Independent Inheritance of Erucic Acid Content and Flower Colour in the C-Genome of Brassica napus L.

The synthetic Brassica napus L. line No7076 was obtained from a cross between yellow-flowered and zero-erucic turnip rape (B. campestris) Sv85-38301 and white-flowered and high-erucic (41.4%) B. oleracea ssp. alboglabra No6510. This synthetic B. napus is pale-flowered and has an average erucic acid content of 25.8 %. It was crossed with the yellow-flowered and zero-erucic B. napus line SvS4-2S053 and segregation of the erucic acid content and flower colour was studied in F₁ and F₂ generations. The high erucic acid content was controlled by a single gene in the C-genome and was additively inherited. Strong evidence was obtained in support of independent segregation of the erucic-arid content and the flower colour characters controlled by the C-genome of B. napus.     

Inheritance of petal colour and its independent segregation from seed colour in Brassica rapa

The inheritance of petal (flower) colour and seed colour in Brassica rapa was investigated using two creamy‐white flowered, yellow‐seeded yellow sarson (an ecotype from Indian subcontinent) lines, two yellow‐flowered, partially yellow‐seeded Canadian cultivars and one yellow‐flowered, brown‐seeded rapid cycling accession, and their F₁, F₂, F₃ and backcross populations. A joint segregation of these two characters was examined in the F₂ population. Petal colour was found to be under monogenic control, where the yellow petal colour gene is dominant over the creamy‐white petal colour gene. The seed colour was found to be under digenic control and the yellow seed colour (due to a transparent coat) genes of yellow sarson are recessive to the brown/partially yellow seed colour genes of the Canadian B. rapa cvs.‘Candle’ and ‘Tobin’. The genes governing the petal colour and seed colour are inherited independently. A distorted segregation for petal colour was found in the backcross populations of yellow sarson × F₁ crosses, but not in the reciprocal backcrosses, i.e. F₁× yellow sarson. The possible reason is discussed in the light of genetic diversity of the parental genotypes.     

The increase of seed fertility of Brassicoraphanus through cytological irregularity

Summary Amphidiploids (Brassicoraphanus) were produced by means of colchicine treatment of F₁ hybrids between Brassica japonica Sieb. and Raphanus sativus L. The cytology of the amphidiploids was studied from F₁ to F₃ generations. Some plants had the euploid chromosome number 2n=38, whereas others had the aneuploid number 2n=37. One or two of either quadrivalents or trivalents, as well as some univalents, were seen in most of the plants examined. All the plants showed a low seed fertility. In F₃ generation there arose some yellow-flowered plants, all of which showed a higher seed fertility than normal white-flowered plants. It is postulated that the change of flower colour might originate in the segmental exchange of only partially homologous chromosomes following multivalent formation. A gene causing white flower colour was perhaps closely linked to a gene causing sterility, and both genes were probably excluded together through the segmental exchange of the chromosomes. Therefore, it can be said that the increase of fertility was induced by cytological irregularity.     

Genetics of fertility restoration in A2-based cytoplasmic genetic male sterility system in pigeonpea (Cajanus cajan)

The three short duration cytoplasmic genetic male sterility (CGMS) hybrids developed using A₂ (Cajanus scarabeoides) cytoplasm-based male sterile lines (CORG 990047A, CORG 990052A and CORG 7A) and the restorer inbred AK 261322 and their segregating populations (F₂ and BC₁F₁) were subjected to the study of inheritance of fertility restoration in pigeonpea. The fertility restoration was studied based on three different criteria, namely, anther colour, pollen grain fertility and pollen grain morphology and staining. The F₂ and BC₁F₁ populations of the three crosses, namely, CORG 990047A × AK 261322, CORG 990052A × AK 261322 and CORG 7A × AK 261322, segregated in the ratio of 3:1 and 1:1, for anther colour (yellow:pale yellow), pollen grain fertility (fertile:sterile) and for pollen grain morphology and staining. The above study confirmed that the trait fertility restoration was controlled by single dominant gene. This finding can be utilized for the identification of potential restorers, which can be further used in the development of CGMS-based hybrids in pigeonpea.     

