CN102246690B - Method for breeding all sterile line with genetic stability of rape recessive epistasis genic male sterile line by means of molecular marker - Google Patents
Method for breeding all sterile line with genetic stability of rape recessive epistasis genic male sterile line by means of molecular marker Download PDFInfo
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Abstract
The invention, belonging to the field of molecule breeding of rape, relates to a method for breeding an all sterile line with genetic stability of a rape recessive epistasis genic male sterile line by means of molecular marker. According to the invention, by the close link of molecular marker and fertility genes, the genotype of molecular mark is tracked and selected to select a program of the genotypes of a two-type line and a temporary maintainer line, so that the breeding strategy of the sterile line controlled by polygenes is improved from a process of complete phenotype identification to blanket search of genotype identification to a high efficiency breeding process of well-targeted marker genotype selection first to actual phenotype identification finally, by backcrossing and selfing multi-generation and field phenotype identification, the homozygous two type line and temporary maintainer line with same genetic background and uniform growth in fields are obtained, and then all sterile line seeds with genetic stability are produced for largely plantation. Therefore, the current situation that seeds for largely plantation of the sterile line are actual three-way cross hybrids is changed, and the problems of bad uniformity and declined advantages caused by the current situation are avoided.
Description
Technical field
The invention belongs to the rape molecular breeding field, be specifically related to a kind of method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line.
Background technology
Male sterile is swede type rape heterosis utilization important channel.Male sterility system involved in the present invention is a kind of line with genic sterile system (Chen Fengxiang etc. that done mutually to control by recessive sterile gene and recessive epistasis suppressor gene, Acta Agronomica Sinica, 1998,24 (4): 431-438), this sterile system has the sterile degree height, sterility is stable; Petal Full-open, hybrid seed yield height and ordinary cole kind are the characteristics of its natural restorer.The propositions such as Chen Fengxiang obtain complete sterile line again by " amphitypy system " and " temporary maintainer line " hybridization of isozygotying and " restorer " (arbitrary common variety) hybridization generation F1 hybrid " three are " utilizes the pattern (patent No.: 97125803.1), need to manually pull out the drawback of 50% fertile plant when having overcome traditional recessive cytoblast sterile " bilinear method " production of hybrid seeds.This sterile system is breeding man and many breeding units concerns extremely both at home and abroad after proposing, and successfully cultivate so far a plurality of cross combinations and are applied to production practice.Yet finding in Breeding Application has two difficult points to limit application and the promotion rate of this sterile system.The one, filial generation that the controlled by multiple genes system brings heredity separate complex, the improvement of amphitypy system and the temporary maintainer line inefficiency that wastes time and energy; The 2nd, big area is used for that the sterile line of the production of hybrid seeds is actual to be the F1 hybrid, and real cross-fertilize seed is triple hybrid, and this can cause advantage decay and purity decline to a certain extent, may become this sterile system and be able on a large scale bottleneck problem with large-area applications.To for many years field fertility investigation of the hybrid separation offspring of this system, the Molecular Identification of fertile gene and repeatedly practising of marking supplementary breeding, think that the ordinary cole kind all contains the dominant allelotrope of educating in recessive sterile gene site according to us; Recessive epistasis suppressor gene site exists recessive and dominant two kinds of allele distributions; Second pair of sterile overlapping genes of recessiveness is ms in most kinds
4Ms
4Genotype or do not have the 2nd sterile gene site.
Summary of the invention
In order to solve the technological deficiency of above-mentioned existence, the present invention seeks to propose a kind of method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line, " two types that the isozygoty system " and the temporary maintainer line that utilize pattern to select to have identical improved seeds (being) genetic background according to Recessive epistatic GMS " three are ", thereby obtain the complete sterile line of inheritance stability, realize a large amount of F of production
1The purpose of (seed of single cross) cross rape reduces and avoids triple hybrid to be applied to produce the problem that the appearance that may occur is inconsistent and advantage descends.
