WO2018032925A1 - 水稻小粒型不育系的育种方法及杂交水稻的轻简制种方法 - Google Patents

水稻小粒型不育系的育种方法及杂交水稻的轻简制种方法 Download PDF

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WO2018032925A1
WO2018032925A1 PCT/CN2017/093249 CN2017093249W WO2018032925A1 WO 2018032925 A1 WO2018032925 A1 WO 2018032925A1 CN 2017093249 W CN2017093249 W CN 2017093249W WO 2018032925 A1 WO2018032925 A1 WO 2018032925A1
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rice
parent
hybrid
female parent
small
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French (fr)
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唐文帮
熊跃东
丁新才
易国良
许强
冯宁
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湖南希望种业科技股份有限公司
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Priority to US16/638,758 priority Critical patent/US11246275B2/en
Publication of WO2018032925A1 publication Critical patent/WO2018032925A1/zh

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    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
    • A01H1/02—Methods or apparatus for hybridisation; Artificial pollination ; Fertility
    • A01H1/022—Genic fertility modification, e.g. apomixis
    • A01H1/023—Male sterility
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H1/00—Processes for modifying genotypes ; Plants characterised by associated natural traits
    • A01H1/04—Processes of selection involving genotypic or phenotypic markers; Methods of using phenotypic markers for selection
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H5/00—Angiosperms, i.e. flowering plants, characterised by their plant parts; Angiosperms characterised otherwise than by their botanic taxonomy
    • A01H5/10—Seeds
    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01H—NEW PLANTS OR NON-TRANSGENIC PROCESSES FOR OBTAINING THEM; PLANT REPRODUCTION BY TISSUE CULTURE TECHNIQUES
    • A01H6/00—Angiosperms, i.e. flowering plants, characterised by their botanic taxonomy
    • A01H6/46—Gramineae or Poaceae, e.g. ryegrass, rice, wheat or maize
    • A01H6/4636—Oryza sp. [rice]

Definitions

  • the invention relates to the field of rice seed production, in particular to a method for breeding a rice small-sized sterile line. Furthermore, the present invention relates to a method of seed production comprising hybrid rice.
  • hybrid rice seed production requires seeding, transplanting, and different field management to obtain the ideal flowering stage.
  • the manual pollination and parental manual harvesting are complicated, labor-intensive, and costly.
  • agricultural labor is becoming increasingly scarce, which makes the labor-intensive and intensive farming method of hybrid rice seed production lack labor enforcement, which will seriously affect the safe production and supply of hybrid rice seeds.
  • the invention provides a breeding method of rice small-sized male sterile line and a light-smelting seed production method of hybrid rice, so as to solve the technical problem of mixed mixing of parental parents during seed production.
  • the present invention provides a breeding method for a rice small-sized male sterile line, comprising the following steps:
  • hybrid Hybrid F1 was obtained, F1 was planted and hybridized with H155S as the female parent, and the re-crossed hybrid F1' was obtained.
  • the replanted hybrid F1′ was obtained to obtain the F2′ generation, in the F2′ generation and F3' generation separation Seeds with ideal plant type, strong tillering ability, short planting, and small grain size are planted, and then the seeds of F5' generation and above obtained by directed breeding for more than 2 generations are rice small-type sterile lines.
  • each generation from the F1' generation alternately under the condition of short-day low temperature and long-day low temperature is that the illumination length is less than 12 hours and the daily average temperature is lower than 23 °C
  • the long-term low temperature condition is that the illumination length is greater than 12 hours
  • the daily average temperature is lower than 23 °C.
  • the rice granule-type sterile line is a seed of the F5'-F9' generation.
  • the rice small-sized sterile line has a thousand-grain weight of 8 to 22 g and a grain thickness of less than 2.1 mm.
  • a method for seed production of hybrid rice comprises the following steps:
  • the rice large-grain type restorer line was used as the male parent and the above-mentioned rice small-type sterile line was used as the female parent group to simultaneously seed and seed production.
  • the husband and the mother are mixed and harvested at the same time.
  • the parent and female seeds are screened and separated, and the parent seeds are hybrids.
  • the rice large-grain type restorer line includes R9311, Wanghui 780, Xinhui 1998 and Wanghui 091.
  • Wanghui 780 is obtained through breeding, and the breeding process includes the following steps:
  • Hybrids R9311 and Shuhui 498 were used to obtain hybrid F1", and F1" and F2" were propagated.
  • the F3"-F5" generations were screened for strains with a grain size greater than 30 g, a grain thickness greater than 2.2 mm, and an excellent rice quality.
  • the seed is planted for the next generation, and the obtained F6" or higher seed is the rice large-grain type restorer.
  • the rice large-sized restorer has a thousand particles larger than 30 g and a grain thickness of 2.2 mm or more.
  • the sow form of the male parent and the female parent includes the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the male parent, the male parent
  • the proportion of the father's bud valley and the father's dry valley is 1:1.
  • the seeding method of the live broadcast includes a live broadcast of the parent and the parent, a live broadcast of the parent and the parent, or a live broadcast of the parent.
  • the screened pore size is 2.1 mm wide and 20 mm long.
  • the invention has the following beneficial effects: the above-mentioned rice small-sized sterile line breeding method uses C815S as the female parent, Qigui B as the male parent to obtain the hybrid F1, and F1 and H155S as the female parent to obtain the hybrid F1', Directed cultivation of rice small-sized sterile lines.
  • the rice small-sized sterile line prepared by the above-mentioned breeding method of the sterile line has the advantages of high out-of-grain rate, high germination potential and seedling rate, strong storage resistance, and low black powder rate.
  • FIG. 1 is a comparison diagram of grain shapes of Zhuo 201S and Wanghui 091 according to a preferred embodiment of the present invention
  • FIG. 2 is a seedling stage diagram of a mixed live seeding system of Zhuo 201S and Wanghui 780 according to a preferred embodiment of the present invention
  • Figure 3 is a diagram showing the seedling stage of Zhuo 201S and Xinhui 1998 in a preferred embodiment of the present invention
  • FIG. 4 is a mature stage diagram of the mixed seed production of Zhuo 201S and Wanghui 780 according to a preferred embodiment of the present invention
  • Figure 5 is a comparison of the maturity of Zhuo 201S, N111S and Wanghui 091 in a preferred embodiment of the present invention.