水稻新质源(CMS-FA)雄性不育恢复基因的遗传研究

发掘水稻新型雄性不育细胞质源CMS-FA,育成系列优质米不育系和系列新质源恢复系,组配成强优势杂交稻组合的基础上研究新质源雄性不育恢复系的恢复基因遗传。采用新质源(CMS-FA)不育系金农1A与恢复系金恢3号杂交获得杂交F₁代种子,种植F₁代,收获自交F₂代种子。用F₁分别与不育系或保持系回交,获得(不育系//不育系/恢复系和不育系/恢复系//保持系)2个测交群体。同时种植P₁、P₂、F₁、F₂、B₁F₁和B₂F₁等群体,考察花粉染色率、套袋结实率和自然结实率,卡平方测验遗传分离适合度。结果表明,不育系与恢复系杂交F₁代正常可育,育性恢复(可育)基因为显性遗传。F₂代分离出可育︰不育适合3︰1,育性恢复(可育)基因为1对显性基因控制。B₁F₁和B₂F₁代2个测交群体的可育︰不育都适合1︰1分离规律,验证了F₂代育性恢复(可育)单基因的遗传模式。暂时确定新质源(CMS-FA)核质互作三系的基因型为不育系S(SS)、保持系F(SS)和恢复系S(FF)。     

Chromosomal location of fertility restoring genes for ‘wild abortive’ cytoplasmic male sterility using primary trisomics in rice

Summary Identification and location of fertility restoring genes facilitates their deployment in a hybrid breeding program involving cytoplasmic male sterility (CMS) system. The study aimed to locate fertility restorer genes of CMSWA system on specific chromosomes of rice using primary trisomics of IR36 (restorer), CMS (IR58025A) and maintainer (IR58025B) lines. Primary trisomic series (Triplo 1 to 12) was crossed as maternal parent with the maintainer line IR58025B. The selected trisomic and disomic F₁ plants were testcrossed as male parents with the CMS line IR58025A. Plants in testcross families derived from disomic F₁ plants (Group I crosses) were all diploid; however, in the testcross families derived from trisomic F₁ plants (Group II crosses), some trisomic plants were observed. Diploid plants in all testcross families were analyzed for pollen fertility using 1% IKI stain. All testeross families from Group I crosses segregated in the ratio of 2 fertile: 1 partially fertile+partially sterile: 1 sterile plants indicating that fertility restoration was controlled by two independent dominant genes: one of the genes was stronger than the other. Testcross families from Group II crosses segregated in 2 fertile: 1 partially fertile+ partially sterile: 1 sterile plants in crosses involving Triplo 1, 4, 5, 6, 8, 9, 11 and 12, but families involving triplo 7 and triplo 10 showed significantly higher X² values, indicating that the two fertility restorer genes were located on chromosome 7 and 10. Stronger restorer gene (Rf-WA-1) was located on chromosome 7 and weaker restorer gene (Rf-WA-2) was located on chromosome 10. These findings should facilitate tagging of these genes with molecular markers with the ultimate aim to practice marker-aided selection for fertility restoration ability.     

Inheritance of root traits and phosphorus uptake in common bean (Phaseolus vulgaris L.) under limited soil phosphorus supply

Heterosis is an important way to improve yield and quality for many crops. Hybrid rice and hybrid maize contributed to enhanced productivity which is essential to supply enough food for the increasing world population. The success of hybrid rice in China has led to a continuous interest in hybrid wheat, even when most research on hybrid wheat has been discontinued in other countries for various reasons including low heterosis and high seed production costs. The Timopheevii cytoplasmic male sterile system is ideal for producing hybrid wheat seeds when fertility restoration lines with strong fertility restoration ability are available. To develop PCR-based molecular markers for use in marker-assisted selection of fertility restorer lines, two F₂ populations derived from crosses R18/ND36 and R9034/ND36 were used to map fertility restoration genes in the two elite fertility restorer lines (R-lines) R18 and R9034. Over 678 SSR markers were analyzed, and markers closely linked to fertility restoration genes were identified. Using SSR markers, a major fertility restoration gene, Rf3, was located on the 1B chromosome in both populations. This gene was partially dominant in conferring fertility restoration in the two restorer lines. SSR markers Xbarc207, Xgwm131, and Xbarc61 are close to this gene. These markers may be useful in marker-assisted selection of new restorer lines with T. timopheevii cytoplasm. Two minor QTL conferring fertility restoration were also identified on chromosomes 5A (in R18) and 7D (in R9034) in two R-lines.     