In order to realize above-mentioned technical purpose, the present invention has adopted following technical scheme:
By the method for the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line, the method comprises the steps:
1) at first with treat the kind of transformation or strain respectively with known stable amphitypy system in the sterile strain and stablize the temporary maintainer line plant and hybridize respectively F of isozygotying
1Obtain F for selfing
2Segregating population;
2) according to treating transformation kind or strain and stable temporary maintainer line hybridization F
2The Fertility segregation situation is determined next step transformation approach from generation to generation, treats transformation kind or strain and temporary maintainer line hybridization F if there is Fertility segregation then to utilize
2In generation, is as next round transformation colony; If without Fertility segregation, then with isozygoty in the amphitypy system sterile strain with treat transformation kind or incross F
2Colony is as next round transformation colony;
3.1) treat transformation kind or strain and temporary maintainer line hybridization F
2In generation, utilized MS during as transformation colony
3/ ms
3Select temporary maintainer line genotype individual plant with Rf/rf gene compact linkage molecule mark, and as male parent with treat that the kind of transformation or strain backcross, and obtain BC after the selfing generation
1F
2In generation, then repeat above-mentioned same method and carry out Marker Identification, backcross and selfing, until BC
3F
2Or highlyer backcross, self-generation;
3.2) treat the sterile strain hybridization F that isozygotys in transformation kind or strain and the amphitypy system
2Generation is during as transformation colony, the sterile strain of the rear molecular markers for identification of blooming, choose isozygoty sterile pnca gene type with treat that the kind of transformation or strain backcross, and obtain BC after the selfing generation
1F
2From generation to generation, then repeat same method and carry out Marker Identification, backcross and selfing, until BC
3F
2Or highlyer backcross, self-generation;
4) to transformation BC
3F
2Or the higher self-generation that backcrosses, use first MS
3/ ms
3All individual plants of compact linkage molecule Marker Identification filter out MS
3Ms
3And ms
3Ms
3Continue behind the marker genetype further to detect with Rf/rf linkage molecule mark, selecting at last the transformation amphitypy is MS
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf marker genetype individual plant;
5) the homozygous two-type line MS to selecting
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf genotype plant carries out the field phenotypic evaluation, to treat that transformation kind or product are contrast, carry out the comparison of coherence test of growing way appearance, the results seed carries out Quality Detection, finally choose appearance and treat that transformation kind or strain are consistent, quality reaches the homozygous two-type line MS of two substandards
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3The rfrf Genotype Strains with the alternate plantation homozygous two-type line of certain proportion and temporary maintainer line, is pulled out fertile plant in the amphitypy system after first the spending of buddingging in isolation environment, gather in the crops seed in the sterile strain when ripe, and the complete sterile line that namely obtains inheritance stability provides the land for growing field crops production of hybrid seeds.
6) temporary maintainer line Genotype selfing breeding, the sterile line that isozygotys is handed over procreation by brother and sister between homozygous two-type line fertile plant and sterile strain.
As preferably, to the marker assisted selection of segregating generation, extract the genomic dna of rape single-strain blade, use and MS
3/ ms
3M1 and the M2 codominant marker primer chain with Rf/rf carry out pcr amplification, and the PCR product is carried out polyacrylamide gel electrophoresis, filter out homozygous two-type line MS
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf genotype individual plant.
As preferably, above-mentioned and MS
3/ ms
3The upstream and downstream sequence of linkage molecule mark M1 primer is respectively: 5 '-GTCTGTTTCTCTTCCCGTTGTC-3 ' and 5 '-GTGCTGCTCCGGTGTTATC-3 '; With the upstream and downstream sequence of Rf/rf linkage molecule mark M2 primer be respectively: 5 '-AGGAAGCCCAACAGGACTTT-3 ' and 5 '-AATTCGATTCTCCATCGTGC-3 '.Certainly that need among the present invention and MS
3/ ms
3The linkage molecule mark also can adopt document " Theor Appl Genet (2007) 115:113-118, Theor Appl Genet (2008) 117:11-18 "; With Rf/rf linkage molecule labeled primer also can be with reference to " Plant Breeding (2008), 127,145-149 " and Euphytica (2008), 164 (2): the upper disclosed primer of 377-384.
As preferably, step 3.1) described temporary maintainer line genotype individual plant and the known sterile strain ms that isozygotys
3Ms
3The RfRf test cross, the summer numerous added-generation is identified the temporary maintainer line genotype.
