Production of yellow-seeded Brassica napus through interspecific crosses

Yellow‐seeded Brassica napus was developed from interspecific crosses between yellow‐seeded Brassica rapa var.‘yellow sarson’ (AA), black‐seeded Brassica alboglabra (CC), yellow‐seeded Brassica carinata (Bbcc) and black‐seeded B. napus (AACC). Three different interspecific crossing approaches were undertaken. Approaches 1 and 2 were designed directly to develop yellow‐seeded B. napus while approach 3 was designed to produce a yellow‐seeded CC genome species. Approaches 1 and 2 differed in the steps taken after trigenomic interspecific hybrids (ABC) were generated from B. carinata×B. rapa crosses. The aim of approach 1 was to transfer the yellow seed colour genes from the A to the C genome as an intermediate step in developing yellow‐seeded B. napus. For this purpose, the ABC hybrids were crossed with black‐seeded B. napus and the three‐way interspecific hybrids were self‐pollinated for a number of generations. The F₇ generation resulted in the yellowish‐brown‐seeded B. napus line, No. 06. Crossing this line with the B. napus line No. 01, resynthesized from a black‐seeded B. alboglabra x B. rapa var.‘yellow sarson’ cross (containing the yellow seed colour genes in its AA genome), yielded yellow‐seeded B. napus. This result indicated that the yellow seed colour genes were transferred from the A to the C genome in the yellowish‐brown seed colour line No. 06. In approach 2, trigenomic diploids (AABBCC) were generated from the above‐mentioned trigenomic haploids (ABC). The seed colour of the trigenomic diploid was brown, in contrast to the yellow seed colour of the parental species. Trigenomic diploids were crossed with the resynthesized B. napus line No. 01 to eliminate the B genome chromosomes, and to develop yellow‐seeded B. napus with the AA genome of ‘yellow sarson’ and the CC genome of B. carinata with yellow seed colour genes. This interspecific cross failed to generate any yellow‐seeded B. napus. Approach 3 was to develop yellow‐seeded CC genome species from B. alboglabra×B. carinata crosses. It was possible to obtain a yellowish‐brown seeded B. alboglabra, but crossing this B. alboglabra with B. rapa var.‘yellow sarson’ failed to produce yellow seed in the resynthesized B. napus. The results of approaches 2 and 3 demonstrated that yellow‐seeded B. napus cannot be developed by combining the yellow seed colour genes of the CC genome of yellow‐seeded B. carinata and the AA genome of ‘yellow sarson’.     

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.     

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.     

Inheritance of siliqua locule number and seed coat colour in Brassica juncea

The inheritance of siliqua locule number and seed coat colour in Brassica juncea was investigated, using three lines each of tetralocular brown seeded and bilocular yellow seeded. Three crosses of tetralocular brown seeded × bilocular yellow seeded lines were attempted and their F₁, F₂ and backcross generations were examined for segregation of these two traits. Brown seed colour and bilocular siliqua characters were found to be dominant over yellow seed and tetralocular siliqua, respectively. Chi‐square tests indicated that each trait is controlled by different sets of duplicate pairs of genes. Bilocular siliquae or brown seeds can result from the presence of either of two dominant alleles, whereas tetralocular siliquae or yellow seeds are produced when alleles at both loci are recessive. A joint segregation analysis of F₂ data indicated that the genes governing siliqua locule number and seed colour were inherited independently.     

Maternal and embryonal control of seed colour by different Brassica alboglabra chromosomes

Most oilseed rape, Brassica napus, cultivars are black‐seeded. The progenitor species, Brassica rapa, has either yellow or black seeds, while known cultivars of the other progenitor species Brassica oleracea/alboglabra have black seeds. To determine which chromosomes of B. alboglabra are carriers of seed colour genes, B. rapaalboglabra monosomic addition lines were produced from a B. napus resynthesized from yellow‐seeded B. rapa and brown/black‐seeded B. alboglabra. Eight out of nine possible lines have been developed and transmission frequencies of the alien chromosomes were estimated. Three B. alboglabra chromosomes in three of these lines influenced seed colour. B. rapa plants carrying alien chromosome 1 exhibited a maternal control of seed colour and produced only brown seeds, which gave rise to plants with either yellow or brown seeds. However, B. rapa plants carrying alien chromosome 4 or another as yet unidentified alien chromosome exhibited an embryonal control of seed colour and produced a mixture of yellow and brown seeds. The yellow seeds gave rise to yellow‐seeded plants, while the brown seeds gave rise to plants that yielded a mixture of yellow and brown seeds, depending on the absence or presence, respectively, of the B. alboglabra chromosome. Consequently, both maternal and embryonal control of seed colour are expected to contribute to the black‐seeded phenotype of oilseed rape.     