Identification and Inheritance of Fertility Restorer Genes for 'tour' CMS in Rapeseed (Brassica napus L.)

Eighteen genotypes of Brassica napus were crossed to a cytoplasmic male sterile (CMS) line of B. napus BO 15 carrying B. tournefortii cytoplasm (‘tour’ cytoplasm). Fourteen genotypes were found to be stable maintainers of the ‘tour’ CMS. Of the remaining four genotypes, GSL-1 and ‘Asahi-natane’ were found to be heterozygous and ‘Mangun’ and ‘Yudal’ were homozygous for the restorer gene. Analysis of the F₁ and F₂ progenies of (CMS) BO 15 בMangun’ and (CMS) BO 15 בYudal’ showed that fertility restoration is controlled by a single dominant gene. The availability of a number of stable maintainer lines and the simple inheritance pattern of fertility restorer gene makes ‘tour’ CMS a useful system for hybrid seed production in rapeseed.     

Inheritance and mapping of a restorer gene for the rapeseed cytoplasmic male sterile line 681A

A novel cytoplasmic male sterility‐fertility restoration system has been developed in rapeseed (Brassica napus). The cytoplasmic male sterile line 681A was derived from a spontaneous male sterile mutant in a newly released double‐low rapeseed cultivar ‘Xiangyou 13′. The restorer line 714R was identified in the interspecific progeny from a B. napus×B. juncea‐cross. Genetic analysis showed that fertility restoration for 681A cytoplasmic male sterility was controlled by a single dominant nuclear gene which might originate from B. juncea. The RAPD marker S1039_‐520 was found to be linked to the restorer gene in F₂ progeny of 681A × 714R with a recombination frequency of 5.45%.     

Comparison of two fertility restoration systems against photoperiod-sensitive cytoplasmic male sterility in wheat

A ‘two‐line system’ using photoperiod‐sensitive cytoplasmic male sterility (PCMS) caused by Aegilops crassa cytoplasm under a long‐day photoperiod ( 15 h) has been proposed as a new means of producing hybrid varieties in common wheat. The PCMS line is maintained by self‐pollination under short‐day conditions, and hybrid seeds can be produced through outcrossing of the PCMS line with a pollinator under long‐day conditions. Two kinds of fertility restoration systems against the PCMS are known. One is involved with a set of multiple fertility‐restoring (Rf) genes in the wheat cultivar ‘Norin 61’ located on (at least) chromosomes 4A, 1D, 3D and 5D. The other is controlled by a single dominant major Rf gene, Rfd1, located on the long arm of chromosome 7B in the wheat cultivar ‘Chinese Spring’. To examine the degree of fertility restoration by these two systems, nine PCMS lines were crossed with ‘Norin 61’ and ‘Chinese Spring’ as the restorer lines, and the F₁ hybrids were investigated. The degree of fertility restoration was estimated by comparing the seed set rates in the F₁ hybrids having the Ae. crassa cytoplasm and those with normal cytoplasm. The results revealed that the fertility restoration ability of a set of multiple Rf genes in ‘Norin 61’ was higher than that of the Rfd1 gene in ‘Chinese Spring’.     

Investigation of possible associations between partial fertility restoration and agronomic traits in Triticum aestivum L.