  • a preferred embodiment of the present invention provides a method for breeding a rice small-sized male sterile line comprising the following steps:
  • hybrid Hybrid F1 was obtained, F1 was planted and hybridized with H155S as the female parent, and the re-crossed hybrid F1' was obtained.
  • the replanted hybrid F1′ was obtained to obtain the F2′ generation, in the F2′ generation and The F3' generation isolates the seeds of the plant with the ideal plant type, strong tillering ability, short plant and small grain size, and then the seeds of the F5' generation and above obtained by the directional cultivation of more than 2 generations are the rice small-sized infertility. system.
  • the invention has the following beneficial effects: the above-mentioned rice small-grain type sterile line breeding method, the hybrid F1 is obtained by hybridization of C815S as the female parent and Qigui B as the male parent, and the hybrid F1′ is obtained by hybridizing F1 and H155S into the female parent.
  • the rice small-sized sterile line prepared by the above-mentioned breeding method of the sterile line has the advantages of high out-of-grain rate, high germination potential and seedling rate, strong storage resistance, and low black powder rate.
  • each generation from the F1′ generation alternately under the condition of short-day low temperature and long-day low temperature, and the short-day low temperature condition is that the illumination length is less than 12 hours, and the daily average temperature is lower than At 23 ° C, the conditions of long day low temperature are that the illumination length is greater than 12 hours and the daily average temperature is below 23 ° C.
  • the conditions of short-day low temperature and long-day low temperature can be achieved by selecting suitable breeding bases and breeding seasons, such as short-term low temperature in Hainan, long-term low temperature in Changsha summer, and short-day low temperature in Changsha autumn.
  • suitable breeding bases and breeding seasons such as short-term low temperature in Hainan, long-term low temperature in Changsha summer, and short-day low temperature in Changsha autumn.
  • the experimental equipment was adjusted.
  • H155S was used as the female parent
  • the two-line sterile line C815S and Qigui B hybrid F1 were used as the male parent to cross the Hainan short-term low temperature, Changsha autumn short-term low temperature stress selection and human control.
  • the water temperature pool fertility pressurization selection through the 5th and 9th generation of directional cultivation, in 2016, the small-grain dual-purpose genic male sterile line Zhuo 201S with high quality, ideal plant type and low temperature of infertility was bred.
  • Zhuo 201S is deposited in the China Center for Type Culture Collection.
  • the Latin name is Oryza sativa L.Zhuo 201S, the deposit date is August 12, 2016, and the deposit number is CCTCC NO: P201615.
  • the shuttle breeding under short-term low temperature and long-day low temperature conditions can ensure that the low temperature sensitive genic male sterile line is stable under more complicated ecological conditions. Fertility conversion characteristics. In Hainan, Sanya, February and March In the short-day low temperature conditions, the selected sterile lines showed lower temperature at the starting point in Hunan long-day conditions, and even the plants that were difficult to breed had the characteristics of light and temperature linkage.
  • the rice granule-type sterile line is a seed of the F5'-F9' generation.
  • the rice small-sized sterile line has a thousand-grain weight of 8 to 22 g and a grain thickness of less than 2.1 mm.
  • Another aspect of the invention provides a method for seed production of hybrid rice comprising the following steps:
  • the rice large-grain type restorer line was used as the male parent and the above-mentioned rice small-type sterile line was used as the female parent group to simultaneously seed and seed production.
  • the husband and the mother are mixed and harvested at the same time.
  • the parent and female seeds are screened and separated, and the parent seeds are hybrids.
  • the hybrids produced by the rice small-type sterile line and the rice large-type restorer line are controlled by the genotype of the rice small-sized sterile line, and the male parent is controlled by the genotype of the large-sized rice restorer line. of. If the parent has a recovery weight of 26g or even more weight, you can mix the sowing or sowing, at least the harvesting (parent, maternal), especially mechanized mixed harvesting, and the harvested mixed seeds are divided by a suitable mesh sieve.
  • Sorting distinguishing between hybrids and male parents, the principle is that when hybrids are screened, small seeds (maternal hybrids) with a grain thickness of less than 2.1 mm pass through the mesh, while normal and large seeds Sorting is not possible through the mesh screen, and the sorting can be performed manually or mechanically. Since the small particles are controlled by a single recessive gene, the hybrids are sturdy and large after sowing, and thus do not affect the production and expression of the hybrid rice.
  • the rice small-sized sterile line is used for seed production, and the rice large-grain type restorer line with large grain size and suitable growth period is matched, and the male parent and the female parent are simultaneously broadcasted, so that the seed production method is simplified, no seedling transplanting is needed, labor is saved, and The pollination process can be simplified (or no supplementary pollination is required), and the parents can mix and harvest at the same time. After the parents have spent the flowers, they do not need to separately harvest the male parent, but use the harvester to mix and harvest at the same time to reduce the production cost.
  • the fathers and hybrids mixed after harvesting use the principle of separating the 1000-grain weight and the different grain size by using the sieve.
  • the sieve holes of different sizes By designing the sieve holes of different sizes, the small-grain hybrids and large grains or grains on the small-type sterile lines of rice can be obtained. Normal recovery is separated.
  • the invention selects the rice small-sized sterile line and the rice large-sized type restore line with similar initiality and large difference in grain type, and uses the difference of grain thickness to screen, for example, the specific mesh type can be designed by using the sorting machinery modification technology. Parents mechanized sorting to achieve the full mechanization of seed production.
  • the 1000-grain weight of the rice cultivar cultivar Zhuo 201S bred during the breeding of rice cultivar is a 14.6 g, while the current rice recovery line has a 1000-grain weight of more than 25 g, which is more different and easier to separate. Ensure the purity of the hybrids.
  • the rice seedlings are used for seed production.
  • the breeding coefficient of hybrids can be improved, which not only saves the production of rice fields, but also reduces the cost of farmers purchasing hybrid rice seeds.
  • it can reduce labor input, simplify seed production procedures, and reduce production costs.
