The stabilization of chromosome numbers and the maintenance of euploidy in Brassicoraphanus

Summary To examine whether chromosome numbers of Brassicoraphanus (amphidiploids between Brassica japonica Sieb. and Raphanus sativus L.) are stable or not, the following four items were studied with some plants during the 2nd–11th generations: (1) chromosome numbers of open-pollinated progenies from eu-, hyper-, and hypoploids, (2) chromosome distribution at metaphase II in these plants, (3) frequency of euploids in relation to flower colour and generation, (4) seed fertility in eu-and aneuploids in relation to flower colour. In early generations, hyper-and hypoploids were frequently produced from euploids. In later generations, however, the chromosome number became less unstable. In euploids (2n=38), chromosome numbers at metaphase II showed some variation, and the mean frequency of the euploid chromosome number (n=19) was approximately 78%. This value was almost the same in white-and yellow-flowered plants through early and late generations. Nevertheless, yellow-flowered plants tended to produce euploids more frequently than did white-flowered ones. It is assumed that the difference in euploid productivity between yellow-and white-flowered plants is due to the difference in seed fertility between them. The progeny of each hypoploid showed higher chromosome numbers than their parents. The progeny of each hyperploid showed lower chromosome numbers than their parents: they were approaching to euploidy. This phenomenon, together with higher fertility of euploids and lower fertility of aneuploids, will favor the maintenance of euploidy of this strain.     

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.     

Inheritance of neurotoxin (ODAP) content, flower and seed coat colour in grass pea (Lathyrus sativus L.)

Summary A strong epidemiological association is known to exist between the consumption of grass pea and lathyrism. A neurotoxin, -N-Oxalyl-L-, -diaminopropanoic acid (ODAP) has been identified as the causative principle. This study was undertaken to investigate the mode of inheritance of the neurotoxin ODAP, flower and seed coat colour in grass pea. Five grass pea lines with low to high ODAP concentration were inter-crossed in all possible combinations to study the inheritance of the neurotoxin. Parents, F₁ and F₂ progenies were evaluated under field condition and ODAP analyzed by an ortho-phthalaldehyde spectrophotometric method. Many of the progenies of low x low ODAP crosses were found to be low in ODAP concentration indicating the low ODAP lines shared some genes in common for seed ODAP content. The F₁ progenies of the low ODAP x high ODAP crosses were intermediate in ODAP concentration and the F₂ progenies segregated covering the entire parental range. This continuous variation, together with very close to normal distribution of the F₂ population both of low x low and low x high ODAP crosses indicated that ODAP content was quantitatively inherited. Reciprocal crosses, in some cases, produced different results indicating a maternal effect on ODAP concentration. Blue and white flower coloured lines of grass pea were inter-crossed to study the inheritance of flower colour. Blue flower colour was dominant over the white. The F₂ progenies segregated in a 13:3 ratio indicating involvement of two genes with inhibiting gene interactions. The gene symbol LB for blue flower colour and LW for white flower colour is proposed.     

Variation in progenies of an Arachis hypogaea x diploid wild species hybrid

Summary Derivatives of a cross between cultivated peanuts, Arachis hypogaea L. (2n=40), and the wild species collection GKP 10017 (2n=20) were compared morphologically, for leafspot resistance and for yield. The objective of the study was to determine the effects of wild species germplasm on the A. hypogaea genome. The sterile F₁ hybrid which resulted from crossing the two species was treated with colchicine to restore fertility at the 6x ploidy level. The resulting hexaploid was cytologically unstable and progeny lost chromosomes until stability was regained at the 2n=40 chromosome level. Forty-seven characters were used to analyze the variation among plants in the tetraploid interspecific hybrid population. The plants were compared to four cultivated lines plus GKP 10017. Many hybrids were intermediate to the two parents in morphology. Individual traits such as growth habit, pod and seed size, elongation of the constricted area between pods, nodulation and leaflet size were altered by the presence of GKP 10017 germplasm in many of the hybrid plants. Cercospora arachidicola Hori and Cercosporidium personatum (Berk. & Curt.) Deighton resistances were evaluated for all plants. Several hybrids had few lesions due to either leafspot pathogen. In addition, 24 largeseeded interspecific hybrid selections were compared to the cultivated variety NC 5 for yield. Five selections were superior to both parents at p=0.01. Morphology, disease resistance and yields appeared to be greatly influenced by the wild species GKP 10017 germplasm in plants of the interspecific hybrid population. The potentials of using wild species for improvement of the cultivated peanut are discussed.Paper number 5948 of the journal series of the North Carolina Agricultural Research Service, Raleigh, NC 27650. The investigation was supported in part by ICRISAT and SEA-CR grant no. 701-15-51.     

