、Sexual Plant Reproduction (2005) 18,91-100)。この高温障害時には、大規模な遺伝子発現の変化が生じていることが、DNAマイクロアレイによる網羅的な発現解析により明らかになった。なかでもオーキシンによりその遺伝子発現が抑制される遺伝子であるオーキシン・リプレスド・プロティン遺伝子(auxin repressed protein genes)の発現が幼穂の高温障害時に誘導される(Oshinoら、Molecular Genetics and Genomics (2007) 278,31-42)。すなわち、植物ホルモンの1つで、細胞の分裂や成長、発生分化に重要な役割を担うオーキシンが、高温条件下では葯特異的に発現が低下する可能性が考察された。特に、5葉期開始時前述のような高温環境に置かれるとタペート細胞並びに花粉母細胞の発生および分化が止まり、その後形成される葯中には花粉が形成されないことが知られている(Oshinoら、Molecular Genetics and Genomics (2007) 278,31-42)。
これらの問題を解決するためには、高温や低温ストレスにより耐性を示す品種が従来の育種や遺伝子組換え技術による組換体の作出などによって開発が続けられている。
より具体的には、本発明は以下の通りである。
(1)オーキシン濃度が10-4 M~10-7 M(M:モル濃度)となるようにイネ科植物へ散布するための、雄性不稔を生じたイネ科植物用稔性回復剤。
(2)遅くとも幼穂期における第5葉展開日またはその前までの間に少なくとも1回散布するための、(1)記載の稔性回復剤。
(3)第5葉展開日から連続して夜間気温が25℃以上である日が3日以上続くことが予測される際に、第5葉展開日の1日前までに少なくとも1回散布するための、(1)記載の稔性回復剤。
(4)第5葉展開日から5日後までの間に少なくとも1回散布するための、(1)記載の稔性回復剤。
(5)オーキシンが、3-インドール酢酸(IAA)、4-クロロインドール酢酸、フェニル酢酸、2,4-ジクロロフェノキシ酢酸(2,4-D)、α-ナフタレン酢酸(NAA) 、2,6-ジクロロ安息香酸、インドール酪酸(IBA)、4-クロロフェノキシ酢酸、5-クロロインタゾール酢酸エチル、ナフトキシ酢酸または2,4,5-トリクロロフェノキシ酢酸である、(1)~(4)のいずれかに記載の稔性回復剤。
(6)オーキシン濃度が10-4 M~10-6 Mとなるようにムギ類へ散布することを含む、雄性不稔を生じたイネ科植物の雄性不稔を回復させる方法。
(7)遅くとも幼穂期における第5葉展開日またはその前までの間に少なくとも1回オーキシンを散布する、(6)記載の方法。
(8)第5葉展開日から連続して夜間気温が25℃以上である日が3日以上続くことが予測される際に、第5葉展開日の1日前までに少なくとも1回オーキシンを散布する、(6)記載の方法。
(9)第5葉展開日から5日後までの間に少なくとも1回オーキシンを散布するための、(6)記載の方法。
(10)オーキシンが、3-インドール酢酸(IAA)、4-クロロインドール酢酸、フェニル酢酸、2,4-ジクロロフェノキシ酢酸(2,4-D)、α-ナフタレン酢酸(NAA) 、2,6-ジクロロ安息香酸、インドール酪酸(IBA)、4-クロロフェノキシ酢酸、5-クロロインタゾール酢酸エチル、ナフトキシ酢酸または2,4,5-トリクロロフェノキシ酢酸である、(6)~(9)のいずれかに記載の方法。
本発明において、イネ科植物にはイネ、ムギ類植物が含まれる。ムギ類植物には、オオムギ属、コムギ属、ライギ属およびカラスムギ属植物が含まれる。
本明細書において雄性不稔の「稔性回復」とは、植物において花粉形成過程の不全を抑制し、正常な花粉を形成する能力を回復させることをいう。
一般にムギ類植物を含む単子葉植物および双子葉植物において、葯の初期発生過程では高いオーキシン活性がみられるが、高温により顕著にその活性が低下することが明らかになった。その結果、高温によるオーキシンの欠乏が生じ、花粉母細胞ならびに葯壁細胞の細胞分裂の停止と早期崩壊が生じ、最終的に花粉不稔、種子稔性の低下に至ると考えられる。一方、イネの低温障害は、通常の葯の成熟に伴って低下すべきオーキシンシグナルが、低温により低下しなくなり、その結果葯壁タペート細胞の崩壊が阻害され、雄性不稔を生じると考えられる。
従って、コムギやオオムギなどのムギ類植物の葯の発生分化過程において高温ストレスが生じた際には、オーキシン作用を有する物質を人為的に散布することで、高温ストレスにより想定される葯特異的なオーキシンの欠乏を補い、正常に花粉母細胞ならびに葯壁細胞の細胞分裂、発生分化を進行させることにより、雄性不稔を生じたムギ類植物の稔性を回復させることができる。