Inheritance of black testa colour in lentil (Lens culinaris Medik.)

The inheritance of testa (seed coat) colour and interaction of cotyledon and testa colours were studied in seven crosses of lentil (Lens culinaris Medik.) involving parents with black, brown, tan or green testa and with orange, yellow or dark green cotyledons. Analysis of F₂ and F₃ seed harvested from F₁ and F₂ plants, respectively, revealed that although black testa is dominant over nonblack testa, its penetrance is not complete since both F₁ plants and heterozygous F₂ plants produced varying proportions of seeds with either black or nonblack testa. The F₂ populations of the crosses between parents with brown and tan, as well as brown and green, testa segregated in the ratio of 3 brown : 1 tan and 3 brown : 1 green, respectively, indicating monogenic dominance of brown testa colour over tan or green. The expression of testa colour was influenced by cotyledon colour when parents with brown or green testa are crossed with those having orange or green cotyledons. Thus F₂ seeds from these crosses with a green testa always had green cotyledons and never orange cotyledons. F₂ seeds from these crosses with a brown testa always had orange cotyledons and never green cotyledons. These results suggest diffusion of a soluble pigment from the cotyledons to the testa. This revised version was published online in July 2006 with corrections to the Cover Date.     

Development of Strains with Yellow Seedcoat in Indian Mustard (Brassies juncea Czern. & Coss.)

A mutant with yellow seedcoat colour was isolated by Nayar (1968) in the mustard variety‘Rai-5′. This mutant was crossed to the national check cultivar ‘Varuna’ in order to develop improved strains with yellow seedcoat. Four such strains with yellow seeds were evaluated for their seed yield, yield components and percent oil. Two strains TM-9 and TM-17 were more productive than ‘Varuna’ in seed yield. All the yellow seeded strains showed higher oil percentage as compared to ‘Varuna’. The seedcoat in the yellow seeded strains accounts for 14-15% of the seed weight as compared to 18% in the black seeded ‘Varuna’. The higher proportion of the cotyledons and embryo accounts for the increased oil percentage in the yellow seeded types.     

Development of Yellow Seeded Brassica napus Through Interspecific Crosses

Yellow seeded Brassica napus was developed through interspecific crosses with the two mustard species, B. juncea and B. carinata. The objective of these two interspecific crosses was the introgression of genes for yellow seed colour from the A genome of B. juncea and C genome of B. carinata into the A and C genomes of B. napus, respectively. The interspecific F₁ generations were backcrossed to B. napus in an attempt to eliminate B genome chromosomes and to improve fertility. Backcross F₂ plants of the (B. napus×B. juncea) ×B. napus cross were then crossed with backcross F₂ plants of the (B. napus×B. carinata) ×B. napus cross. The objective of this intercrossing was to combine the A and C genome yellow seeded characteristics of the two backcross populations into one genotype. The F₂ generation of the backcross F₂ intercrosses was grown in the field, plants were individually harvested and visually rated for seed colour. Ninety-one yellow seeded plants were identified among the 4858 plants inspected. This result indicated that the interspecific crossing scheme was successful in developing yellow seeded B. napus.     

Dissection of a major QTL for seed colour and fibre content in Brassica napus reveals colocalization with candidate genes for phenylpropanoid biosynthesis and flavonoid deposition

A major quantitative trait locus (QTL) influencing seed fibre and colour in Brassica napus was dissected by marker saturation in a doubled haploid (DH) population from the black‐seeded oilseed rape line ‘Express 617’ crossed with a yellow‐seeded B. napus line, ‘1012–98’. The marker at the peak of a sub‐QTL with a strong effect on both seed colour and acid detergent lignin content lay only 4 kb away from a Brassica (H+)‐ATPase gene orthologous to the transparent testa gene AHA10. Near the peak of a second sub‐QTL, we mapped a copy of the key phenylpropanoid biosynthesis gene cinnamyl alcohol dehydrogenase, while another key phenylpropanoid biosynthesis gene, cinnamoyl co‐a reductase 1, was found nearby. In a cross between ‘Express 617’ and another dark‐seeded parent, ‘V8’, Bna.CCR1 was localized in silico near the peak of a corresponding seed fibre QTL, whereas in this case Bna.CAD2/CAD3 lay nearby. Re‐sequencing of the two phenylpropanoid genes via next‐generation amplicon sequencing revealed intragenic rearrangements and functionally relevant allelic variation in the three parents.     

