Increased anthocyanin content in purple pericarp × blue aleurone wheat crosses

Due to their antioxidant activity, anthocyanins are of increasing interest for nutritionists, food scientists and plant breeders. Anthocyanins in wheat grains are expressed in either the pericarp or aleurone layer. Previous studies revealed that different anthocyanins are present in wheat varieties carrying genes for either the purple pericarp or the blue aleurone trait. Progeny from crosses between red‐, purple‐ and blue‐grained wheat varieties were selected over several cycles for grain colour by visual scoring. Bulked F₅ grains were evaluated for their total anthocyanin content by UV‐VIS spectrophotometry and HPLC‐MS. The results demonstrate that it is possible to increase the anthocyanin content by the combination of the different genetic backgrounds for purple pericarp and blue aleurone, even though the majority of progeny were within the range of the purple‐ and blue‐grained check varieties. Visual scoring for grain colour is efficient, reliable and fast for selection in early breeding generations. Advanced breeding lines with high anthocyanin content can be identified by simple extraction methods and spectrophotometric measurements.     

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.     

Wheats with purple and blue grains: a review

Summary In addition to white and red grains, wheats with purple and blue grains may occur. Purple grain colour is caused by anthocyanins in the pericarp whereas blue colour is caused by anthocyanins in the aleurone layer. Purple grains occur in tetraploid wheats from Ethiopia, and in one bread wheat accession apparently native to China.Although the use of the purple and blue grain characters as markers has been suggested, their expression is often erratic, especially when heterozygous.No hexaploid wheat with blue grains was described prior to the artificial introgression of genes from diploid wheat and Agropyron species. The number of different sources of blue aleurone gene(s) from Agropyron elongatum is unknown. It is possible that with exchange between researchers the same or related accessions have been used at several research stations. Accessions of diploid wheats are known to possess blue aleurone. The breeding history of a number of purple and blue grained accessions is described. Research should indicate the source species of the gene for blue aleurone of the blue-grained Barevna.     

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.     

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.     

Quantitative trait loci mapping of seed colour,hairy leaf,seedling anthocyanin,leaf chlorosis and days to flowering in F2 population of Brassica rapa L.

A diversity arrays technology (DArT) map was constructed to identify quantitative trait loci (QTL) affecting seed colour, hairy leaf, seedling anthocyanin, leaf chlorosis and days to flowering in Brassica rapa using a F₂ population from a cross between two parents with contrasting traits. Two genes with dominant epistatic interaction were responsible for seed colour. One major dominant gene controls the hairy leaf trait. Seedling anthocyanin was controlled by a major single dominant gene. The parents did not exhibit leaf chlorosis; however, 32% F₂ plants showed leaf chlorosis in the population. A distorted segregation was observed for days to flowering in the F₂ population. A linkage map was constructed with 376 DArT markers distributed over 12 linkage groups covering 579.7 cM. The DArT markers were assigned on different chromosomes of B. rapa using B. rapa genome sequences and DArT consensus map of B. napus. Two QTL (RSC1‐2 and RSC12‐56) located on chromosome A8 and chromosome A9 were identified for seed colour, which explained 19.4% and 18.2% of the phenotypic variation, respectively. The seed colour marker located in the ortholog to Arabidopsis thaliana Transparent Testa2 (AtTT2). Two QTL RLH6‐0 and RLH9‐16 were identified for hairy leaf, which explained 31.6% and 20.7% phenotypic variation, respectively. A single QTL (RSAn‐12‐157) on chromosome A7, which explained 12.8% of phenotypic variation was detected for seedling anthocyanin. The seedling anthocyanin marker is found within the A. thaliana Transparent Testa12 (AtTT12) ortholog. A QTL (RLC6‐04) for leaf chlorosis was identified, which explained 55.3% of phenotypic variation. QTL for hairy leaf and leaf chlorosis were located 0–4 cM apart on the same chromosome A1. A single QTL (RDF‐10‐0) for days to flowering was identified, which explained 21.4% phenotypic variation.     

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.     

