CN109997683B - Rice double haploid breeding method based on haploid induction line - Google Patents
Rice double haploid breeding method based on haploid induction line Download PDFInfo
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- A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
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- A—HUMAN NECESSITIES
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Abstract
The invention provides a haploid breeding method of rice based on a haploid induction line, and belongs to the technical field of agriculture. The invention screens out the red rice maintainer line carrying dominant red rice gene after hybridizing the rice three-line maintainer line material with the rice material with dominant red rice gene, and screens out the mutant with centromere gene function after chemical directional induction, and hybridizes the red rice three-line sterile line to obtain the maintainer line and sterile line carrying red rice gene and centromere mutant gene, and the colorless seed coat hybrid seed is reserved after hybridizing the hybrid F1 prepared according to breeding target with the red rice haploid induced sterile line; and (3) processing the colorless seed coat hybrid seeds by using colchicine to obtain a diploid material, namely forming a new strain. The method can quickly homozygize the gene, overcomes the defect of long generation of traditional crossbreeding and segregation, and is applied to rice breeding.
Description
Technical Field
The invention relates to a haploid breeding method of rice based on a haploid induction line, and belongs to the technical field of agriculture.
Background
The haploid breeding technology is to obtain purified doubled haploids (DH plants) from naturally or artificially cultured haploid plants in a chromosome group doubling mode, and due to the characteristics of rapidness, accuracy and high efficiency, breeders pay more and more attention, at present, 250 crop species apply doubled haploid breeding all over the world, and 300 doubled haploid pure lines from cultivars are cultured from l2 species. The core of the traditional breeding technology is that the favorable genes are fixed and homozygous through continuous selection, the process usually needs a plurality of generations of selfing or backcrossing processes, for example, a relatively pure line needs more than 6 generations when a traditional conventional breeding method is used for breeding a maize selfing line, and a double haploid breeding method generally only needs 2 generations, so that the breeding process is greatly accelerated. In addition, because each locus of the doubled double haploid is homozygous, the gene interaction can be simplified, the super-dominant effect can be removed, the favorable additive and additive epistatic effects can be reserved, the harmful, lethal and semi-lethal recessive genes can be eliminated, and the selection efficiency of the favorable genes can be accelerated. If the dihaploid technology is organically combined with the improvement of germplasm amplification and breeding materials, the target inducing materials are effectively recombined and improved, enough favorable gene loci are accumulated, and then the dihaploid technology is used for obtaining a pure line, so that the advantage of the dihaploid breeding can be greatly exerted.
The breeding of new varieties by utilizing a doubled haploid technology has become a trend of corn breeding development. Dupont pioneer, Monsanto, Sineda, etc. have been used as an important method for corn breeding, and are called breeding techniques "black horses". At present, the cross-country species companies all present large-scale, high-throughput and flow breeding modes in the aspect of DH breeding, and the number of DH pure lines obtained in a year reaches more than 50000.
The natural haplotypes have very low probability of occurrence, generally not exceeding 0.01%, and artificial induction is necessary to obtain a large amount of haplotypes. There are many artificial inducing methods, mainly including distant hybridization, anther culture, parthenogenesis, haploid inducing line hybridization induced haploid, etc. The methods of distant hybridization, anther culture, parthenogenesis and the like have obvious defects: the induction rate of distant hybridization is extremely low; the anther culture has the defects of low anther culture efficiency, genotype obstacle, low callus induction rate and low green seedling differentiation rate; extremely low parthenogenesis frequency, unstable effect, large difference between genotypes, small quantity of obtained seeds and the like. In view of various defects of distant hybridization, anther culture and parthenogenesis induction to generate haploids, the induction of haploids by utilizing the haploid induction line for hybridization is widely regarded. The discovery of a corn parthenogenesis haploid induction line Stock6 brings breakthrough progress for corn haploid breeding.
Disclosure of Invention
Aiming at the problems existing in the prior rice haploid induction, the invention obtains haploid inducing material with application value by directional screening of mutant gene of centromere control gene CENH3, transforms the haploid inducing material into sterile line with red seed coat marker character, utilizes the sterile line to hybridize with breeding material to be induced, screens haploid seeds with red seed coats, utilizes colchicine to realize doubling, identifies the doubled material and breeds new varieties.