  • Small-grain combination is also an effective way to improve the quality of hybrid rice. Small grain seed filling is fast and the degree of filling is good, which can effectively reduce the occurrence of chalkiness, especially in the high-temperature and high-humidity conditions of the early indica rice filling stage. The advantages are very obvious.
  • the outcrossing rate is 21.7% higher than Y58S, and the seed production is 18.6% higher.
  • the germination potential and seedling rate of hybrid seeds were high, and the germination potential and seedling rate were 6.9% and 7.3% higher than that of Y58S, respectively.
  • Hybrid seeds have strong storage resistance and are stored at room temperature for 1 year. The seedling rate is 87.4%, which is 33.9% higher than Y58S.
  • Hybrid seeds had less black disease and the rate of black powder infection was 77.6% less than that of Y58S.
  • Y58S is a variety selected by the Hunan Hybrid Rice Research Center and applied for the protection of new plant variety rights.
  • the rice large-grain type restorer line includes R9311, Wanghui 780, Xinhui 1998 and Wanghui 091.
  • Wanghui 780 is preserved in the China Center for Type Culture Collection. Its Latin name is Oryza sativa L. Wanghui 780, the deposit date is August 12, 2016, and the deposit number is CCTCC NO: P201616. Wanghui 780 is obtained through breeding.
  • the breeding process includes the following steps: hybridization of R9311 and ⁇ 498 to obtain hybrid F1”, breeding F1”, F2”, mixed, and screening thousands of grains in the F3”-F5 generation for more than 30g, grain Seeds with a thickness greater than 2.2 mm and excellent rice quality are planted for the next generation, and the resulting F6" or higher seeds are rice large-grain recovery lines.
  • the final selection is Wanghui 780.
  • Wanghui 780 has a thousand-thickness, about 34 grams, a grain thickness of more than 2.3mm, and the advantages of excellent rice quality and strong combining ability.
  • Yangdao 6 is a conventional japonica rice variety selected by the agricultural department of Lixiahe District, Jiangsu province. It was approved by Shaanxi Province in 2003 and Shaanxi Judao 2003001, which is a commercially available product.
  • Xinhui 1998 and Wanghui 091 have been selected for Hunan Hope Seed Industry Technology Co., Ltd., and have applied for the protection of new plant variety rights. If the mixture is not filtered, all the seeds obtained by planting F1" are planted, and all the seeds obtained by the same F2" are planted.
  • the rice large-sized restorer has a thousand particles larger than 30 g and a grain thickness of 2.2 mm or more.
  • the sow form of the male parent and the female parent includes the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the female parent, the male parent, the male parent
  • the seeding method is the female parent
  • the proportion of the father's bud valley and the father's dry valley is 1:1.
  • the method of seeding includes a live broadcast of the parent and the parent, a live broadcast of the parent and the parent, or a live broadcast of the parent.
  • seeding can be performed by any of the above methods.
  • the paternal and maternal broadcasts have a long duration, it is preferred to use the sow mode of the male parent and the female parent.
  • the paternal duration of the paternal is relatively late, and the "920" can be sprayed on the male parent. Adjust the flowering period of the parents to make the father and the mother meet each other. Other methods of sowing, obviously inconvenient to operate, are not conducive to adjusting the flowering period of the father. As shown in Fig.
  • the rice small-type sterile line Zhuo 201S and the rice large-type restorer line Wanghui 780 were mixed and live seedling stage.
  • the parents grew normally and vigorously, and the female parent grew around the male parent to facilitate the acceptance of pollen.
  • Zhuo 201S and Xinhui 1998 were seeded at the seedling stage, and the parents were growing neatly and the quality of the seeding was good.
  • the screened pore size is 2.1 mm wide and 20 mm long.
  • the sieve of the aperture can effectively screen and separate the parent seed and the female seed.
  • the breeding process of the small-sized sterile line is as follows:
  • Zhuo 201S showed male sterility in heading and flowering in Hunan from the end of June to the beginning of July to the beginning of August. In Sanya, Hainan, heading and flowering from the end of March showed male sterility.
  • Zhuo 201S uses 23.0 ° C, 23.5 ° C, 24.0 ° C The temperature was treated for 5 days at a constant temperature (Table 3), and the fertile pollen rate was zero, indicating that the sterile point temperature of Zhuo 201S was lower than 23.0 °C.
  • the pollen fertility identification results of Zhuo 201S are shown in Table 3.
  • Zhuo 201S is planted in early Sanya in Sanya, planted on January 28, transplanted around February 28, generally starting on April 24, and the beginning of the broadcast is about 84 days; early rice planting in Changsha, May 26 Sowing on the day, moving around June 12 Planted, before the August 13th, the beginning of the sowing, the beginning of the calendar period of about 79 days.
  • Zhuo 201S plant height is about 70cm, plant type is moderately dispersed, the early leaves are planted, the latter three leaves are more upright, the blade leaves are 25 cm long and 1.5 cm wide, the angle is around 45 °C, the leaves are light green, the leaf sheath and the tip All are colorless.
  • the stalks are medium in size and strong in tillering ability.
  • the effective ear diameter per plant is 17-19, the average ear is 15, the ear length is 28 cm, the average per panicle is 178, the valley length is 6.4 mm, and the aspect ratio is 3.3.
  • the grain is full, and the 1000-grain weight is about 14.6 grams.
  • Zhuo 201S has good outcrossing characteristics, large panicle shape, slightly curved when heading flowering, 4-5 days of flowering period, early flowering time, flowering period is 10:30-12:00, flowering peak is obvious, noon Flowers account for more than 75%, and the flower encounter rate is high.
  • the rate of spike neck is about 10%, and the stigma exsertion rate is high.
  • the total exudation rate of stigma is 88.2%, of which the bilateral exsertion rate is 62.2%, the stigma is strong, and the outcrossing rate is 80.
  • About % the seed production is easy to produce. It is more sensitive to "Nine-two". When the heading is 15-20%, 15 grams of "92" per acre can be used to remove the neck.