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.     

Cytomorphological studies in an intergeneric hybrid between Gossypium hirsutum L. (2n=52) and Hibiscus panduraeformis Burm.

Summary An intergeneric hybrid (2n=38) between Gossypium hirsutum L. (2n=52) × Hibiscus panduraeformis Burm. (2n=24) was obtained by pollinating about 2000 flower buds of G. hirsutum var. Gregg Male Sterile with pollen from H. panduraeformis. The F₁ hybrid was intermediate in plant habit, but possessed gossypol glands and nectaries on the leaves, bolls containing seeds with fuzz and lint as dominant characters of G. hirsutum. Flowers with yellow corolla and anthers; purple petal spot, profuse growth of epidermal hairs on all plant parts including the boll sutures, and jassid tolerance were dominant characters of H. panduraeformis. The partial fertility of the F₁ indicated the possibilities of combining jassid and drought tolerance of H. panduraeformis with the desired economic characters of G. hirsutum for rainfed cultivation.The F₁ hybrid showed various meiotic irreguarities and about 40% pollen sterility. Formation of the normal bivalents occurred quite frequently suggesting a close relationship between the parental species. The sterility observed in the hybrid may be due to small structural differences between the chromosomes of the two genera and meiotic abnormalities.     

Chromosomal and genic structure of Brassicoraphanus related to seed fertility and the presentation of an instance of improvement of its fertility

Summary In order to elucidate the mechanism of low fertility of Brassicoraphanus, i.e., amphidiploids between Brassica japonica Sieb. and Raphanus sativus L., the chromosome number of 253 plants was studied during the 3rd–9th generations for their seed fertility. Meiotic irregularity showed no connection with degree of sterility. Brassicoraphanus consisted of euploids (2n=38), hyperploids (2n=39–43) and hypoploids (2n=34–37) with white or yellow flowers. The number of plants was highest in euploids and became lower as the chromosome number diverged from the euploid number. Further, seed fertility was highest and the range of its variation widest in euploids. The seed fertility of aneuploids became lower and its variation narrower in proportion to the number of chromosomes additional to or missing from the euploid number. Yellow-flowered plants were superior in seed fertility to white-flowered plants. Seed fertility of plants is primarily affected by their chromosome numbers and secondarily modified by genic effects. As a whole, seed fertility of Brassicoraphanus increased gradually and its variation widened with the advance of generations. This was explained mainly by the increase of balanced combinations of genes.     

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.     

Barriers to hybridization of Solanum bulbocastanum Dun. and S. Verrucosum Schlechtd. and structural hybridity in their F1 plants

Summary Results of reciprocal crosses between Solanum verrucosum (2n=2x=24) and S. bulbocastanum (2n=2x=24) are described in terms of pollen tube behaviour in styles, of berry and seed set, of fertility and of meiotic behaviour of the F₁ hybrids. Pollen tube growth of S. verrucosum is strongly inhibited in styles of S. bulbocastanum, whereas no inhibition is observed in the reciprocal cross. Therefore S. bulbocastanum x S. verrucosum fails to set berries or seeds, whereas the reciprocal cross produces both berries (54.4% berry set) and seeds (0.3 per berry). Only 14.6% of the seeds germinate. Both the diploid and corresponding tetraploid hybrid plants are vigorous, flower abundantly, have a rather regular meiosis (mainly rod bivalents), but show a high degree of cytoplasmic-genic male sterility. Crossability of the diploid hybrid plants is nil when used as pollen parents and near to zero when used as pistillate parents. The barriers to hybridization of the parent species (unilateral inhibition of pollen tube growth, somatoplastic sterility, cytoplasmic-genic male sterility and structural differences of the parental chromosomes) are discussed and methods are suggested to overcome these barriers.     

Crosses between Beta intermedia bunge and B. vulgaris L.