The invention is realized by the following technical scheme:
a rice double haploid breeding method based on a haploid induction line comprises the following steps:
s1, selecting an excellent rice three-line maintainer line material, hybridizing the excellent rice three-line maintainer line material with a rice material with a dominant red rice gene, backcrossing a plurality of generations by using the maintainer line, breeding a red rice maintainer line carrying the dominant red rice gene, and controlling the rice seed coat to be red by using the dominant red rice gene;
s2, soaking the screened red rice maintainer line material carrying the dominant red rice gene in EMS with the concentration of 0.3-1.0% for 12-24 h to obtain the red rice maintainer line material containing the functional mutant of the rice centromere gene;
s3, directionally screening functional mutants of the centromere gene of the rice, selfing the screened mutants for 2 generations to form a stable genetic material, and identifying that the haploid induction efficiency of the mutants is higher than 5 percent to obtain the stable genetic material, wherein the haploid induction efficiency of the mutants refers to that the mutants are hybridized with a normal variety, wherein the mutants are used as female parents, and the ratio of the haploid seeds in the obtained seeds is counted;
s4, hybridizing the stable genetic material screened in the step S3 with a rice three-line sterile line, backcrossing for 5 generations by taking a red rice maintainer line mutant as a recurrent parent, and completing the replacement of the nuclear gene of the sterile line to obtain a red rice haploid induction maintainer line and a sterile line carrying a red rice gene and a centromere mutant gene;
s5, hybridizing a hybrid F1 prepared according to a breeding target with a red rice haploid induction sterile line, identifying the seed coat color of the obtained hybrid seeds, eliminating red seed coat hybrid seeds, and reserving colorless seed coat hybrid seeds;
s6, treating the achromatous seed coat hybrid seeds by using colchicine to obtain a doubled diploid material, and identifying the agronomic characters or genotypes of the diploid material to form a new strain, wherein the identification of the agronomic characters or genotypes of the diploid material refers to the investigation of economic character indexes such as the growth period, the maturing rate, the thousand seed weight, the number of ears and the number of ears of the diploid material, and the genotypes refer to the investigation of the genotypes of the quality or resistance genes of the diploid.
Preferably, the targeted screening method in S3 is to detect the GGA → GAG sequence difference of CENH3 gene by sequencing, and the material different from the control is mutant.
Preferably, the red rice maintainer line mutant is hybridized with the rice three-line sterile line in S4, the red rice maintainer line mutant is backcrossed for 5 generations by taking the red rice maintainer line mutant as a recurrent parent, the replacement of the sterile line nuclear gene is completed, red seed coat seeds are selected from each backcross generation to carry out the next backcross, and the red seed coat seeds are diploid and carry heterozygous red seed coat genes.
Preferably, the seed coat color identification of the obtained hybrid seeds is performed in S5, wherein the seeds with red seed coats are non-haploid seeds, the seeds with colorless seed coats are haploid seeds, and the screening for the seed coat color can be achieved manually or mechanically.
Preferably, the colorless seed coat hybrid seeds in S6 are treated with colchicine, which means that seeds with embryo germination are soaked for 8 hours by using colchicine with the concentration of 0.6%, and haploid doubling can be realized after soaking.
Compared with the prior art, the invention has the following beneficial effects:
1. the existing rice haploid breeding technology depends on anther tissue culture, has tissue culture genotype dependence, for example, japonica rice is easy to obtain tissue culture individuals, while the tissue culture capacity of indica rice is poor, and a lot of indica rice materials can not obtain haploids by an anther culture method, so that the application of the haploid breeding technology is greatly limited;
2. the time for obtaining the haploid by the existing anther culture technology is longer, the time for obtaining tissue culture differentiation individuals from the obtained anther is 3-4 months, the time for obtaining the haploid induction line from hybridization to haploid seeds is only 1 month, and the breeding time is obviously shortened;
3. the existing haploid breeding technology of rice depending on anther culture is not strong in universality, particularly the technical process of anther culture is almost not universal, and a large amount of early-stage research is needed from a culture medium formula to hormone proportion;
4. the haploid seeds are distinguished by identifying the seed coat color, batch and automation are easy to realize, and the traditional haploid identification technology needs phenotype and chromosome observation and has higher technical difficulty;
5. because the complete inactivation of centromere gene leads to the death of individual plants, the invention proposes that physical or chemical mutagenesis is utilized to induce gene to generate point mutation, and mutants with reduced gene function are created instead of gene mutants with complete loss of function, and the mutants with the type have reduced fertility, but can still breed seeds;
6. the haploid induction system of the corn is to utilize pollen to induce a target individual to generate haploid seeds, the stamens of a female parent are required to be manually removed, the work is manually finished, the required labor cost is higher, the haploid induction sterile line is provided, the sterile line is used as the female parent to directly receive the pollen of the target individual, the hybridization efficiency is higher, and the mechanized seed production is easy to realize;
7. the haploid induced sterile line can propagate sterile line seeds only through a matched maintainer line, so that a breeder can control the source of the induced line seeds more conveniently, and intellectual property is protected.