  • the brown rice rate was 80.3%
  • the polished rice rate was 71.8%
  • the whole polished rice rate was 59.1%
  • the grain length was 6.1mm
  • the aspect ratio was 3.3
  • the white grain rate was 11.0. %, chalkiness 1.0%, transparency grade 1, alkali value 6.0, gel consistency 65 mm, linear ingot powder content 15.8%, protein content 9.5%.
  • Zhuo 201S In 2015, it was identified by rice blast in Dawei Mountain, Liuyang City, with 3 nursery grades, 5 grades of spikes, and a comprehensive rating of 3.1; resistance to bacterial blight was grade 5;
  • Fig. 1 The comparison between the granules of Zhuo 201S and Wanghui 091 is shown in Fig. 1.
  • the small granules are Zhuo 201S
  • the large granules are the recovery system of Hunan Hope Seed Industry Technology Co., Ltd. Conducive to the separation of the male and female seeds.
  • the control experiment was carried out under the same conditions with Y58S as the female parent while seeding with Zhuo 201S as the female parent.
  • the outcrossing rate and seed production yield of the group with Zhuo 201S as the female parent and the group with Y58S as the female parent were counted, and the hybrids prepared by each group were planted separately, and the germination potential of each group of hybrids was counted. Seedling rate, seed splitting rate and black powder infection rate.
  • Table 4 The statistical results of the relevant data of each group are shown in Table 4:
  • the outcrossing rate of the Zhuo 201S group was 21.7% higher than that of the Y58S group, and the seed production was 18.6% higher than that of the Y58S group.
  • the germination potential and seedling rate of the hybrid seeds prepared by Zhuo 201S group were high, and the germination potential and seedling rate were 6.9% and 7.3% higher in the Y58S group, respectively.
  • the seed splitting rate of the hybrid seeds prepared by Zhuo 201S group was 1.47%, which was 8.23% less than that of the Y58S group, and the hybrid seeds prepared by Zhuo 201S group were not infected with smut, while the black powder disease rate of the Y58S group was 13.64%.
  • the control experiment was carried out under the same conditions with N111S as the female parent while seeding with Zhuo 201S as the female parent.
  • the outcrossing rate and seed production yield of the group with Zhuo 201S as the female parent and the group with N111S as the female parent were counted as shown in Fig. 5, and the left side was used.
  • the hybrids prepared by each group were planted separately, and the germination potential, seedling rate, seed splitting rate and black powder disease infection rate of each group of hybrids were counted.
  • the statistical results of the relevant data of each group are shown in Table 5:
  • the outcrossing rate of Zhuo 201S group was 58.5% higher than that of N111S group, and the seed production was 53.3% higher than that of N111S group.