As preferably, step 3.2) described sterile strain and the known stable temporary maintainer line ms of isozygotying
3Ms
3The rfrf test cross, the sterile pnca gene type that isozygotys that summer numerous added-generation checking is selected.
The present invention the invention has the advantages that owing to adopted above-mentioned technical scheme:
1, the present invention proposes to utilize the assisting sifting strategy with Recessive Male sterility and the closely linked codominant marker of recessive epistasis suppressor gene, can be like a cork from segregating generation target pick out exactly temporary maintainer line genotype (1/13) and the sterile line genotype (1/3) of isozygotying continues to backcross; Again through once hybridization, then continuous backcross adds selfing 3-4 wheel until segregating population genetic background and treat that the transformation kind is basic identical.At last from BC
3-4F
2In the segregating generation, utilize molecule marker to filter out simultaneously homozygous two-type line and temporary maintainer line genotype.This invention will significantly reduce the seed selection workload, greatly improve seed selection accuracy rate and success ratio.The proposition of the method is incited somebody to action so that application potential and the using value of swede type rape Recessive epistatic GMS system are doubled and redoubled.
2, the Breeding strategy that proposes according to the present invention, be expected in identical good genetic background colony, obtain simultaneously homozygous two-type line and temporary maintainer line genotype with the 4-5 year, further under isolation environment, produce the complete sterile line seed application of inheritance stability in the big area production of hybrid seeds.The hybrid of results is common seeds of single cross, changed the present big area of this system produce with kind actual be the present situation of triple hybrid, avoid the risk of the poor advantage decline of reguarity that may bring.
3, the present invention is directed to rape variety in esse two kinds of genotype patterns on two fertility-related genes, proposed for MS
3MS
3The appropriate to the occasion selection temporary maintainer line of the genotypic kind transformation of RfRf genotype ms
3Ms
3Rfrf is donor parents; And for MS
3MS
3Then to select the sterile line ms that isozygotys during the genotypic transformation parent of rfrf
3Ms
3RfRf is donor, can guarantee to occur among the transformation offspring various possible genotype; In the transformation program " Xia Fan " non-segregating generation is proposed in addition, normal growth season plantation segregating generation, the pattern that hockets.This is to consider that strange land " Xia Fan " should simplify procedures, avoid selecting error (in the situation of impossible marker detection), and under the positive season growing environment, utilize the mark assisting sifting, from the segregating population that expands, filter out the individual backcross transformation that continues of minority target gene type, embody the ingenious combination on the space-time from each orientation, reached best and the fastest selection effect.
Description of drawings
Fig. 1 is techniqueflow chart of the present invention.
Embodiment
The method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line as shown in Figure 1, the technical scheme that the method is taked may further comprise the steps:
1, kind recessive epistasis suppressor gene site allelotrope affirmation to be turned: errorless for guaranteeing, kind at first to be turned simultaneously with stable amphitypy system in isozygoty sterile strain and temporary maintainer line (for male parent) hybridization, the F1 selfing has or not Fertility segregation to judge that kind to be turned is MS from F2 from generation to generation
3MS
3RfRf (there is sterile strain to separate with temporary maintainer line hybridization F2 generation, sterile/as can to educate: 3/13) or MS
3MS
3Rfrf (hybridizing F2 for separating without sterile strain with temporary maintainer line) genotype.
If 2 treat that the transformation kind is MS
3MS
3The RfRf genotype, continue according to the following steps:
1) treats that the transformation kind is maternal MS
3MS
3RfRf and known temporary maintainer line ms
3Ms
3The rfrf plant carries out conventional sexual hybridization, F
1The selfing of Dai Xiafan bagging produces F
2Generation.Normal growth season plantation F
2Segregating population is pressed individual plant seedling stage and is extracted DNA, uses first MS
3/ ms
3All individual plants of compact linkage molecule marker detection are selected ms
3Ms
3Continue behind the marker genetype to screen with Rf/rf compact linkage molecule mark, obtain temporary maintainer line marker genetype individual plant.