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In order to more clearly illustrate the embodiments of the present invention or the technical solutions in the prior art, the drawings needed to be used in the embodiments will be briefly described below, and it is obvious that the drawings in the following description are only some embodiments of the present invention, and it is obvious for those skilled in the art to obtain other drawings without inventive exercise.
FIG. 1 is a technical scheme of rice double haploid breeding based on haploid inducer line in example 1.
Detailed Description
The invention is further illustrated by the following examples and figures.
A method for breeding rice double haploid based on haploid induction line is shown in figure 1, and comprises the following steps:
1. maintainer line H353-B carrying multiple favorable genes was transformed into a red rice maintainer line.
Three-line maintainer line H353-B carrying 3 excellent genes (amylose content gene Wx, aroma gene fgr and rice blast resistance gene Pita) is bred in the early stage of the research group, a homonuclear heterogeneous sterile line H353-A is bred in a matched mode, and the homonuclear heterogeneous sterile line H353-A are hybridized to obtain F1 high sterility. A material H22 (male parent) carrying a dominant red seed coat gene RR is hybridized with H353-B (female parent), seed coats of F1 generation seeds are red, H353-B is used as a recurrent parent, red seed coat individuals are selected from each backcross generation to backcross with H353-B, the red seed coat individuals are selected for 1 time of selfing after 6 generations of backcross, red seed coat plants without separation are selected from selfed offspring, cytoplasm of the plants is the same as that of the H353-B, and the similarity between cell nucleus and the H353-B reaches 99%, and the plants are named as H353-BR. Except for the seed coat color, the characteristics of H353-BR are highly similar to those of H353-B.
2. H353-BR was mutagenized using EMS.
Selecting plump H353-BR dry seeds without plant diseases and insect pests 20000 grains, soaking the seeds in 0.5% EMS for 12H, and washing the seeds with sterile water for 12H. The treated seeds are planted in the field to form a first mutagenesis generation, the seeds are managed by conventional water and fertilizer, the seedling height is investigated in the 3-leaf stage, the plant height is investigated in the mature stage, and the seed setting rate is counted. The data show that after 0.5% EMS treatment, the height of the rice seedling is reduced by 21% compared with the control, the plant height is reduced by 10%, and the maturing rate is reduced by 15%, which shows that the EMS concentration is a relatively proper mutation dosage. When the plants are mature, 10 seeds are taken from each main spike and mixed and sowed to form 20000 mutagenic second-generation groups. The second generation of mutagenesis is divided into single plants and the number of each plant is 5cm or so of leaves for genotype analysis in a laboratory.
3. And (3) directionally screening the point mutation type of the centromere control gene CENH3 to obtain a haploid induction line H353-BRI carrying the red rice gene.
CENH3 is a centromere control gene reported in Arabidopsis, and complete inactivation of the gene leads to plant death, so the point mutation of GGA → GAG in the sequence screened by the invention may lead to reduced gene function rather than inactivation. BLAST was used to search for a homologous gene of CENH3 in rice, primers were designed to amplify the coding region, and sequencing was used to analyze whether a GGA → GAG point mutation was present. And (4) breeding individual plants with point mutations, and performing repeated sequencing verification. The real mutant is used as a female parent, the indica rice variety Huahang I is used as a male parent, the two are hybridized, the haploid proportion in the hybrid seeds is counted, and the mutant which can produce 5% of haploid seeds is selected for expanding propagation and is named as H353-BRI. The mutant is a haploid induction line H353-BRI carrying the red rice gene.
4. Hybridizing a haploid induction line H353-BRI carrying the red rice gene with a sterile line H353-A, and continuously backcrossing for 5 times by taking the H353-BRI as a recurrent parent to cultivate a haploid induction line sterile line H353-ARI carrying the red rice gene.
Since H353-BRI can still be selfed to obtain seeds, male flowers need to be manually removed when the H353-BRI is hybridized with a target individual, and a large amount of hybrid seeds are not obtained. In order to overcome the defects, H353-BRI is hybridized with a sterile line H353-A, H353-BRI is used as a recurrent parent for continuous backcross for 6 times, and each backcross generation selects red seed coat and sterile individuals for hybridization to finally obtain a haploid induction line sterile line H353-ARI with red rice genes (figure 1).