一方、イネの葯の発生分化過程で低温ストレスにさらされた場合には、オーキシン作用を阻害する物質をイネに与えることにより、低温ストレスにより想定される葯特異的なオーキシン作用の過剰を補い、正常に葯壁細胞の崩壊を進行させる、その結果雄性不稔から回復させることができる。
また本発明において使用するオーキシン作用を阻害する物質には、アミノオキシ酢酸(AOA)、L-α-(2-アミノエトキシビニル)グリシン(AVG)、p-クロロフェノキシイソ酪酸(PCIB)、トリヨード酢酸(TIBA)およびナフチルサラミック酸(NPA)が含まれる。
本発明によりイネ科植物、特にムギ類の稔性を回復させる目的でオーキシンを与える場合は、オーキシンの濃度は10-4 M~10-8 Mが好ましく、10-4 M~10-7 Mがより好ましく、10-5 M~10-6 Mが特に好ましい。特にムギ類以外単子葉植物の場合は10-4 M~10-7 Mがより好ましく、10-5 M~10-6 Mが特に好ましい。双子葉植物は一般にオーキシンへの感受性が単子葉植物よりも高いので、10-5 M~10-8 Mがより好ましく、10-6 M~10-7 Mが特に好ましい。散布するオーキシン濃度が高すぎる場合は、オーキシン誘導性のエチレン合成が植物体内で生じ、その作用として老化や成長阻害につながる可能性がある。従って、散布するオーキシン濃度が比較的高い場合は、この誘導性エチレン合成の阻害効果を示すアミノオキシ酢酸(AOA)またはL-α-(2-アミノエトキシビニル)グリシン(AVG)を加えることもできる。これによりオーキシン誘導性のエチレン合成による副作用を軽減することができる。
場合により、本発明の稔性回復剤の植物体への展着性を向上させるため本発明の稔性回復剤に界面活性剤を添加することもできる。または本発明によりイネ科植物へオーキシンを与える場合にも必要により界面活性剤,例えばTween20を添加することもできる。界面活性剤の濃度は約0.1%(v/v)が好ましい。
イネの低温障害では、穂ばらみ期(花粉母細胞の減数分裂直後)に、最高気温が20℃に満たない低温ストレスに数日間曝されることにより生じるもので、従って、花粉母細胞の減数分裂直後の時期に低温が予想される場合、オーキシン活性を阻害する物質の散布が有効である。また、処理濃度はそれぞれの物質を用いた既知の生理学的影響を及ぼす範囲内での使用方法に準ずる。
モデル植物の1つであるシロイヌナズナを用いて葯特異的および発生時期特異的なオーキシン発現および高温条件下における葯特異的なオーキシン低下を解析した。
実験にはオーキシン応答性の転写活性化配列DR5にβ-グルクロニダーゼ遺伝子(GUS)を融合させた融合遺伝子を組み込んだシロイヌナズナは組換え体(Plant Cell 9,1963-1971,1997)を使用した。この株はオーキシンに応答して融合遺伝子DR5-β-グルクロニダーゼを発現する。なお本株はTom Guilfoyle氏(University of Missouri, Columbia)から入手した。
DR5-GUS系統のシロイヌナズナを直径8センチのポットに約10-20粒播種して、23℃のインキュベーターで育成すると約4週間で花芽の形成が開始した。この花芽をサンプリングして、アセトン処理後に、β-グルクロニダーゼの基質であるX-Glucを2mMの濃度で作用させ、β-グルクロニダーゼ(Gus)を発現している細胞を検出した。23℃の条件下では、花粉の減数分裂期前後から小胞子期までの期間において、葯壁タペート細胞と花粉母細胞に強いGUSの発現(青緑色に染色)が観察された(図1)。その後、葯壁タペート細胞の崩壊の時期から花粉形成が進行するに従ってGUS発現は低下し、成熟花粉においては完全に消失した(図1)。すなわち、葯におけるオーキシンレベルは、葯の発生時期に最も強く、葯壁タペート細胞が消失して花粉が成熟する過程で徐々に低下するという、発生時期特異的にダイナミックな変化をすることが示された。
一方で、雌蕊や花弁などの他の組織においてはGUSの発現シグナルが、高温で増加することが確認された(図2)。これまで、芽生えなど栄養成長組織においては、高温はオーキシンの発現シグナルを増加させることが報告されており、雌蕊や花弁での発現増加はこの点で一致するが、雄蕊の葯の発生分化過程においては高温が全く逆な影響を及ぼすことは、高温障害が雄蕊により顕著に生じ易い原因であろうと思われる。