Summary A tetraploid annual male sterile form of Beta vulgaris L. (2n=4x=36) was crossed with the wild beet species Beta intermedia Bunge (2n=36). The resulting F₁-plants were male sterile annuals being two or three times back-crossed to diploid and tetraploid sugar and fodder beets in the next years. Apart from tetraploid material (36 chromosomes) hexaploid (54 chromosomes) and a number of aneuploid plants developed.The results obtained justify the conclusion that, at a tetraploid level the material mostly propagates apomictically after the F₁ generation. The presence of penta-, hexa-, septa-and even octaploid plants might be explained by assuming that no meiosis has taken place in the crossing partners. Triploid plants are sometimes found in the progeny of hexaploid material and may presumably be considered haploids. Moreover some pentaploid plants were found in the progeny of the open pollinated F₁ which after two generations of bagging are still pentaploids although they produce no pollen. This is another clear indication of apomictic reproduction.The tetraploid generation from the cross between the hexaploid material and diploid sugar beets probably contains the best prospects for breeding.     

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.     

Production of intergeneric hybrids between Brassica juncea and Diplotaxis virgata through ovary culture, and the cytology and crossability of their progenies

The cytogenetic study was investigated in the intergeneric F₁ hybrid, F₂and backcross progenies (BC₁). The plants used were Brassica juncea(2n=36) and Diplotaxis virgata(2n=18). Three intergeneric F₁ hybrids between two species were produced through ovary culture. They showed 36 chromosomes. It might consist one genome of B. juncea and two genomes of D. virgata. The morphology of the leaves resembled that of B. juncea. The color of the petals was yellow that was like in D. virgata. The size of the petal was similar to that of B. juncea. The mean pollen fertility was15.3% and the chromosome associations in the first meiotic division were(0–1)_IV+(0–2)_III+(8–12)_II+(12–20)_I. Many F₂ and BC₁seeds were harvested after open pollination and backcross of the F₁ hybrids withB. juncea, respectively. The F₂seedlings showed different chromosome constitutions and the range was from 28 to54 chromosomes. Most seedlings had 38chromosomes followed by 36, 40 and 54. The BC₁ seedlings also showed different chromosome constitutions and the range was from 29 to 62. Most seedlings had both 40and 54 chromosomes followed by 36, 46 and52. In the first meiotic division of F₂ and BC₁ plants, a high frequency of bivalent associations was observed in all the various kinds of somatic chromosomes. Many F₃ and BC₂ seeds were obtained by self-pollination and open pollination of both F₂ and BC₁ plants, and by backcrossing both F₂ and BC₁plants with B. juncea, respectively,especially, three type progeny with 36, 40or 54 chromosomes. The somatic chromosomes of the F₃ and BC₂ plants were further investigated. The bridge plants between B. juncea and D. virgata with 36 chromosomes may be utilized for breeding of other Brassica crops as well as B. juncea. This revised version was published online in July 2006 with corrections to the Cover Date.     

Intergeneric hybrid between Brassica napus and Diplotaxis harra through ovary culture and the cytogenetic analysis of their progenies

Brassica napus (2n = 38) and Diplotaxis harra (2n = 26) were used to investigate gene transfer from D. harra to B. napus. Intergeneric F₁ hybrids (dihaploid 2n = 32 chromosomes) were obtained through ovary culture. The chromosome associations in the first meiotic division was (0–2)_III + (2–10)_II + (12–28)_I. Many seeds were harvested in the F₁ hybrid after backcrossing with B. napus, and from open pollination of the F₁ hybrid. Somatic chromosome numbers of BC₁ and hybrid plants varied from 2n = 26 to 52. In the first meiotic division, high frequencies of bivalent association and relatively low pollen fertility were observed. BC₂ plants generated from the BC₁ plants with 2n = 38 chromosomes, 69.6% showed 2n = 38 chromosomes. Many aneuploids with addition and deletion of chromosomes were also obtained. A bridge plant between B. napus and D. harra with 2n = 32 chromosomes should be valuable material for the breeding of brassica crops.     

Relationships between African species of the genus Cucumis L. Estimated by the production,vigour and fertility of F1 hybrids

Summary The production, vigour, and fertility of F₁ hybrids between nine African species of the genus Cucumis L. were studied as a measure of the relationships between the species. Hybrid plants were obtained from 29 out of the 72 possible cross combinations. Two F₁ hybrids died as seedlings, and 27 hybrids were raised to maturity. Pollen production and stainability varied greatly amongst these hybrids, as did fruit and seed set following self pollination and backcrossing with either parental species. The fruit shape of the hybrids was always intermediate between that of both parental species.Two species appeared to be closely related: C. prophetarum L. and C. anguria var. longipes A. Meeuse. Most other species produced highly to moderately fertile F₁ hybrids with at least one other species. C. metuliferus Naud. produced only sterile hybrids with C. zeyheri Sond. 2x. The results sustain the recent taxonomic classification of the genus (Jeffrey, 1980).     