2) the temporary maintainer line marker genetype ms that selects
3Ms
3The rfrf individual plant, continue with treat transformation kind MS on one side
3MS
3RfRf is that female parent backcrosses, and produces BC1F1 from generation to generation, on one side with the known sterile strain ms that isozygotys
3Ms
3The RfRf test cross, the summer numerous added-generation is identified the temporary maintainer line genotype.Simultaneously BC1F1 bagging selfing produces BC1F2 from generation to generation.Next repeats first step from generation to generation, and normal growth is planted segregating population season, selects the sterile strain of isozygotying during mark is auxiliary and continues to backcross; The summer breeding is planted non-segregating population, only need certainly accompany each other generation, until BC3F2 from generation to generation.
3) enlarge the BC3F2 kind and plant colony's (more than 1000 strains), extract individual plant DNA seedling stage, use first MS
3/ ms
3All individual plants of compact linkage molecule Marker Identification filter out MS
3Ms
3And ms
3Ms
3Continue behind the marker genetype to detect with Rf/rf linkage molecule labeled analysis, selecting at last the transformation amphitypy is MS
3Ms
3RfRff, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf marker genetype individual plant.Can repeat simultaneously first step and continue to select temporary maintainer line ms
3Ms
3The rfrf marker genetype backcrosses, and produces more transformation amphitypy system and the temporary maintainer line of advanced lines.
4) the auxiliary transformation amphitypy of mark is sterile ms
3Ms
3RfRf and fertile plant MS
3Ms
3Brother and sister hand between RfRf; Transformation temporary maintainer line ms
3Ms
3Rfrf and the known sterile strain ms that isozygotys
3Ms
3The RfRf test cross; The transformation sterile strain ms that isozygotys
3Ms
3RfRf and known temporary maintainer line ms
3Ms
3The rfrf test cross; The transformation amphitypy is fertile plant MS
3Ms
3RfRf and known isozygoty sterile strain test cross and bagging selfing.The next generation is investigated all test crosses and self progeny's fertility performance, if transformation temporary maintainer line and the known sterile strain test cross that isozygotys, offspring's complete sterility, then temporary maintainer line transformation success.If isozygoty sterile strain and the known temporary maintainer line test cross of transformation, offspring's complete sterility, and the transformation amphitypy is fertile plant MS
3Ms
3RfRf and the known sterile strain test cross that isozygotys, offspring's 1: 1 (can educate: sterile) separates, and self progeny's 3: 1 (can educate: sterile) separates, and then transformation amphitypy system succeeds.
5) amphitypy system and the temporary maintainer line strain of all transformation successes are carried out field phenotypic evaluation and Quality Detection, select yield level, plant forms, resistance requirement and quality standard and amphitypy system and the temporary maintainer line for the treatment of that the transformation kind is consistent.Yet according to (Acta Agronomica Sinicas such as Chen Fengxiang, 1998,24 (4): the method that 431-438) proposes, the amphitypy of alternate plantation seed selection system and temporary maintainer line under isolation environment, initial bloom stage is pulled out the sterile strain in the amphitypy system, spontaneous pollination between temporary maintainer line and the sterile strain is gathered in the crops seed in the sterile strain when ripe, namely obtains inheritance stability and basically identical complete sterile line seed is used for the big area production of hybrid seeds with treating the transformation variety and genetype.
If 3 treat that the transformation cultivar identification is MS
3MS
3The rfrf genotype then follows these steps to continue:
1) treats that the transformation kind is male parent MS
3MS
3Rfrf and the known sterile line ms that isozygotys
3Ms
3The RfRf plant carries out conventional sexual hybridization, and the bagging selfing produced F2 generation when F1 was numerous for the summer.Normal growth season plantation F2 segregating population, budding or just spend after extract sterile strain leaf DNA by individual plant, screen with Rf/rf compact linkage molecule mark, select marker site and be the sterile strain ms of RfRf of isozygotying
3Ms
3RfRf.