  • the germination potential and seedling rate of the hybrid seeds prepared by Zhuo 201S group were high, and the germination potential and seedling rate were 9.5% and 13.1% higher respectively in the N111S group.
  • the seed splitting rate of hybrid seeds prepared by Zhuo 201S group was 2.54%, which was 33.06% less than that of N111S group, and the hybrid seeds prepared by Zhuo 201S group were not infected with smut, while the black powder disease rate of N111S group was 15.12%.
  • the control experiment was carried out under the same conditions with Y58S as the female parent while seeding with Zhuo 201S as the female parent.
  • the outcrossing rate and seed production yield of the group with Zhuo 201S as the female parent and the group with Y58S as the female parent were counted, and the hybrids prepared by each group were planted separately, and the germination potential of each group of hybrids was counted. Seedling rate, seed splitting rate and black powder infection rate.
  • Table 6 The statistical results of the relevant data of each group are shown in Table 6 below:
  • the outcrossing rate of the Zhuo 201S group was 17.6% higher than that of the Y58S group, and the seed production was 11.5% higher than that of the Y58S group.
  • the germination potential and seedling rate of the hybrid seeds prepared by Zhuo 201S group were high, and the germination potential and seedling rate were 5.8% and 5.4% higher in the Y58S group, respectively.
  • the seed splitting rate of the hybrid seeds prepared by Zhuo 201S group was 1.15%, which was 10.55% less than that of Y58S group, and the hybrid seeds prepared by Zhuo 201S group were not infected with smut, while the black powder rate of 8.26% in Y58S group.
  • Example 2 is a preferred embodiment.

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Abstract

本发明公开了一种水稻小粒型不育系的育种方法及杂交水稻的轻简制种方法，所述水稻小粒型不育系的育种方法包括以下步骤：以C815S为母本，七桂B为父本进行杂交得到杂种F1，种植F1并与H155S为母本进行杂交，得到杂种F1'，种植F1'得到F2'世代，在F2'、F3'世代分离出株型理想、分蘖力强、植株较矮、粒型小的单株，再经2个世代以上的定向培育得到水稻小粒型不育系。上述不育系的育种方法培育的水稻小粒型不育系具有异交结实率高、发芽势和成苗率高、耐储存性强、黑粉病率低的优点。在此基础上，水稻小粒型不育系与大粒型恢复系制种，父本和母本同时混合收割，根据谷粒厚度的差异进行分选，从而实现杂交水稻轻简化制种。

Description

水稻小粒型不育系的育种方法及杂交水稻的轻简制种方法 技术领域
本发明涉及水稻制种领域，特别地涉及一种水稻小粒型不育系的育种方法。此外，本发明还涉及一种包括杂交水稻的轻简制种方法。
背景技术
杂交水稻的选育成功并大面积推广，在促进粮食稳定增产，确保我国粮食安全上发挥了重要作用。传统杂交水稻制种需要分别将父母本播种、移栽、进行不同田间管理才能获得理想的花期相遇状态，需要人工授粉、父母本人工分开收割这一方式工序繁琐、劳动强度大、成本高。而近些年来随着我国社会经济和城镇化发展，农业劳力日益紧缺，使得杂交水稻制种的这种劳动力密集型、精耕细作的方式缺少劳动力执行，将严重影响了杂交水稻种子的安全生产与供应，亟需集成研究实现杂交水稻轻简高效规模化制种的方法，以大幅减少劳力投入和劳动强度，提高工作效率，以确保杂交水稻种子的安全高效生产及供应，继续保持我国在杂交水稻技术领域的国际领先地位。
目前国内外在杂交水稻机械化制种技术研究方面部分环节实现了机械化作业。美国水稻技术公司、德国拜耳公司等在父母本的田间管理上采用机械旱条播、收获，机械或农用飞机喷药、施肥，直升机赶粉等制种技术，实现了大面积半机械化制种。韩国、日本分别在80-90年代在耕翻、育秧、植保、收获、干燥等生产环节基本实现机械化。国内在土地耕整，母本机械条播和机械插秧、喷施农药及赤霉素、收割、烘干等环节实现机械操作。目前国外正在研究父母本播始历期一致的强优组合，母本与父本之间的籽粒颜色、粒重或粒形有明显差异，如采用水稻小粒型不育系实现制种时父母本混播混收的技术来实现杂交水稻机械化制种，但还没有取得研究进展。而杂交水稻机械化父母本混植制种技术的关键是实现父母本种子机械化分选。迄今为止，受不育系的限制，并无一种合适的机械化分选方法在生产中得到推广应用。
发明内容
本发明提供了一种水稻小粒型不育系的育种方法及杂交水稻的轻简制种方法，以解决制种时父母本混播混收的技术问题。
本发明采用的技术方案如下：
本发明一方面提供了一种水稻小粒型不育系的育种方法，包括以下步骤：