Identification and inheritance of a partially dominant gene for yellow seed colour in Brassica napus

A yellow‐seeded doubled haploid (DH) line no. 2127‐17, derived from a resynthesized Brassica napus L., was crossed with two black‐seeded Brassica cultivars ‘Quantum’ and ‘Sprint’ of spring type. The inheritance of seed colour was investigated in the F₂, and BC1 populations of the two crosses and also in the DH population derived from the F1 of the cross ‘Quantum’× no. 2127‐17. Seed colour analysis was performed with the colorimeter CR‐300 (Minolta, Japan) together with a visual classification system. The immediate F1 seeds of the reciprocals in the two crosses had the same colour as the self‐pollinated seeds of the respective black‐ and yellow‐seeded female parents, indicating the maternal control of seed colour. The F1 plants produced yellow‐brown seeds that were darker in colour than the seeds of no. 2127‐17, indicating the partial dominance of yellow seed over black. In the segregating BC1 progenies of the two crosses, the frequencies of the black‐ and yellow‐seeded plants fit well with a 1 : 1 ratio. In the cross with ‘Quantum’, the frequencies of yellow‐seeded and black‐seeded plants fit with a 13 : 3 ratio in the F₂ progeny, and with a 3 : 1 ratio in the DH progeny. However, a 49 : 15 segregation ratio was observed for the yellow‐seeded and black‐seeded plants in the F₂ progeny of the cross with ‘Sprint’. It was postulated from these results that seed colour was controlled by three pairs of genes. A dominant yellow‐seeded gene (Y) was identified in no. 2127‐17 that had epistatic effects on the two independent dominant black‐seeded genes (B and C), thereby inhibiting the biosynthesis of seed coat pigments.     

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.     

A New Mutation in Wheat Producing Three Pistils in a Floret

A floret carries only one pistil that will develop into one grain after fertilization in normal cultivated common wheat (Triticum aestivum L.) plants. A new cultivated common wheat mutation line ‘Three Pistils’ is described. It carries three pistils in a floret, all with the potential to develop into grains. The floret morphology of this line is reported. Genetic analysis of the three pistils trait was carried out by crossing Three Pistils with the normal common wheat variety Chinese Spring. F₂ population segregation analysis revealed that the three pistils trait is controlled by a single dominant gene. This conclusion was confirmed by the backcross test.     

Genetics of novel corolla colours in periwinkle

Inheritance of a novel corolla colour in periwinkle [Catharanthus roseus (L) G. Don], viz. magenta, was studied by crossing an accession MJ, possessing this corolla colour, with cultivar Nirmal, possessing white corolla. The accession MJ was also crossed with another accession OR, possessing another novel corolla colour, viz. orange-red, to determine the relationship between genes governing magenta corolla and orange-red corolla. The F₁ plants of the cross MJ× Nirmal had pink corolla and red eye. In the F₂ generation, five kinds of corolla colours were observed: (i) pink corolla and red eye, (ii) rose corolla and red eye, (iii) magenta corolla and red eye, (iv) white corolla and red eye and (v) white corolla. The observed frequencies of the five kinds of plants fitted a ratio of 144:27:9:12:64. The progeny of the backcross, F₁ × MJ, segregated into three kinds of plants, (i) pink corolla and red eye, (ii) rose corolla and red eye and (iii) magenta corolla and red eye, in the ratio of 2:1:1, while the backcross, F₁ × Nirmal, segregated into two kinds of plants, (i) pink corolla and red eye and (ii) white corolla, in the ratio of 1:1. Two new genes (proposed symbols O^m and J) appeared to be involved in the determination of magenta and rose corolla colours. Interaction between four independent genes R, W, O^m and J, appeared to explain the observed segregation in the cross MJ × Nirmal. The F₁ plants of the cross MJ × OR had scarlet-red corolla and red eye. The segregation data of F₂ and backcross generations suggested that genes governing orange-red corolla and magenta corolla were allelic to each other. Two new and non-parental corolla colours viz., rose corolla and scarlet-red corolla, were observed in the progeny of the crosses of the present study.     