2) the sterile marker genetype ms that isozygotys that filters out
3Ms
3The RfRrf individual plant, continue with treat transformation kind MS on one side
3MS
3Rfrf backcrosses, and produces BC1F1 from generation to generation, on one side with known stable temporary maintainer line test cross ms
3Ms
3Rfrf, the sterile pnca gene type that isozygotys that summer numerous added-generation checking is selected.Simultaneously BC1F1 bagging selfing produces BC1F2 from generation to generation.Next repeats first step from generation to generation, and normal growth is used the sterile strain of Rf/rf compact linkage molecule Marker Identification season, and filtering out marker genetype is the plant ms that isozygotys sterile
3Ms
3RfRrf continues with treating transformation kind MS
3MS
3Rfrf backcrosses, and test cross, summer breeding are planted non-segregating generation, and the generation of certainly accompanying each other is until the BC3F2 generation.
3) obtain BC3F2 for seed after, on one side be MS by transformation genotype in above-mentioned the 2nd joint
3MS
3The the 3rd to the 5th step the during kind of RfRf is carried out.Can continue with treating transformation kind MS on one side
3MS
3Rfrf backcrosses, enter higher one take turns backcross, test cross and selfing.
Sequence table
＜110〉Zhejiang Academy of Agricultural Science
＜120〉by the method for the upper line with genic sterile of molecular selection rapeseed plant recessive system inheritance stability complete sterile line
<160>4
<210>1
<211>22
<212>DNA
＜213〉primer
<222>(1)…(22)
<400>1
GTCTGTTTCTCTTCCCGTTGTC 22
<210>2
<211>19
<212>DNA
＜213〉primer
<222>(1)…(19)
<400>2
GTGCTGCTCCGGTGTTATC 19
<210>3
<211>20
<212>DNA
＜213〉primer
<222>(1)…(20)
<400>3
AGGAAGCCCAACAGGACTTT 20
<210>4
<211>20
<212>DNA
＜213〉primer
<222>(1)…(20)
<400>4
AATTCGATTCTCCATCGTGC 20
Claims (5)
1. by the method for the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line, it is characterized in that the method comprises the steps:
1) at first with treat the kind of transformation or strain respectively with known stable amphitypy system in the sterile strain and stablize the temporary maintainer line plant and hybridize respectively F of isozygotying
1Obtain F for selfing
2Segregating population;
2) according to treating transformation kind or strain and stable temporary maintainer line hybridization F
2The Fertility segregation situation is determined next step transformation approach from generation to generation, treats transformation kind or strain and temporary maintainer line hybridization F if there is Fertility segregation then to utilize
2In generation, is as next round transformation colony; If without Fertility segregation, then with isozygoty in the amphitypy system sterile strain with treat transformation kind or incross F
2Colony is as next round transformation colony;
3.1) treat transformation kind or strain and temporary maintainer line hybridization F
2In generation, utilized and MS during as transformation colony
3/ ms
3Gene and select temporary maintainer line genotype individual plant, described and MS with the closely linked molecule marker of Rf/rf gene
3/ ms
3The upstream and downstream sequence of the closely linked molecule marker M1 of gene primer is respectively: 5 '-GTCTGTTTCTCTTCCCGTTGTC-3 ' and 5 '-GTGCTGCTCCGGTGTTATC-3 '; With the upstream and downstream sequence of the closely linked molecule marker M2 of Rf/rf gene primer be respectively: 5 '-AGGAAGCCCAACAGGACTTT-3 ' and 5 '-AATTCGATTCTCCATCGTGC-3 '; And as male parent with treat that the kind of transformation or strain backcross, and obtain BC after the selfing generation
1F
2In generation, then repeat above-mentioned same method and carry out Marker Identification, backcross and selfing, until BC
3F
2Or highlyer backcross, self-generation;
3.2) treat the sterile strain hybridization F that isozygotys in transformation kind or strain and the amphitypy system
2In generation,, the sterile strain of the rear molecular markers for identification of blooming was screened with the closely linked molecule marker M2 of Rf/rf gene primer during as transformation colony, chose to isozygoty sterile pnca gene type and treat that the kind of transformation or strain backcross, and obtain BC after the selfing generation
1F
2From generation to generation, then repeat same method and carry out Marker Identification, backcross and selfing, until BC
3F
2Or highlyer backcross, self-generation;
4) to transformation BC
3F
2Or the higher self-generation that backcrosses, use first MS
3/ ms
3All individual plants of compact linkage molecule Marker Identification filter out MS
3Ms
3And ms
3Ms
3Continue behind the marker genetype further to detect with Rf/rf linkage molecule mark, selecting at last the transformation amphitypy is MS
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf marker genetype individual plant;
5) the homozygous two-type line MS to selecting
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf genotype plant carries out the field phenotypic evaluation, to treat that transformation kind or product are contrast, carry out the comparison of coherence test of growing way appearance, the results seed carries out Quality Detection, finally choose appearance and treat that transformation kind or strain are consistent, quality reaches the homozygous two-type line MS of two substandards
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3The rfrf Genotype Strains with the alternate plantation homozygous two-type line of certain proportion and temporary maintainer line, is pulled out fertile plant in the amphitypy system after first the spending of buddingging in isolation environment, gather in the crops seed in the sterile strain when ripe, and the complete sterile line that namely obtains inheritance stability provides the land for growing field crops production of hybrid seeds;
6) temporary maintainer line Genotype selfing breeding, the sterile line that isozygotys is handed over procreation by brother and sister between homozygous two-type line fertile plant and sterile strain.