以C815S为母本，七桂B为父本进行杂交得到杂种F1，种植F1并与H155S为母本进行杂交，得到复交杂种F1′，种植复交杂种F1′得到F2′世代，在F2′世代和F3′世代分离 出株型理想、分蘖力强、植株矮、粒型小的单株所结种子进行种植，再经2个世代以上的定向培育所得F5′世代以上种子即为水稻小粒型不育系。
进一步地，在育种过程中，自F1′世代起各世代逐代交替在短日低温、长日低温的条件下胁迫选择，短日低温的条件为光照长度小于12小时、日平均温度低于23℃，长日低温的条件为光照长度大于12小时、日平均温度低于23℃。
进一步地，水稻小粒型不育系为F5′～F9′世代的种子。
进一步地，水稻小粒型不育系的千粒重为8～22g且籽粒厚度小于2.1mm。
本发明另一方面一种杂交水稻的轻简制种方法，包括以下步骤：
以水稻大粒型恢复系为父本和上述的水稻小粒型不育系为母本配组同时直播制种。
对父本和母本同时混合收割。
根据谷粒厚度的差异，筛选分离父本种子和母本种子，母本种子为杂交种。
可选地，水稻大粒型恢复系包括R9311、望恢780、新恢1998和望恢091。
望恢780通过育种得到，育种过程包括以下步骤：
用R9311与蜀恢498杂交得到杂种F1”，繁殖F1”、F2”，混收，在F3”～F5”世代均筛选千粒重大于30g、谷粒厚度大于2.2mm且米质优的株系所结种子种植下一世代，所得F6”代以上种子即为水稻大粒型恢复系。
可选地，水稻大粒型恢复系的千粒重大于30g且籽粒厚度为2.2mm以上。
可选地，父本和母本的播种形式包括父本芽谷母本干谷、父本干谷母本芽谷、父本母本均播芽谷或父本母本均播干谷。
其中播种方式为父本芽谷母本干谷时，父本芽谷和父本干谷的比例为1：1。
可选地，直播的播种方式包括父本和母本混直播、父本和母本条直播或父本抛秧母本直播。
可选地，筛选的筛孔孔径为2.1毫米宽，20毫米长。
本发明具有以下有益效果：上述水稻小粒型不育系的育种方法，以C815S为母本，七桂B为父本进行杂交得到杂种F1，F1与H155S为母本进行杂交得到杂种F1′，经定向培育得到水稻小粒型不育系。上述不育系的育种方法制得的水稻小粒型不育系具有异交结实率高、发芽势和成苗率高、耐储存性强、黑粉病率低的优点。
除了上面所描述的目的、特征和优点之外，本发明还有其它的目的、特征和优点。下面将参照图，对本发明作进一步详细的说明。
附图说明
构成本申请的一部分的附图用来提供对本发明的进一步理解，本发明的示意性实施例及其说明用于解释本发明，并不构成对本发明的不当限定。在附图中：
图1是本发明优选实施例的卓201S与望恢091的粒型对比图；
图2是本发明优选实施例的卓201S与望恢780混直播制种苗期图；
图3是本发明优选实施例的卓201S与新恢1998条播制种苗期图；
图4是本发明优选实施例的卓201S与望恢780混播制种成熟期图；
图5是本发明优选实施例的卓201S、N111S分别与望恢091制种成熟期对比图。
具体实施方式
以下结合附图对本发明的实施例进行详细说明，但是本发明可以由权利要求限定和覆盖的多种不同方式实施。
参照图1，本发明的优选实施例提供了一种水稻小粒型不育系的育种方法，包括以下步骤：
以C815S为母本，七桂B为父本进行杂交得到杂种F1，种植F1并与H155S为母本进行杂交，得到复交杂种F1′，种植复交杂种F1′得到F2′世代，在F2′世代和F3′世代分离出株型理想、分蘖力强、植株矮、粒型小的单株所结种子进行种植，再经2个世代以上的定向培育所得F5′世代以上种子即为水稻小粒型不育系。
本发明具有以下有益效果：上述水稻小粒型不育系的育种方法，以C815S为母本、七桂B为父本进行杂交得到杂种F1，F1与H155S为母本进行杂交得到杂种F1′，经定向培育得到水稻小粒型不育系。上述不育系的育种方法制得的水稻小粒型不育系具有异交结实率高、发芽势和成苗率高、耐储存性强、黑粉病率低的优点。
可选地，在育种过程中，自F1′世代起各世代逐代交替在短日低温、长日低温的条件下胁迫选择，短日低温的条件为光照长度小于12小时、日平均温度低于23℃，长日低温的条件为光照长度大于12小时、日平均温度低于23℃。
短日低温、长日低温的条件可通过选择合适的育种基地及育种季节达到，如海南为短日低温、长沙夏季为长日低温、长沙秋季为短日低温，也在室内的实验环境可通过实验设备调节达到。
以前一种方式为例，2011年夏季用H155S为母本，用两系不育系C815S与七桂B杂交F1为父本杂交，经海南短日低温、长沙秋季短日低温胁迫选择及人控水温池育性增压选择，通过5年9代的定向培育，于2016年育成具有优质、株型理想、不育起点温度较低的小粒型两用核不育系卓201S。卓201S保藏于中国典型培养物保藏中心，其拉丁文名称为Oryza sativa L.Zhuo 201S，保藏日期为2016年8月12日，保藏编号为CCTCC NO:P201615。根据水稻两用核不育系在海南和湖南的育性转换特点，开始短日低温和长日低温条件下穿梭育种，可确保育成的低温敏核不育系在较复杂的生态条件下有稳定的育性转换特性。在海南三亚2、3月 份短日低温条件下当选的不育系，在湖南长日条件下起点温度表现为更低的，甚至难以繁殖的株系，具有光温连动特点。
可选地，水稻小粒型不育系为F5′～F9′世代的种子。
可选地，水稻小粒型不育系的千粒重为8～22g且籽粒厚度为小于2.1毫米。
本发明另一方面提供了一种杂交水稻的轻简制种方法，包括以下步骤：
以水稻大粒型恢复系为父本和上述水稻小粒型不育系为母本配组同时直播制种。
对父本和母本同时混合收割。
根据谷粒厚度的差异，筛选分离父本种子和母本种子，母本种子为杂交种。
该水稻小粒型不育系与水稻大粒型恢复系生产出的杂交种受母体即水稻小粒型不育系的基因型控制是小粒的，父本种子受水稻大粒型恢复系的基因型控制是大粒的。如果父本千粒重在26g甚至更大粒重的恢复系，则可以混合播种或条播，至少可以混合收割(父、母本)，尤其是机械化混合收割，收割后的混合种子通过合适的网筛进行分拣，区分杂交种和父本，其原理是杂交种过网筛清选时，因谷粒厚度小于2.1毫米的小粒种子(母本上结的杂交种)通过网筛，而正常粒和大粒种子无法通过网筛，从而实现分选，具体分选时可采用人工或机械进行分选。由于小粒受单一隐性基因控制，杂交种播种后结实为大粒，因而不影响杂交稻产量的发挥和表达。
利用水稻小粒型不育系进行制种，与籽粒大粒、生育期合适的水稻大粒型恢复系配组，父本与母本同时直播，使得制种方式简化、无需育秧移栽、节省劳力，并且可以简化授粉环节(或者不需要辅助授粉)，父母本同时混合收割，在父母本扬花过后无需单独收获父本，而是用收割机同时混合收割，降低生产成本。
收获后混合的父本及杂交种利用网筛分离千粒重不同即籽粒大小不同的原理，通过设计大小不同的筛孔，可以将水稻小粒型不育系上所结的小粒型杂交种与大粒或籽粒正常的恢复系分离出来。本发明通过选育播始历期相近、粒型差异大的水稻小粒型不育系和水稻大粒型恢复系，并利用籽粒厚度区别筛选，如可利用分选机械改装技术设计特定的网筛对父母本机械化分选，实现种子生产的全程机械化。
尤其，水稻小粒型不育系的育种过程中育成的水稻小粒型不育系卓201S的千粒重为14.6g，而当前的水稻恢复系籽粒千粒重一般在25g以上，差异更大，更易于分离，可保证杂交种的纯度。