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.     

Genetic characterization of the red coloured corolla phenotype for Gossypium arboreum accession PI 529731

Flower corolla colour is an important trait for the attraction of pollinators and for the horticultural industry. Gossypium arboreum (L.) accessions from the United States Department of Agriculture germplasm collection frequently show flowers with a yellow coloured corolla. Accession PI 529731 is unique in that the flowers have a red coloured corolla. Genetic characterization of corolla pigmentation was conducted by crossing PI 529731 with two white flower accessions. Flowers with a red corolla were observed in the F₁ generations suggesting a dominant trait. Variation in corolla colour was observed for plants in the F₂ populations including dark red, red, light red, white, yellow and white with petals having red coloured margins. These data support a single dominant gene conferring the four red corolla phenotypes. The yellow corolla phenotype also supported a single dominant gene model. Dominant alleles at both loci are required for expression of the PI 529731 phenotype and data support a two gene model with a 9:3:3:1 segregation ratio. These data are useful for the characterization of genetic mechanisms controlling tissue‐specific pigmentation.     

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.     

Effect of selection for pollen grain size on various traits in common bean

Summary Selection among microgametophytes usually exploits variation in pollen grain germination. Studies of variation in pollen grain size in common bean (Phaseolus vulgaris L.) suggested that selection for size might lead to changes in sporophytic traits. To determine whether microgametophytic selection based on size would affect pollen grain size in subsequent generations or sporophytic traits that were correlated with pollen grain size, pollen grains from three crosses were separated into two size categories by sieving and then used to pollinate cv. Diacol Calima. Selection resulted in changes in pollen grain diameter for pollen from F₁, F₂ and F₃ plants for all crosses. In vitro germination indicated no differences between vigor of large and small grains, but extraction and sieving reduced germinability. F₁ seed from two of the crosses with size-selected pollen varied in weight according to pollen grain size, but in subsequent generations, the effect disappeared. Both size categories of selected pollen resulted in F₂ progeny with reduced numbers of seeds per pod as compared to controls, suggesting that the size selection process may have resulted in indirect selection for traits reducing seed set. The overall results suggested that genes determining pollen grain size in bean have little or no effect on sporophytic traits such as seed size and seed yield.     

Inheritance of Eye Pattern and Seed Coat Colour in Cowpea (Vigna unguiculata [L.] Walp.)

The F₂ progenies of crosses between several cowpea (V. unguiculata) lines were investigated for variation of eye pattern and seed coat colour. It was found that three (W, H, O) and five (R, P, B, M, N) major genes control eye pattern and seed coat colour, respectively. The recessive gene (GO) for restricted eye pattern enables the underlying basic white or cream seed coat colour to be observed. A similar effect is obtained with the recessive gene (rr) for colour expression. The expression of mottling (V), possibly a seed coat pattern, may for be observed when it is combined with the genes for certain eye patterns. The significance of these findings in breeding for consumer preference for specific seed coat colour is discussed.     

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.     

Mapping of Re,a gene conferring the red leaf trait in ornamental kale (Brassica oleracea L. var. acephala)

Variegated leaf colour is an important agronomic trait that affects the market value of ornamental kale (Brassica oleracea L. var. acephala). The red leaf phenotype in kale is due to anthocyanin accumulation. To investigate the pattern of inheritance of this trait, we constructed an F₂ population by crossing ‘Y005‐15’, a double haploid with red leaves, with a white‐leaved double haploid, ‘Y011‐13‐38’, followed by self‐pollination. An F₂ population consisting of 4284 individuals was used to study the inheritance of this trait, which showed that the character was controlled by a dominate gene. All of the 1050 white leaf trait plants in the F₂ were used for mapping and developing markers linked to Re gene. Results showed that Re was mapped to a locus on linkage group C09 of Brassica oleracea, and the locus was mapped between six SSR markers (C9Z1, C9Z16‐1, C9Z90, C9Z94, C9Z96 and C9Z99), with a genetic distance of 6.7, 1.0, 0.3, 2.0, 2.1 and 0.4 cM from Re gene, respectively. These results may facilitate marker‐assisted selection of the red leaf trait in kale breeding as well as map‐based cloning of the red leaf trait gene.     