Inheritance of siliqua orientation and seed coat colour in Brassica tournefortii

The inheritance of siliqua orientation and seed coat colour in Brassica tournefortii was investigated using four genotypes varying in these two characters. The F₁, F₂ and backcross generations of two crosses were used for studying the segregation pattern of the traits. The plants were classified for seed colour as having brown or yellow seeds and for siliqua orientation as having upright, semi‐spread or spread siliqua. Seed colour was found to be under monogenic control with brown being dominant over yellow. Siliqua orientation was under digenic polymeric gene action: upright siliqua was produced by the presence of two dominant genes and spread siliqua by two recessive genes. The absence of even a single dominant gene resulted in a third type of siliqua orientation, semi‐spread siliqua.     

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.     

Blue aleurone trait diagnoses developmental origin of three pistils in a floret in common wheat

The common ‘three‐pistil’ (TP) wheat mutation line expresses TPs in a floret normally containing TPs forming three grains set close back‐to‐back. The developmental origin of the TP trait in common wheat had been diagnosed non‐destructively using the blue aleurone trait. The aleurone colour of F₂ seeds grown in F₁ plants of cross TP/UC66049 was evaluated. Due to xenia, the hue of blue grain colour depended on dose of the Ba1 gene for blue aleurone in the triploid endosperm. The TP trait produced four types of segregation in three‐seed clusters: (i) white grain only, (ii) two white grains and one blue, (iii) one white grain and two blue, and (iv) three blue grains only. The observed frequency of blue–white seed within clusters followed the binominal distribution ₃C_r (0.75)^r·(0.25)^3–r, where r is the number of colour variants in three‐seed clusters (r = 0–3). Intrafloret segregation of seed colour and F₂ segregation derived from aleurone colour of F₃ seeds indicated an independent origin of the TP trait.     

Effects of Wheat Chromosome 1D Telosomes on Hybrid Seed Development in Wheat × Rye Crosses

Pollination of ‘Chinese Spring,’ monosome 1D plants with rye results in failure of hybrid seed development in a proportion of the F₁ seeds corresponding to the transmission rate of the nullisomic 1D egg cells. Development and viability of these hybrid seeds closely resemble that normally observed in T. aurum× rye crosses. Using ‘Chinese Spring’ chromosome ID telosomic plants in crosses with rye, it was possible to illustrate that the observed effect was associated with the long arm of this chromosome.     

The linkage between the stem rust resistance gene Sr2 and pseudo-black chaff in wheat can be broken

Recessively inherited gene Sr2 has provided the basis of durable resistance to stem rust (caused by Puccinia graminis tritici) in wheat (Triticum aestivum L.) worldwide. The associated earhead and stem melanism or ‘pseudo‐black chaff’ is generally used as a marker for this gene. Sr2 has been postulated in many wheat cultivars of India including ‘Lok 1’, based on associated pseudo‐black chaff in adult plants, and leaf chlorosis in seedlings. However, dominant inheritance of the resistance factor operating in ‘Lok 1’, and a 13 : 3 (resistant : susceptible) F₂ segregation in the ‘Sr2‐line’ (‘Chinese Spring’⁶ × ‘Hope’ 3B) × ‘Lok 1’ cross confirmed that Sr2 was absent in ‘Lok 1’. Susceptible plants with a pseudo‐black chaff phenotype were observed in F₂ populations of ‘Agra Local’ (susceptible) × ‘Lok 1’, and the ‘Sr2‐line’ × ‘Lok 1’ crosses. Most of the F₃ families derived from the susceptible F₂ segregants with pseudo‐black chaff phenotypes were true breeding for the expression of pseudo‐black chaff with susceptibility to stem rust. Thus, linkage of pseudo‐black chaff with Sr2 in wheat can be broken, and hence, caution may be exercised in using pseudo‐black chaff as a marker for selecting Sr2 in breeding programmes.     