并且用水稻小粒型不育系进行制种，一方面可提高杂交种的繁殖系数，不仅节约制种稻田，而且可减少农民用于购买杂交稻种子的成本。另一方面，可减少劳动力的投入、简化制种程序，降低生产成本。小粒型组合也是提高杂交水稻米质的有效途径，小粒种子灌浆快，充实度好，可有效降低垩白的发生，尤其在南方早籼稻灌浆期高温高湿气候条件下，小粒型种子的这种优势表现得十分明显。
同时，本申请的杂交水稻的轻简制种方法中，杂交制种时，异交结实率比Y58S高21.7％，制种产量高18.6％。杂交种子发芽势和成苗率高，与同一个父本配组，其发芽势和成苗率分别比Y58S高6.9％、7.3％。杂交种子耐储存性强，常温条件保存1年，其成苗率达到87.4％，比Y58S高33.9％。杂交种子黑份病少，其感染黑粉病率比Y58S少77.6％。Y58S为湖南杂交水稻研究中心选育的品种，并且申请了植物新品种权保护。
可选地，水稻大粒型恢复系包括R9311、望恢780、新恢1998和望恢091。望恢780保藏于中国典型培养物保藏中心，其拉丁文名称为Oryza sativa L.Wanghui 780，保藏日期为2016年8月12日，保藏编号为CCTCC NO:P201616。望恢780通过育种得到，育种过程包括以下步骤：用R9311与蜀恢498杂交得到杂种F1”，繁殖F1”、F2”，混收，在F3”～F5”世代均筛选千粒重大于30g、谷粒厚度大于2.2mm且米质优的株系所结种子种植下一世代，所得F6”代以上种子即为水稻大粒型恢复系。最终选育得到望恢780，望恢780具有千粒重大，达到34克左右，谷粒厚度大于2.3mm，并且米质优，配合力强的优点。
R9311即扬稻6号是江苏里下河地区农科所选育的常规籼稻品种，2003年通过陕西省审定，陕审稻2003001，为市售产品。新恢1998、望恢091为湖南希望种业科技股份有限公司选育，已经申请了植物新品种权保护。混收即不进行筛选，种植F1”所得所有种子均进行种植，同样的F2”所得所有种子均进行种植。
可选地，水稻大粒型恢复系的千粒重大于30g且籽粒厚度为2.2mm以上。
可选地，父本和母本的播种形式包括父本芽谷母本干谷、父本干谷母本芽谷、父本母本均播芽谷或父本母本均播干谷。其中播种方式为父本芽谷母本干谷时，父本芽谷和父本干谷的比例为1：1。
可选地，播种的方式包括父本和母本混直播、父本和母本条直播或父本抛秧母本直播。当父本和母本的播始历期相近时，可采用上述任一种方式进行播种。当父本和母本的播始历期较大时，优选采用父本和母本条播的播种方式，一般父本的播始历期相对较晚，可通过对父本喷洒“920”等措施调节父母本的花期，使父本和母本花期相遇。而其他播种方式，显然操作有所不便，不利于调节父本的花期。如图2所示，水稻小粒型不育系卓201S与水稻大粒型恢复系望恢780混直播制种苗期，父母本均生长正常且旺盛，母本生长在父本周围，便于接受花粉。如图3所示，卓201S与新恢1998条播制种苗期，父母本生长整齐，条播质量好。
可选地，筛选的筛孔孔径为2.1毫米宽，20毫米长。该孔径的筛网可有效的筛选分离筛选分离父本种子和母本种子。
实施例1
小粒型不育系的选育过程如下：
2011年夏季用H155S为母本，用自育的两系不育系C815S与七桂B杂交F1为父本杂交，经海南短日低温、长沙夏季长日低温、长沙秋季短日低温胁迫选择及人控水温池育性增压选 择，通过5年9代的定向培育，于2016年育成具有米质优、株型理想、不育起点温度较低的小粒型两用核不育系卓201S。选育过程如表1所示。
表1卓201S选育流程图
Figure PCTCN2017093249-appb-000001
三、特征特性
1、育性表现
从表2可知，卓201S在湖南从6月底7月初抽穗开花至8月下旬抽穗开花均表现雄性不育。在海南三亚从3月底左右抽穗开花表现雄性不育。卓201S用23.0℃、23.5℃、24.0℃的 恒温分别处理5天(表3)，可育花粉率均为零，这表明卓201S的不育起点温度低于23.0℃。卓201S的花粉育性鉴定结果如表3所示。
表2卓201S在湖南、海南两地生态实用性鉴定试验中的育性表现
Figure PCTCN2017093249-appb-000002
表3水稻两用核不育系卓201S花粉育性鉴定结果
Figure PCTCN2017093249-appb-000003
2、生育特性
卓201S在三亚地区作早稻种植，1月28日播种，2月28日左右移栽，一般在4月24日始穗，播始历期84天左右；在长沙地区作早稻种植，5月26日播种，6月12日左右移 栽，8月13日前齐穗，播始历期79天左右。
3、形态特征和农艺性状
卓201S株高约70cm，株型紧散适中，前期叶片植披，后三叶较直立，剑叶长25厘米，宽1.5厘米，夹角在45℃左右，叶色淡绿，叶鞘、稃尖均无色。茎秆粗细中等，分蘖力强，一般单株有效穗17～19个，平均成穗15个，穗长28厘米左右，平均每穗颖花178个左右，谷长6.4毫米，长宽比3.3，谷粒饱满，千粒重14.6克左右。
4、异交特性
卓201S具有良好的异交特性，穗形较大，抽穗开花时稍呈弧形，盛花期4-5天，花时早，盛花期时间在10：30-12：00，开花高峰明显，午前花占75％以上，花时相遇率高。穗包颈率10％左右，柱头外露率高，在不喷施九二0条件下，柱头总外露率为88.2％，其中双边外露率62.2％，柱头生活力强，异交结实率可达80％左右，制种易高产。对“九二0”较敏感，在抽穗15-20％时，每亩喷施15克“九二0”可解除包颈。
5、稻米品质
2015年冬季在海南三亚市海棠湾镇繁殖的卓201S稻谷，经分析，糙米率80.3％，精米率71.8％，整精米率59.1％，粒长6.1mm，长宽比3.3，垩白粒率11.0％，垩白度1.0％，透明度1级，碱消值6.0级，胶稠度65mm，直链锭粉含量15.8％，蛋白质含量9.5％。
6、卓201S抗逆性
卓201S 2015年经浏阳市大围山稻瘟病鉴定，苗瘟3级，穗瘟5级，综合评级3.1级；白叶枯病抗性5级；纹枯病轻。
7、卓201S的粒型
取卓201S与望恢091进行粒型对比如图1所示，其中小粒为卓201S，大粒为湖南希望种业科技股份有限公司选育的恢复系望恢091，可见二者厚度存在明显差异，有利于父本种子和母本种子分离。
实施例2
2016年4月在海南三亚海棠湾杂交制种时，以大粒型父本望恢780为父本，卓201S为母本。父、母本1月25日同时混播，父母本比例为1：20，父本干谷和母本芽谷比例为1：1；父母本4月18日同时始穗，4月20日父母本同时混合施用“九二零”15克/亩，有微风即不 赶粉，无风状态下人工辅助授粉1次，5月20日成熟，如图4所示，卓201S与望恢780混播制种成熟期，父本占的空间小，比例少，制种异交结实率高。5月20日混合收割。在筛孔孔径为2.1毫米宽，20毫米长的条件下，进行机械筛选，顺利分离父本种子和母本种子。母本种子即杂交种的纯度为99.76％。