2. the method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line according to claim 1 is characterized in that: to the marker assisted selection of segregating generation, extract the genomic dna of rape individual plant, use and MS
3/ ms
3With Rf/rf respectively chain M1 and M2 codominant marker primer carry out pcr amplification, the PCR product is carried out polyacrylamide gel electrophoresis, filter out homozygous two-type line MS
3Ms
3RfRf, ms
3Ms
3RfRf and temporary maintainer line ms
3Ms
3Rfrf genotype individual plant.
3. the method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line according to claim 1 is characterized in that: step 3.1) described temporary maintainer line genotype individual plant and the known sterile strain ms that isozygotys
3Ms
3The RfRf test cross, the summer numerous added-generation is identified the temporary maintainer line genotype.
4. the method by the upper line with genic sterile system of molecular selection rapeseed plant recessive inheritance stability complete sterile line according to claim 1 is characterized in that: step 3.2) described sterile strain and the known stable temporary maintainer line ms of isozygotying
3Ms
3The rfrf test cross, the sterile pnca gene type that isozygotys that summer numerous added-generation checking is selected.
5. be used for the molecule marker of selectively breeding hybrid rape Recessive epistatic GMS system inheritance stability complete sterile line, it is characterized in that: with the upstream and downstream sequence of MS3/ms3 linkage molecule mark M1 primer be respectively: 5 '-GTCTGTTTCTCTTCCCGTTGTC-3 ' and 5 '-GTGCTGCTCCGGTGTTATC-3 '; With the upstream and downstream sequence of Rf/rf linkage molecule mark M2 primer be respectively: 5 '-AGGAAGCCCAACAGGACTTT-3 ' and 5 '-AATTCGATTCTCCATCGTGC-3 '.
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Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN1559187A (en) * | 2004-03-09 | 2005-01-05 | 华中农业大学 | Method for selectively breeding cabbage type rape dominant karyon male sterility isozygotic wo-purpose line |
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2010
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Patent Citations (1)
| Publication number | Priority date | Publication date | Assignee | Title |
|---|---|---|---|---|
| CN1559187A (en) * | 2004-03-09 | 2005-01-05 | 华中农业大学 | Method for selectively breeding cabbage type rape dominant karyon male sterility isozygotic wo-purpose line |
Non-Patent Citations (4)
| Title |
|---|
| 王贵春.甘蓝型油菜隐性细胞核雄性不育两型系9012AB雄性不育基因的分子标记开发.《中国博士学位论文全文数据库》.2009,(第2期), |
| 甘蓝型油菜细胞核雄性不育性的遗传研究――Ⅰ.隐性核不育系9012A的遗传;陈凤祥等;《作物学报》;19980430(第04期);全文 * |
| 甘蓝型油菜隐性细胞核雄性不育两型系9012AB雄性不育基因的分子标记开发;王贵春;《中国博士学位论文全文数据库》;20090215(第2期);全文 * |
| 陈凤祥等.甘蓝型油菜细胞核雄性不育性的遗传研究――Ⅰ.隐性核不育系9012A的遗传.《作物学报》.1998,(第04期), |
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