Stay-green expression in early generation sorghum [<Emphasis Type="Italic">Sorghum bicolor</Emphasis> (L.) Moench] QTL introgression lines

Reduced leaf senescence (stay-green) has been demonstrated to improve tolerance of post-flowering moisture stress in grain sorghum. A number of quantitative trait loci (QTLs) associated with stay-green have been identified in sorghum, to facilitate transfer of this trait into adapted genetic backgrounds. This study reports initial evaluations, in both well watered and post-flowering stress environments, following partial introgression (BC₂F₃/BC₁F₄ generations) of four stable stay-green QTLs (StgB, Stg1, Stg3 and Stg4) from donor parent B35 to senescent variety R 16. The majority of the introgression lines had higher leaf chlorophyll levels at flowering (a distinctive trait of the donor parent) and a greater percentage green leaf area during the latter part of grain filling, than did R 16, indicating that the stay-green QTLs were expressed phenotypically in the R 16 background. None of the QTL introgression lines achieved the same level of stay-green as B35, however. Maintenance of a greater relative green leaf area during the latter half of grain filling was related to a greater relative grain yield in two of three post-flowering moisture deficit environments in which the materials were evaluated (r ² = 0.34 in 2004–2005 and r ² = 0.76 in 2005–2006), as was a direct measure of leaf chlorophyll in one of the post-flowering stress environments in which this was measured (r ² = 0.42, P < 0.05). Thus the study provided useful evidence that the marker-assisted backcross transfer of stay-green QTLs from B35 into an adapted, but senescent background has the potential to enhance tolerance of post-flowering drought stress in sorghum.     

Quantitative trait loci for agronomic and seed quality traits in an F2 and F4:6 soybean population

Molecular breeding is becoming more practical as better technology emerges. The use of molecular markers in plant breeding for indirect selection of important traits can favorably impact breeding efficiency. The purpose of this research is to identify quantitative trait loci (QTL) on molecular linkage groups (MLG) which are associated with seed protein concentration, seed oil concentration, seed size, plant height, lodging, and maturity, in a population from a cross between the soybean cultivars ‘Essex’ and ‘Williams.’ DNA was extracted from F₂ generation soybean leaves and amplified via polymerase chain reaction (PCR) using simple sequence repeat (SSR) markers. Markers that were polymorphic between the parents were analyzed against phenotypic trait data from the F₂ and F_4:6 generation. For the F₂ population, significant additive QTL were Satt540 (MLG M, maturity, r² = 0.11; height, r² = 0.04, seed size, r²= 0.06], Satt373 (MLG L, seed size, r² = 0.04; height, r² = 0.14), Satt50 (MLG A1, maturity r² = 0.07), Satt14 (MLG D2, oil, r² = 0.05), and Satt251 (protein r² = 0.03, oil, r² =0.04). Significant dominant QTL for the F₂ population were Satt540 (MLG M,height, r² = 0.04; seed size, r² = 0.06) and Satt14 (MLG D2, oil, r² = 0.05). In the F_4:6 generation significant additive QTL were Satt239 (MLGI, height, r² = 0.02 at Knoxville, TN and r² = 0.03 at Springfield, TN), Satt14 (MLG D2, seed size, r² = 0.14 at Knoxville, TN), Satt373 (MLG L, protein, r² = 0.04 at Knoxville, TN) and Satt251 (MLG B1, lodging r² = 0.04 at Springfield, TN). Averaged over both environments in the F_4:6 generation, significant additive QTL were identified as Satt251 (MLG B1, protein, r² = 0.03), and Satt239 (MLG I, height, r² = 0.03). The results found in this study indicate that selections based solely on these QTL would produce limited gains (based on low r² values). Few QTL were detected to be stable across environments. Further research to identify stable QTL over environments is needed to make marker-assisted approaches more widely adopted by soybean breeders. This revised version was published online in July 2006 with corrections to the Cover Date.