在以卓201S为母本进行制种的同时，以Y58S为母本在相同条件下进行对照实验。统计以卓201S为母本的组别和以Y58S为母本的组别的异交结实率、制种产量，并取各组制得的杂交种分别种植，统计各组杂交种的发芽势、成苗率、种子裂颖率和黑粉病感染率。各组的相关数据的统计结果如表4所示：
表4不育系与望恢780的轻简制种结果
Figure PCTCN2017093249-appb-000004
在相同条件下，以卓201S组的异交结实率比Y58S组高21.7％，制种产量比Y58S组高18.6％。卓201S组制得的杂交种子发芽势和成苗率高，其发芽势和成苗率分别Y58S组高6.9％、7.3％。卓201S组制得的杂交种子的种子裂颖率为1.47％，比Y58S组少8.23％，并且卓201S组制得的杂交种子不感染黑粉病，而Y58S组感染黑粉病率13.64％。
实施例3
2016年4月在海南三亚海棠湾杂交制种时，以大粒型父本望恢091为父本，卓201S为母本，N111S为母本对照。父本1月15日用抛秧盘播种、母本1月25日直播，父母本比例为1：20；父母本4月18日同时始穗，4月21日父母本同时混合施用“九二零”15克/亩，有微风即不赶粉，无风状态下人工辅助授粉1次，5月20日成熟混合收割。在筛孔孔径为2.1毫米宽，20毫米长的条件下，进行机械筛选，顺利分离父本种子和母本种子。母本种子即杂交种的纯度为99.51％。
在以卓201S为母本进行制种的同时，以N111S为母本在相同条件下进行对照实验。统计以卓201S为母本的组别和以N111S为母本的组别的异交结实率、制种产量，如图5所示，左侧为。并取各组制得的杂交种分别种植，统计各组杂交种的发芽势、成苗率、种子裂颖率和黑粉病感染率。各组的相关数据的统计结果如表5所示：
表5不育系与望恢091的轻简制种结果
Figure PCTCN2017093249-appb-000005
Figure PCTCN2017093249-appb-000006
在相同条件下，以卓201S组的异交结实率比N111S组高58.5％，制种产量比N111S组高53.3％。卓201S组制得的杂交种子发芽势和成苗率高，其发芽势和成苗率分别N111S组高9.5％、13.1％。卓201S组制得的杂交种子的种子裂颖率为2.54％，比N111S组少33.06％，并且卓201S组制得的杂交种子不感染黑粉病，而N111S组感染黑粉病率15.12％。
实施例4
2016年4月在海南三亚海棠湾杂交制种时，以大粒型父本新恢1998为父本，卓201S为母本。父、母本1月25日同时条播，父母本比例为2：9，父本播干谷、母本播芽谷；父母本4月18日同时始穗，4月20日父母本同时混合施用“九二零”15克/亩，每天人工辅助授粉1次，5月20日成熟混合收割。在筛孔孔径为2.1毫米宽，20毫米长的条件下，进行机械筛选，顺利分离父本种子和母本种子。母本种子即杂交种的纯度为99.05％。
在以卓201S为母本进行制种的同时，以Y58S为母本在相同条件下进行对照实验。统计以卓201S为母本的组别和以Y58S为母本的组别的异交结实率、制种产量，并取各组制得的杂交种分别种植，统计各组杂交种的发芽势、成苗率、种子裂颖率和黑粉病感染率。各组的相关数据的统计结果如下表6所示：
表6不育系与新恢1998的轻简制种结果
Figure PCTCN2017093249-appb-000007
在相同条件下，以卓201S组的异交结实率比Y58S组高17.6％，制种产量比Y58S组高11.5％。卓201S组制得的杂交种子发芽势和成苗率高，其发芽势和成苗率分别Y58S组高5.8％、5.4％。卓201S组制得的杂交种子的种子裂颖率为1.15％，比Y58S组少10.55％，并且卓201S组制得的杂交种子不感染黑粉病，而Y58S组感染黑粉病率8.26％。
综合比较实施例2～4，以卓201S为母本、望恢780为父本制得杂交种异交结实率最高、制种产量最高、成苗率最高、仅发芽势与种子裂颖率略低于实施例4。此外，卓201S为母本、望恢780的播始历期相近。综合比较之下，实施例2为最佳实施例。
以上所述仅为本发明的优选实施例而已，并不用于限制本发明，对于本领域的技术人员 来说，本发明可以有各种更改和变化。凡在本发明的精神和原则之内，所作的任何修改、等同替换、改进等，均应包含在本发明的保护范围之内。

Claims (10)

  1. 一种水稻小粒型不育系的育种方法，其特征在于，包括以下步骤：
    以C815S为母本，七桂B为父本进行杂交得到杂种F1，种植F1并与H155S为母本进行杂交，得到复交杂种F1′，种植复交杂种F1′得到F2′世代，在F2′世代和F3′世代分离出株型理想、分蘖力强、植株矮、粒型小的单株所结种子进行种植，再经2个世代以上的定向培育所得F5′世代以上种子即为水稻小粒型不育系。
  2. 根据权利要求1所述的水稻小粒型不育系的育种方法，其特征在于，在育种过程中，自F1′世代起各世代逐代交替在短日低温、长日低温的条件下胁迫选择，所述短日低温的条件为光照长度小于12小时、日平均温度低于23℃，所述长日低温的条件为光照长度大于12小时、日平均温度低于23℃。
  3. 根据权利要求1所述的水稻小粒型不育系的育种方法，其特征在于，所述水稻小粒型不育系为F5′～F9′世代的种子。
  4. 根据权利要求1所述的水稻小粒型不育系的育种方法，其特征在于，所述水稻小粒型不育系的千粒重为8～22g且籽粒厚度小于2.1mm。
  5. 一种杂交水稻的轻简制种方法，其特征在于，包括以下步骤：
    以水稻大粒型恢复系为父本和权利要求1～4中任一项所述的水稻小粒型不育系为母本配组同时直播制种；
    对父本和母本同时混合收割；
    根据谷粒厚度的差异，筛选分离父本种子和母本种子，所述母本种子为杂交种。
  6. 根据权利要求5所述的杂交水稻的轻简制种方法，其特征在于，所述水稻大粒型恢复系包括R9311、望恢780、新恢1998和望恢091；
    所述望恢780通过育种得到，育种过程包括以下步骤：
    用R9311与蜀恢498杂交得到杂种F1”，繁殖F1”、F2”，混收，在F3”～F5”世代均筛选千粒重大于30g、谷粒厚度大于2.2mm且米质优的株系所结种子种植下一世代，所得F6”代以上种子即为水稻大粒型恢复系。
  7. 根据权利要求8所述的杂交水稻轻简制种方法，其特征在于，所述水稻大粒型恢复系的千粒重大于30g且籽粒厚度为2.2mm以上。
  8. 根据权利要求5所述的杂交水稻的轻简制种方法，其特征在于，父本和母本的播种形式包括父本芽谷母本干谷、父本干谷母本芽谷、父本母本均播芽谷或父本母本均播干谷；
    其中播种方式为父本芽谷母本干谷时，所述父本芽谷和父本干谷的比例为1：1。
  9. 根据权利要求5所述的杂交水稻的轻简制种方法，其特征在于，所述直播的播种方式包 括父本和母本混直播、父本和母本条直播或父本抛秧母本直播。
  10. 根据权利要求5所述的杂交水稻的轻简制种方法，其特征在于，筛选的筛孔孔径为2.1毫米宽，20毫米长。
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