実験に使用したオオムギは、二条オオムギの栽培品種であるはるな二条(Hordeum vulagare L.cv Haruna-nijo)である。暗黒下4℃で保存した種子を25℃暗黒下で48時間、催芽し、発芽した種子を園芸用育苗培土(全農クレハ培土:窒素 0.4,リン酸 1.9,カリ 0.6,クド 0.2g kg-1)を1リットル詰めた直径11cmの円形ポッドに円形に10粒播種した。それらを人工気象器(株式会社日本医科器械製作所BIOTRON LH300RDS)で、昼温20℃、夜温15℃、日長16時間の条件で育成させた。光源として、20本の植物育成用蛍光灯(16本;National FL40S FR-P、4本;National FL20S FR-P)を用いた。播種当日を0日目とし、第5葉が出現する播種後18~19日目から、昼温30℃、夜温25℃、日長16時間の高温条件で育成し、5日間(120時間)の高温処理を行った。5日経過した播種後23~24日目に昼温20℃、夜温15℃の栽培条件に戻し、高温処理を停止させ、その後の出穂、種子の登熟まで栽培を継続させた。
一方で、第5葉が展開する時期から5日間の高温処理区においては、幼穂長の伸長は対照区と比べ同等もしくはより促進される傾向にあるが、葯壁細胞ならびに花粉母細胞では、細胞分裂が早期停止し、さらに10mmの幼穂長において、花粉母細胞は未成熟な状況での減数分裂が開始されるが、最終的には花粉小胞子も退化する(図4)。
この高温条件下では雌蕊を含むその他の組織や器官における発生分化の異常は生じず、その結果、開花期には花粉を全く持たない葯が形成されるが、雌蕊は稔性を有しており完全な雄性不稔(花粉不全)が生じる(図5)。
これらの幼穂において、遺伝子オーキシン・リプレスド・プロティン遺伝子(auxin repressed protein genes)の発現が、高温条件下で対照区と比較して有意に発現上昇していることが観察された(図6)。すなわち、シロイヌナズナと同様、高温条件下ではオオムギにおいてもオーキシンが低下することが強く示唆された。
オーキシンの効果を確認する実験では、上記の園芸用育苗培土を用いる他に、オーキシン水溶液の処理の制御がし易いように、2つの角型プラントボックス(60mm×60mm×100mm(タテ×ヨコ×高さ)、AGCテクノグラス株式会社 プラントボックス(型番CUL-JAR300))を上下二段に重ねて、上段側の底部中央には直径1cmほどの穴をあけ、結び目を作った長さ6cmのアルコールランプの芯を通し、その上にバーミキュライト(ニッタイ株式会社 農園芸用バーミキュライト)を300ml詰めた。下段のボックスにはHYPONEX 0.2%(v/v)(株式会社ハイポネックスジャパン)を入れ、植物の養水源とした。暗黒下4℃で保存した種子を25℃にて暗黒下で48時間、催芽したオオムギはるな二条を1ポットあたり4粒、上段容器の4隅に播種した。それらを人工気象器(株式会社日本医科器械製作所BIOTRON LH300RDS)で、昼温20℃、夜温15℃、日長16時間の条件で育成させた。光源として、20本の植物育成用蛍光灯(16本;National FL40S FR-P、4本;National FL20S FR-P)を用いた。
この栽培法においても、培養土を用いた場合と同様に、播種当日を0日目と数えると、第5葉が出現し展開を開始するのは播種後18~19日目となる。播種後19日目から昼温30℃、夜温25℃、日長16時間の高温条件下で栽培し、5日間(120時間)の高温処理を行い、その後(播種後24日目)に通常の昼温20℃、夜温15℃、日長16時間の条件に戻した。
播種33~40日目にかけて出穂してきたオオムギの穂から、3粒ずつ開花受粉前の穎花を採取した。これらをFAA固定液(ホルムアルデヒド 5%,酢酸 5%,エタノール 45%,H2O 45%)で4℃暗黒下、24時間固定し、最終的には0.1 Mリン酸バッファー(pH7.2)につけて4℃で保存した。
固定した穎花から雄蕊ならびに雌蕊を実体顕微鏡下(OLYMPUS SZX12)で解剖し、顕微鏡カメラ(OLYMPUS DP70)によりデジタル写真撮影を行った。それぞれの葯長は、デジタル画像を元に、画像処理ソフトのImage J(葯の大きさの測定は、アメリカ国立衛生研究所よりフリーに公開)を用いて測定した。各オーキシンの各濃度において3個体分9粒(1個体あたり3粒)の種子を観察し、それぞれの種子内に3つ存在する葯の大きさを測定し、その平均値と標準誤差を求めた(一つの濃度処理区あたり約27個の葯を測定した)。花粉の成熟度を調べる目的で、ヨードカリウム染色液(MERCK社製 ルゴール溶液(製品番号 109261))を用いて、一つの濃度処理区あたり3個の葯を染色し、顕微鏡(OLYMPUS BX51)による花粉観察と顕微鏡カメラ(OLYMPUS DP70)によりデジタル写真撮影を行った。