5. Haploid generation was induced using H353-ARI.
A breeding scheme is designed, and aims to quickly mature and aggregate the fragrance gene fgr and the rice blast resistance gene Pik into an individual. The specific scheme and implementation are as follows: indica rice material H25 carrying aroma gene fgr is used as a female parent, indica rice material H26 carrying rice blast resistance gene Pik is used as a male parent, the two are hybridized to obtain F1 seeds, the F1 seeds are planted in the field to form F1 plants, pollen of F1 is used for pollinating a sterile line H353-ARI in the flowering period, and the hybrid seeds are harvested. The seed coat of hybrid seeds is of two types, one red and one colorless. Since the seeds of the F1 plant to be induced are not red, individuals of colorless seed coats in the seeds obtained by crossing with the haploid inducing line are haploid only carrying male parent chromosomes, and the red seed coat hybrid seeds are diploid formed by combining the inducing line with F1 pollen. And screening out all colorless seeds, and carrying out the next doubling work.
6. Haploid seeds were doubled with colchicine.
Soaking the obtained haploid seeds with sterile water for 24 hours, then placing the seeds on wet sterile filter paper for germination for 24 hours, soaking the seeds with 0.6% colchicine for 8 hours when the embryo of the seeds germinates, and flushing the seeds with running water for 8 hours. The treated seeds are sowed in a greenhouse for normal fertilizer and water management. The seedlings are observed, because the haploid of the rice is dysplasia, dysplasia or malformed individuals can be eliminated according to the phenotype, and the normally-developed seedlings are the diploid obtained by successful doubling.
7. And (3) carrying out genotype identification on the diploid to obtain an individual containing fgr and Pik homozygous genes.
Taking diploid plant leaves, carrying out genotype identification, reserving individuals containing fgr and Pik homozygous genes simultaneously, achieving the purpose of gene polymerization, and finishing the breeding target formulated in the step 5.
The principles and embodiments of the present invention have been described herein using specific examples, which are provided only to help understand the method and the core concept of the present invention; meanwhile, for a person skilled in the art, according to the idea of the present invention, the specific embodiments and the application range may be changed. In view of the above, the present disclosure should not be construed as limiting the invention.
Claims (4)
1. A rice double haploid breeding method based on a haploid induction line is characterized by comprising the following steps:
s1, selecting an excellent rice three-line maintainer line material, hybridizing the excellent rice three-line maintainer line material with a rice material with dominant red rice genes, backcrossing a plurality of generations by using the maintainer line, and breeding a red rice maintainer line carrying the dominant red rice genes;
s2, soaking the screened red rice maintainer line material carrying the dominant red rice gene in EMS with the concentration of 0.3-1.0% for 12-24 h to obtain the red rice maintainer line material containing the functional mutant of the rice centromere gene;
s3, directionally screening functional mutants of the centromere gene of the rice, and selfing the screened mutants for 2 generations to form a stable genetic material;
s4, hybridizing the stable genetic material screened in the step S3 with a rice three-line sterile line, backcrossing for 5 generations by taking a red rice maintainer line mutant as a recurrent parent, and completing the replacement of the nuclear gene of the sterile line to obtain a red rice haploid induction maintainer line and a sterile line carrying a red rice gene and a centromere mutant gene;
s5, hybridizing a hybrid F1 prepared according to a breeding target with a red rice haploid induction sterile line, identifying the seed coat color of the obtained hybrid seeds, eliminating red seed coat hybrid seeds, and reserving colorless seed coat hybrid seeds;
s6, treating the colorless seed coat hybrid seeds by using colchicine to obtain a doubled diploid material, and identifying the agronomic characters or genotypes of the diploid material to form a new strain;
the directional screening method in step S3 is to detect the GGA → GAG sequence difference of CENH3 gene by sequencing, and the material with difference from the control is mutant.
2. The haploid inducer line-based rice double haploid breeding method as claimed in claim 1, wherein the red rice maintainer line mutant is hybridized with a rice three-line sterile line in S4, the red rice maintainer line mutant is backcrossed for 5 generations by taking the red rice maintainer line mutant as a recurrent parent, the replacement of the sterile line nuclear gene is completed, red seed coat seeds are selected from each backcross generation to carry out the next backcrossing, the red seed coat seeds are diploid, and the red seed coat genes are heterozygous.
3. The haploid induction line-based rice double haploid breeding method as claimed in claim 1, wherein the obtained hybrid seed is subjected to seed coat color identification in S5, wherein the seed with red seed coat is non-haploid seed, the seed with colorless seed coat is haploid seed, and the seed coat color screening can be achieved manually or mechanically.
4. The method for breeding rice doubled haploid based on haploid induction line as claimed in claim 1, wherein the colorless seed coat hybrid seeds in S6 are treated with colchicine, which means that seeds with embryo germination are soaked for 8h with colchicine with concentration of 0.6%, and haploid doubling can be realized after soaking is finished.
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