Summary Many conventional hard red spring wheat (Triticum aestivum L. em Thell) lines, including several North Dakota cultivars, carry a gene (or genes) which restore partial male fertility to male sterile plants with Triticum timopheevi Zhuk. cytoplasm. Since this gene has no fertility restoration function in T. aestivum cytoplasm, the postulation can be made that it is being retained in conventional lines because of pleiotropic effects, favorable linkages or chance. The research reported in this paper examined these possibilities. Forty F₆ lines, derived from a single F₂ plant which was heterozygous for a gene (or genes) for partial fertility restoration, were evaluated for two years in a yield trial planted at Fargo, North Dakota. The 40 lines were testcrossed to a male sterile line having T. timopheevi cytoplasm, and the mean seed set of testcrosses was used as a measure of a line's fertility restoration potential. Twenty-seven lines had the gene for partial fertility, and 13 lines apparently lacked this gene. The 40 lines differed for heading date, anther extrusion, plant height, grain yield, 200-kernel weight, test weight, and grain protein percentage. However, comparisons of lines having the restorer gene with those lacking the gene did not provide any obvious explanation for the retention of the partial fertility restorer gene in the breeding stocks of the North Dakota conventional hard red spring wheat breeding program. The possibility that the restorer gene was linked with genes for resistance to stem rust or leaf rust also was evaluated by testing lines for their reaction to several races of rust. No conclusive association was found.Contribution from the Agric. Exp. Sta., North Dakota State University, Fargo, ND 58105, Journal Article no.     

The mechanism of increased seed fertility accompanied with the change of flower colour in Brassicoraphanus

Summary In Brassicoraphanus (amphidiploids between Brassica japonica Sieb. and Raphanus sativus L.), yellow-flowered plants that occurred among originally white-flowered plants showed an increased seed fertility. It is assumed that the gene Y (yellow-flower gene) from Brassica and the gene W (white-flower gene) from Raphanus are located at corresponding loci of only partially homologous chromosomes. W is dominant (epistatic) over Y. The normal white-flowered plants have the genotype YYWW. A YYYW-plant was found, which is assumed to have arisen through crossing-over following multivalent formation. In the progeny of this plant, yellow-flowered plants (YYYY) as well as white-flowered plants (YYWW, YYYW) appeared. The gene for flower colour is closely linked to a gene which controls the development of embryos (or endosperm). This gene promotes the development of embryos in homozygous condition. Therefore, the embryo having only the yellow-flower gene can develop more easily into viable seed than the embryo having the white-flower gene. It is also possible that the sterility of white-flowered plants is caused by a discordance between the cytoplasm of Brassica and W (or genes linked to W) of Raphanus.     

Development of dominant nuclear male-sterile lines with a blue seed marker in durum and common wheat

In order to develop genie male‐sterile lines with a blue seed marker, male‐sterile plants, controlled by a dominant nuclear gene Ms2, were used as female parents against a 4E disomic addition line ‘Xiaoyan Lanli’(2n= 44, AABBDD+4EII) as the male parent to produce monosomic addition lines with blue seed. Male‐sterile plants from the monosomic addition lines were pollinated with durum wheat for several generations and in 1989 a male‐sterile line with the blue grain gene and the male‐sterile gene Ms2 on the same additional chromosome was detected and named line 89‐2343. Using this line, the blue seed marker was successfully added to a short male‐sterile line containing Ms2 and Rht10. The segregation ratios of male sterility and seed colour as well as the chromosome figurations of different plants indicated that the blue grain genes, Ms2 and Rht10 were located on the same additional chromosome. Cytological analysis showed that the blue marker male‐sterile lines in durum wheat and common wheat were monosomic with an additional chromosome 4E. The inheritance ratio for blue seed male‐sterile plants and white seed male‐fertile plants was 19.7% and 80.3%, respectively, in common wheat. The potential for using blue marker sterile lines in population improvement and hybrid production is discussed.     