Genetic control of crossability of triticale with rye

Limited genetic knowledge is available regarding crossability between hexaploid triticale (2n= 6x= 42, 21″, AABBRR, amphiploid Triticum turgidum L.‐Secale cereale L.) and rye (2n= 14, 7″, RR). Our objectives were to determine (1) the crossability between triticales and rye and (2) the inheritance of crossability between F₂ progeny from intertriticale crosses and rye. First, ‘8F/Corgo’, a hexaploid triticale, was crossed as a female with two landrace ryes, ‘Gimonde’ and, ‘Vila Pouca’ and two derived north European cultivars, ‘Pluto’ and ‘Breno’. These crosses produced 21.7, 20.9, 5.9, and 5.6%, seed‐set or crossability, respectively, showing that the landrace ryes produced higher seed‐set than the cultivars. Second, ‘Gimonde’ rye was crossed as a male with four triticales for 3 years. The control cross, ‘Chinese Spring’ wheat × rye, produced 80‐90% seed‐set. Of the four triticales, ‘Beagle’ produced 35.7‐56.8% seed‐set. The other three triticales produced less than 20% seed‐set, showing that the triticales differ in crossability with ‘Gimonde’ rye. Third, six FiS from intertriticale crosses (‘8F/Corgo’בBeagle’, ‘Beagle’בCachirulo’, ‘Lasko’בBeagle’, ‘8F/Corgo’בCachirulo’, ‘Lasko’בCachirulo’, ‘Lasko’ב8F/Corgo’) were crossed to ‘Gimonde’ rye. Results indicated that lower crossability trait was partially dominant in the two F₁S from crosses involving ‘Beagle’(high crossability) with‘8F/Corgo’ and ‘Cachirulo’(low crossability) and completely dominant in the ‘Beagle’בLasko’ cross, as it happens in wheat. Fourth, segregants in four F₂ populations (‘Lasko’בBeagle’, ‘8F/Corgo’בBeagle’, ‘Lasko’ב8F/Corgo’, and‘8F/Corgo’בCachirulo’) were crossed with rye. Segregation for crossability was observed, although distinct segregation classes were blurred by environmental and perhaps other factors, such as self‐incompatibility alleles in rye. Segregation patterns showed that ‘Beagle’, with high crossability to rye, carries either Kr1 or Kr2. The three triticales with low crossability with rye were most likely homozygous for Kr1 and Kr2. Therefore, it is likely that the Kr loci from A and B genomes acting in wheat also play a role in triticale × rye crosses.     

A Comparison of Genetic Segregation in Traditional and Microspore-Derived Populations of Brassica juncea L. Czern and Coss.

An isolated microspore culture procedure was used to produce doubled haploid lines of Brassica juncea from F¹ plants of reciprocal crosses between the cultivar‘RLM514’and a canola quality breeding line. The inheritance of two qualitative markers, seed color and leaf hairiness, was compared using traditional and microspore-derived populations from these crosses. Chi-square tests indicated that each trait is controlled by different sets of duplicate pairs of genes. Brown seeds or hairy leaves can result from the presence of either of two dominant alleles, whereas yellow seed or glabrous leaves are produced when alleles at both loci are recessive. The segregation of genes controlling seed color and leaf hairs in doubled haploid progeny did not differ significantly from that expected under random assortment, indicating that doubled haploids can be used in this species for genetic studies, and probably cultivar development as well.     

Generation and characterisation of colchicine-induced polyploid Lavandula × intermedia

Double podding in cultivated chickpeas (Cicer arietinum L.) can increase yield and yield stability. In the present study, we performed reciprocal crosses of ‘kabuli’ (double podded) and ‘desi’ (single podded) chickpeas to determine (i) the expressivity and penetrance of double podding, (ii) the correlations of yield and yield components, and (iii) the heritability of double podding, flower color, and stem pigmentation in F₂ plants. Reciprocal crosses were performed with two genotypes, AC 2969 (kabuli) and ICC 4969 (desi), to generate F₁ and F₂ plants. The results indicated hybrid vigor (heterosis) for yield in F₁ plants and better performance of F₂ plants. Yield and yield components of some lines in F₂ were superior to the best parent, indicative of transgressive segregation. In particular, the presence of double podding (‘s’ allele) significantly increased yield in some of the transgressive segregants. Expressivity and penetrance of the ‘s’ allele depends on the background of the female parent. Some of the double podding progeny had greater seed yields than those of the single podding progeny and greater seed yields than the best parents. Double podding, stem pigmentation, and pink flowers each appears to be governed by a single recessive gene. Stem pigmentation and pink flowers appear to be linked traits that depend on the genetic background of the crossed chickpeas. Taken together, our studies of reciprocal crosses of kabuli and desi chickpeas clearly showed that yield could be improved by selection for transgressive phenotypes that have double podding.     