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.     

Expression of barley yellow dwarf virus and Russian wheat aphid resistance genes in and fertility of spring wheat × triticale hybrids and backcross lines

Summary The Barley Yellow Dwarf Virus disease (BYDV) and the Russian wheat aphid (RWA) Diuraphis noxia (Mordvilko) have caused significant losses to wheat and barley in several areas of the world. Important sources of resistance to both BYDV and RWA have been found in Triticale. Different generations of interspecific wheat x Triticale crosses were produced and the progenies were screened for BYDV and RWA tolerance. Plants with equal chromosome numbers showed different levels of fertility. A significant correlation was observed between pollen fertility and seed set in primary florets (r=0.57). In generaL, pollen fertility, seed set and the number of euploid plants (2n=42) increased from one generation to the next. The expression of BYDV tolerance varied from population to population. Additive effects were predominant in F₁ and some backcross populations. A dominant effect of rye tolerance genes was also observed in few populations. A monogenic trait or a quantitative (polygenic) character would not agree with the observed segregation patterns. The heritability of this oligogenic tolerance was quite different between populations and in many populations the tolerance genes were only partially expressed. Some transgressive segregation for tolerance and sensitivity was demonstrated. The genes controlling tolerance to RWA in Triticale lines, Muskox 658 and Nord Kivu were not expressed in advanced lines resistant to BYDV. This indicates that tolerance genes for BYDV and RWA in these lines are located on different chromosomes.     

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.     

Independent Inheritance of Flower Colour and Male-Fertility Restorer Characters in Brassica napus L.

Available material of oilseed (Brassica napus L., AACC) comprises two yellow-flowered breeding lines and a white/pale-flowered line of resynthesized rape. The flower colour white/pale is dominant over yellow, and is controlled by a gene located in the C-genome. The yellow-flowered genotypes acted as restorer lines and the white/pale-flowered genotype as a maintainer line in a cytoplasmic male sterility system. The segregation pattern of flower colour and male fertility restorer characters were studied in F₂ generations of crosses between these lines, also in a three-way cross additionally including a yellow flowered B. campestris (AA) line. Evidense was obtained in support of the conclusion that the flower colour and male fertility restorer characters are monogenically controlled and independently inherited. Whether the male fertility restorer gene is located in the A or C genome remains to be determined.     

Pollen grain germination and pollen tube growth following in vivo and in vitro self and interspecific pollinations in annual Cicer species

Summary Germination of pollen grains and growth of pollen tubes were studied to determine the cause of barreness in crosses among annual Cicer species. In vivo and in vitro time-course studies and fluorescent microscopy revealed no pollination incompatibility among the selfs, crosses and reciprocals of C. arietinum L., C. reticulatum Lad. and C. cuneatum Rich. In general, Cicer pollen grains germinated and grew on styles of Cicer species. Pollen tube growth was characterized by irregularly spaced and intermittent callose deposits. Failure of seed formation in interspecific pollinations may be attributed to the slowness of pollen tube growth or collapse of fertilized ovules. In addition to these causes, shortness of stamens and sparsity of pollen grains were responsible for flower drop in natural selfs. Although the number of pollen tubes entering the micropyle in interspecific pollinations was low, it may be possible to grow the fertilized ovules on an artificial medium to obtain F₁ plants.     

Interspecific hybridization inBrassica juncea × Brassica hirta using embryo rescue

Summary Interspecific hybrids have been obtained in an otherwise incompatible cross betweenBrassica juncea × Brassica hirta through the in vitro culture of hybrid ovules and ovaries. The best response was observed from ovules and ovaries cultured 10–15 and 5–7 days after pollination respectively on a basal medium supplemented with indoleacetic acid, kinetin and casein hydrolysate. In some cases the basal cut end of the ovaries proliferated to form callus and shoots. The in vitro-derived hybrid seeds varied in their colour, size and shape, and the F₁ plants in the field showed a large diversity in their morphological traits. The hybrids were sterile, and had an intermediate number of chromosomes (2n=30).