Selection and genetic improvement of pollen fertility restorer lines with Triticum timopheevii cytoplasm in common wheat

The paper summarizes the selection and improvement of pollen fertility restoration in cytoplasmic male-sterile lines during the past 30 years at Jiangsu Province, China. A fertility restorer line (R16) with a good history of strong and stable restoring ability to different sterile lines was bred by accumulating fertility-restoring genes from derivatives of T797 and other restorer lines such as Primepi. A series of well-performing restorer lines with similar fertility-restoring ability has been bred by improving agronomic characters, disease-resistance and kernel size of R16. The restoring ability of these restorer lines using different male-sterile lines demonstrates that fertility restoration is no longer an obstacle for commercial utilization of hybrid wheat with the Triticum timopheeviii cytoplasmic male-sterile system. Line 2114 is a restorer with a single restoring gene transferred from Aegilops umbellulata. Its restoring ability, using both difficult and easily restored lines was 82% and 93.3% respectively. Maiyou No. 5, one hybrid variety, showed 13.2% yield advantage over the control variety in the Jiangsu Province registration test in 1997-1998 and was superior to nine other varieties adapted to the Jiangsu Province.     

Development and use of a two-gene marker-aided selection system for fertility restorer genes in rice

We have established marker-aided selection strategies for the two major Rf genes (Rf3 and Rf4) governing fertility restoration of␣cytoplasmic-genetic male sterility (CMS) in rice. Polymorphisms between restorer and non-restorer␣lines were observed using RG140/PvuII for Rf3 located on chromosome 1 and S10019/BstUI for Rf4 located on chromosome 10. DNA polymorphisms associated with these two loci in restorer lines of wild abortive (WA), Dissi, and Gambiaca cytoplasm are conserved, suggesting that similar biological processes control pollen fertility in this diverse cytoplasm. Because of their close linkage to Rf genes and distinct banding patterns, STS markers RG140/PvuII and S10019/BstUI are well suited for marker-aided selection, enhanced backcross procedures, and pyramiding of Rf genes in agronomically superior non-restorer lines. The combined use of markers associated with these two loci improved the efficiency of screening for putative restorer lines from a set of elite lines. Positional analyses of Rf4 and the inheritance pattern of the polymorphism in S10019/BstUI suggest that Rf4, governing fertility restoration in WA-CMS in rice, is likely to be the same gene governing fertility restoration in BT- and HL-CMS that has a gametophytic effect, which explains why 100% pollen fertility in hybrids is impossible to attain.     

A unique introgression from Moricandia arvensis confers male fertility upon two different cytoplasmic male-sterile lines of Brassica juncea

A Brassica juncea line carrying an introgression from Moricandia arvensis restored male fertility to two cytoplasmic male‐sterile (CMS) B. juncea lines carrying either M. arvensis or Diplotaxis catholica cytoplasm. Genetics of fertility restoration was studied in the F₁, F₂, F₃ and backcross generations of the cross between CMS and fertility‐restorer lines. No male‐sterile plants were found in F1‐F₃ generations of the cross between CMS [M. arvensis] B. juncea and the restorer. However, a 1: 1 segregation for male sterility and fertility was observed when the F1 was pollinated with non‐restorer pollen from a euplasmic line. These results clearly show that restoration is mono‐genic and gametophytic. In CMS lines carrying D. catholica cytoplasm, the restorer conferred male fertility to the F1 and showed 3: 1 and 1: 1 segregations for male fertility and sterility in F₂ and BC1 generations, respectively, indicating a monogenic, sporophytic mode of fertility restoration. The results were also supported by pollen stainability in the F1 which was about 65% in M. arvensis‐based CMS and >90% in D. catholica‐based CMS. The above results are discussed in the light of previous molecular studies which showed association between CMS and atpA in both systems.     

Development of yellow-seeded Brassica napus of double low quality

Two yellow‐seeded white‐petalled Brassica napus F₇ inbred lines, developed from interspecific crosses, containing 26–28% emcic acid and more than 40 μmol glucosinolates (GLS)/g seed were crossed with two black/dark brown seeded B. napus varieties of double low quality and 287 doubled haploid (DH) lines were produced. The segregation in the DH lines indicated that three to four gene loci are involved in the determination of seed colour, and yellow seeds are formed when all alleles in all loci are in the homozygous recessive state. A dominant gene governed white petal colour and is linked with an erucic acid allele that, in the homozygous condition, produces 26–28% erucic acid. Four gene loci are involved in the control of total GLS content where low GLS was due to the presence of recessive alleles in the homozygous condition in all loci. From the DH breeding population a yellow‐seeded, yellow‐petalled, zero erucic acid line was obtained. This line was further crossed with conventional B. napus varieties of double low quality and, following pedigree selection, a yellow seeded B. napus of double low quality was obtained. The yellow seeds had higher oil plus protein content and lower fibre content than black seeds. A reduction of the concentration of chromogenic substances was found in the transparent seed coat of the yellow‐seeded B. napus.     