Genetic markers for manganese efficiency in durum wheat

Manganese (Mn) deficiency is a major constraint of alkaline soils around the world, particularly for cultivation of durum wheat, which is more intolerant of low Mn levels than either common wheat or barley. Genetic variation for Mn efficiency exists in the current germplasm of durum wheat. Several restriction fragment length polymorphisms (RFLPs) previously shown to be linked to the Mel1 locus for Mn efficiency on chromosome 4HS of barley were tested on 88 selected F₂ plants of the durum cross, ‘Stojocri 2’ (Mn efficient) בHazar’ (Mn inefficient). The Mel1‐linked RFLP marker Xcdo583a was closely linked to the trait and explained over 42% of the total variation for Mn efficiency in the ‘Stojocri 2’/‘Hazar’ F₂ progeny. This marker has the potential to provide a valuable tool for the marker‐assisted selection of Mn‐efficient durum progeny derived from crosses with ‘Stojocri 2’.     

Comparative seed yield performance of high-by-high and low-by-high crosses in flax *

Successful identification of a desirable segregant depends partly on the parents chosen to make the crosses. This experiment was conducted to compare performance and genetic variability for seed yield, yield components, agronomic traits and harvest index of lines derived from low- and high-yielding flax (Linum usitatissimum L.) crosses, and to identify important yield components for flax seed yield improvement. The lines chosen as high yielding parents for this experiment were‘Linott’and 'Summit; and the low yielding parents were‘Grant’and Ci2395. The high-yielding lines produced significantly more seeds per boll and had a higher harvest index than the low-yielding lines. Evaluation of 161 F_2:6 lines from four crosses among these lines showed that the greatest genetic variability, highest cross average, and highest F₆ line seed yield occurred in the low x high cross,‘Grant’x‘Linott’. All low X high crosses exhibited higher genetic variances for seed yield than the high x high crosses. The high x high cross 'Summit’x‘Linott’had low genetic variance for seed yield. Number of bolls per area was determined by linear regression, path coefficient analysis, and stepwise multiple regression analysis to be the most important component of seed yield. This study showed that hybridization of low and high-yielding flax lines may be useful to increase genetic variability and obtain high-yielding flax lines.     

Monosomic analysis of Helminthosporium leaf blight resistance genes in wheat

A set of 21 monosomic (2n ‐ 1) and the disomic (2n) lines of the ‘Chinese Spring’ cultivar were crossed with ‘Chirya‐3′, the CIMMYT synthetic wheat line which has been identified as highly resistant for Helminthosporium leaf blight disease (HLB), in order to locate the genes governing disease resistance. The F₁ and segregating populations were challenged and screened against the most virulent pure mono‐conidial HLB isolate KL‐8 (Karnal, India). The F₁ progenies of the crosses were found to be susceptible because of the recessive nature of resistance. The F₂ progeny of the control cross (‘Chinese Spring’בChirya‐3’), segregated in the ratio of 1: 15 (resistant: susceptible), indicating that resistance to HLB was controlled by a pair of recessive genes. While the F₂ progeny of 19 monosomic crosses segregated in the ratio of 1: 15 (resistant: susceptible), the progeny of the remaining two crosses, 7B and 7D, deviated significantly from the ratio, revealing that 7B and 7D were the critical chromosomes for resistance genes that were located one on each chromosome. Moreover, the critical lines, 7B and 7D, confirmed the digenic complementary recessive nature of gene action by fitting well with the overall pooled F₂ segregation ratio of 13: 51 (resistant: susceptible) as expected for digenic complementary recessive resistance. The F₃ segregation ratios of the critical crosses, based on their pooled F₂ analysis, was estimated as 19: 32: 13 (non‐segregating susceptible: segregating as susceptible and resistant: non‐segregating resistant). F₃ progenies when tested with these ratios showed goodness‐of‐fit, confirming that the two pairs of recessive resistance genes were located on chromosomes 7B and 7D.