高温処理区では、オーキシン効果の比較対照として、DMSOと界面活性剤をそれぞれ0.1%(v/v)の濃度にした混合液、水のみを散布した場合、出穂期の葯の伸長は顕著に抑制され、葯長は1.5mm程度にしか伸長せず(図8)、さらに内部には全く正常な花粉粒の形成がみられず(図9B)、何も散布しない場合と同様で、回復効果は得られなかった。10-6 MのIAA、2,4-DまたはNAAの散布では葯長が1.8mm~2mm程度に、10-5 Mおよび10-4 Mのオーキシン水溶液の散布ではいずれの場合も葯長が2.5mm前後まで伸長し、高温条件下でも有意に回復することが明らかになった(図8)。また、10-6 M、10-5 M、10-4 M のいずれかのオーキシン処理により葯長が回復した葯内はヨードカリウム染色液で黒紫色に染色され、デンプンを十分に蓄積した正常な成熟花粉が形成されていることも確認できた(図9C)。前述のように、高温処理サンプルにおいても、10-4 Mの2,4-D処理区においては早期に葉が枯れはじめ、植物体にとってはオーキシンによる負の影響が生じていたが、IAAならびにNAAでは10-4 Mでも副作用はほとんど観察されなかった。
また、オーキシンの散布を当初の1回にした場合でも、有意な稔性の回復効果がみられるとともに、園芸培養土を用いた栽培においても、全く同様に、オーキシン処理による高温障害の回避が可能であることが確かめられている。
Claims (15)
- オーキシンを有効成分とする、雄性不稔を生じたイネ科植物用稔性回復剤。
- オーキシン濃度が10-4M~10-7Mとなるようにイネ科植物へ散布するための、請求項1記載の稔性回復剤。
- 遅くとも幼穂期における第5葉展開日またはその前までの間に少なくとも1回散布するための、請求項1または2記載の稔性回復剤。
- 第5葉展開日から連続して夜間気温が25℃以上である日が3日以上続くことが予測される際に、第5葉展開日の1日前までに少なくとも1回散布するための、請求項1または2記載の稔性回復剤。
- 第5葉展開日から5日後までの間に少なくとも1回散布するための、請求項1または2記載の稔性回復剤。
- オーキシンが、3-インドール酢酸(IAA)、4-クロロインドール酢酸、フェニル酢酸、2,4-ジクロロフェノキシ酢酸(2,4-D)、α-ナフタレン酢酸(NAA) 、2,6-ジクロロ安息香酸、インドール酪酸(IBA)、4-クロロフェノキシ酢酸、5-クロロインタゾール酢酸エチル、ナフトキシ酢酸または2,4,5-トリクロロフェノキシ酢酸である、請求項1~5のいずれか1項記載の稔性回復剤。
- イネ科植物がムギ類植物である、請求項1~6のいずれか1項記載の稔性回復剤。
- ムギ類植物が、オオムギ属、コムギ属、ライムギ属またはカラスムギ属植物である、請求項7記載の稔性回復剤。
- 濃度が10-4M~10-7Mとなるようにイネ科植物へオーキシンを散布することを含む、雄性不稔を生じたイネ科植物の雄性不稔を回復させる方法。
- 遅くとも幼穂期における第5葉展開日またはその前までの間に少なくとも1回オーキシンを散布する、請求項9記載の方法。
- 第5葉展開日から連続して夜間気温が25℃以上である日が3日以上続くことが予測される際に、第5葉展開日の1日前までに少なくとも1回オーキシンを散布する、請求項9記載の方法。
- 第5葉展開日から5日後までの間に少なくとも1回オーキシンを散布するための、請求項9記載の方法。
- オーキシンが、3-インドール酢酸(IAA)、4-クロロインドール酢酸、フェニル酢酸、2,4-ジクロロフェノキシ酢酸(2,4-D)、α-ナフタレン酢酸(NAA) 、2,6-ジクロロ安息香酸、インドール酪酸(IBA)、4-クロロフェノキシ酢酸、5-クロロインタゾール酢酸エチル、ナフトキシ酢酸または2,4,5-トリクロロフェノキシ酢酸である、請求項9~12のいずれか1項記載の方法。
- イネ科植物がムギ類植物である、請求項9~13のいずれか1項記載の方法。
- ムギ類植物が、オオムギ属、コムギ属、ライムギ属またはカラスムギ属植物である、請求項14項記載の方法。
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CN105794451A (zh) * | 2016-03-15 | 2016-07-27 | 河南师范大学 | 利用snp促使温敏核不育小麦bns不育系花粉彻底败育的方法 |
CN105794509A (zh) * | 2016-03-15 | 2016-07-27 | 河南科技学院 | 一种提高温敏核不育小麦bns结实率的snp化学调控方法 |