An improved cytoplasmic male sterile (Diplotaxis berthautii) Brassica juncea: identification of restorer and molecular characterization

We report the development of an improved cytoplasmic male sterile (CMS) system of Brassica juncea carrying cytoplasm of the wild species Diplotaxis berthautii. Flowers of the CMS line are smaller than the euplasmic line but have improved nectaries. Anthers are slender and fail to extend to the level of stigma. Female fertility of the CMS line is comparable to the euplasmic line. Fertility restorers of Moricandia arvensis and D. catholica-based alloplasmic CMS systems of B. juncea were found capable of restoring male fertility to this new CMS line. The fertility restoration is monogenic and gametophytic. Southern analysis showed that the cytoplasm of the CMS line is different from euplasmic B. juncea and other CMS systems restored by the same restorer lines. Northern analysis of the CMS, fertility restored and euplasmic lines using eight mitochondrial gene probes revealed altered atpA expression associated with male sterility. Cleaved amplified polymorphic sequence (CAPS) markers were identified for the plastid gene psbB, which could be useful for a quick identification of this CMS line. S.R. Bhat and P. Kumar contributed equally to this work.     

Inheritance of seed colour in Brassica campestris L. and breeding for yellow-seeded B. napus L.

Summary Seed colour inheritance was studied in five yellow-seeded and one black-seeded B. campestris accessions. Diallel crosses between the yellow-seeded types indicated that the four var. yellow sarson accessions of Indian origin had the same genotype for seed colour but were different from the Swedish yellow-seeded breeding line. Black seed colour was dominant over yellow. The segregation patterns for seed colour in F₂ (Including reciprocals) and BC₁ (backcross of F₁ to the yellow-seeded parent) indicated that the black seed colour was conditioned by a single dominant gene. Seed colour was mainly controlled by the maternal genotype but influenced by the interplay between the maternal and endosperm and/or embryonic genotypes. For developing yellow-seeded B. napus genotypes, resynthesized B. napus lines containing genes for yellow seed (Chen et al., 1988) were crossed with B. napus of yellow/brown seeds, or with yellow-seeded B. carinata. Yellow-seeded F₂ plants were found in the crosses that involved the B. napus breeding line. However, this yellow-seeded character did not breed true up to F₄. Crosses between a yellow-seeded F₃ plant and a monogenomically controlled black-seeded B. napus line of resynthesized origin revealed that the black-seeded trait in the B. alboglabra genome was possibly governed by two independently dominant genes with duplicated effect. Crossability between the resynthesized B. napus lines as female and B. carinata as male was fairly high. The sterility of the F₁ plants prevented further breeding progress for developing yellow-seeded B. napus by this strategy.     

Genetics of colour traits in common vetch (Vicia sativa L.)

Five parents of common vetch (Vicia sativa L.) having orange/beige cotyledon colour, brown/white testa colour, purple/green seedling colour and purple/white flower colour were crossed as a full diallele set. The inheritance patterns of cotyledon, testa or seed coat colour, flower and seedling colour, were studied by analyzing their F₁, F₂, BC₁ and BC₂ generations. The segregation pattern in F₂, BC₁ and BC₂, showed that cotyledon colour was governed by a single gene with incomplete dominance and it is proposed that cotyledon colour is controlled by two allelic genes, which have been designated Ct₁ and Ct₂. Testa colour was governed by a single gene with the brown allele dominant and the recessive allele white. This gene has been given the symbol H. Two complementary genes governed both flower and seedling colours. These flower and seedling colour genes are pleiotropic and the two genes have been given the symbols S and F.