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CA2867385A1 (en) | 2012-03-13 | 2013-09-19 | Pioneer Hi-Bred International, Inc. | Genetic reduction of male fertility in plants |
US10155961B2 (en) | 2012-03-13 | 2018-12-18 | Pioneer Hi-Bred International. Inc. | Genetic reduction of male fertility in plants |
MX2014011037A (es) | 2012-03-13 | 2015-05-15 | Pioneer Hi Bred Int | Reduccion genetica de la fertilidad masculina en plantas. |
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KR101669887B1 (ko) * | 2015-03-05 | 2016-10-28 | 부산대학교 산학협력단 | 벤조인산을 유효성분으로 포함하는 임신 촉진용 조성물 |
US10934556B2 (en) | 2015-04-16 | 2021-03-02 | Council Of Scientific And Industrial Research | Reversible expression system for transgene expression in plants |
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CN111802203A (zh) * | 2020-06-02 | 2020-10-23 | 佛山科学技术学院 | 一种评估豌豆生长情况的方法 |
CN115669402B (zh) * | 2022-09-16 | 2023-06-20 | 北京林业大学 | 高温恢复百合远缘杂种育性的方法与设备 |
CN117158424B (zh) * | 2023-11-02 | 2024-01-16 | 浙江大学海南研究院 | 花粉发育不同时期不同试剂处理提高白菜花粉耐热性方法 |
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JPH03123420A (ja) * | 1989-10-04 | 1991-05-27 | Zenkoku Nogyo Kyodo Kumiai Rengokai | 稲の障害型冷害時の収量低下防止方法 |
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Cited By (2)
Publication number | Priority date | Publication date | Assignee | Title |
---|---|---|---|---|
CN105794451A (zh) * | 2016-03-15 | 2016-07-27 | 河南师范大学 | 利用snp促使温敏核不育小麦bns不育系花粉彻底败育的方法 |
CN105794509A (zh) * | 2016-03-15 | 2016-07-27 | 河南科技学院 | 一种提高温敏核不育小麦bns结实率的snp化学调控方法 |
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Publication number | Publication date |
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EP2394513A4 (en) | 2014-05-14 |
JPWO2010079805A1 (ja) | 2012-06-28 |
CN102348385A (zh) | 2012-02-08 |
US8361929B2 (en) | 2013-01-29 |
US20110306498A1 (en) | 2011-12-15 |
JP5614647B2 (ja) | 2014-10-29 |
AU2010203981A2 (en) | 2014-06-26 |
EP2394513A1 (en) | 2011-12-14 |
AU2010203981A1 (en) | 2011-07-28 |
CA2749158A1 (en) | 2010-07-15 |
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