Inheritance of cabbage head formation in crosses of cabbage × ornamental cabbage and cabbage × kale

We investigated the inheritance of head formation in Brassica oleracea by using two crosses, cabbage × ornamental cabbage and cabbage × kale. The degree of head formation (DHF) was classified into nine grades ranging from non‐heading to full heading. DHF in the two F₂ populations showed a continuous distribution. The variance of F₃ offspring selected for full heading or non‐heading was large. The DHF distributions in the F₃ offspring selected for moderate heading or randomly selected F₃ populations were similar to those of the respective F₂ populations, but had smaller average values and variance. However, the realized heritability in F₃ offspring was similar in full‐heading and non‐heading selections. Our findings suggest that head formation is a quantitative trait controlled additively with low dominance effects. In comparisons of leaf developmental patterns among the parents, only cabbage showed a change in leaf shape becoming wider because of the shorter petiole length with increasing leaf position. These findings suggest that cabbage acquired the developmental changes in leaves required to form a head during the process of domestication.     

Relationships between water-use traits and photosynthesis in Brassica oleracea resolved by quantitative genetic analysis

The genetic control of water‐use and photosynthetic traits in Brassica oleracea is resolved by genetic analysis of quantitative trait loci (QTL). Variations in leaf conductance, photosynthetic assimilation rate, leaf thickness and leaf nitrogen content were assessed in a segregating population of F₁‐derived doubled haploid (DH) B. oleracea lines. In addition, stable carbon isotope ratios in leaf organic material were used as a surrogate measure of plant water‐use efficiency. Analysis of an existing linkage map for the population revealed significant QTL on seven linkage groups. Single significant QTL explained between 3.4% and 36.6% of the phenotypic variance in each of the traits measured. The locations of QTL for several traits were found to coincide in a physiologically meaningful way; stable carbon isotope discrimination had QTL co‐locating with leaf level water‐use efficiency, photosynthetic capacity with leaf thickness and nitrogen content and stomatal density with leaf thickness. Taken together, these results suggest that single genes or clusters of genes at these loci may have an influence on the expression of physiologically related traits controlling water‐use and photosynthesis.     

Productivity of F hybrids of botanical varieties ofBrassica oleracea L.

Summary F₁ hybrids were obtained by making pair-crosses between nine botanical varieties ofBrassica oleracea. Although non-inbred parents were used, their yield was surpassed by all F₁ hybrids. The highest yields were shown by the crosses marrow-stem kale × sprouting broccoli and marrow-stem kale × cauliflower while the highest consumable yield (assessed by feeding plants to sheep) was that of F₁ hybrids marrow-stem kale × cauliflower and wild cabbage × cauliflower. A scheme for the production of F₁ hybrid seed is suggested, based upon an adaptation ofThompson's (1964) Triple-cross technique.     

Early-bolting trait and RAPD markers in the specific monosomic addition line of radish carrying the e-chromosome of Brassica oleracea

The specific monosomic addition line of radish, Raphanus sativus, carrying the e chromosome of Brassica oleracea (2n = 19, e‐type MAL) with the genetic background of the late‐bolting cv.‘Tokinashi’ was produced by successive backcrossing of the original e‐type MAL of radish that showed early bolting in the genetic background of the cv. ‘Shogoin’. The early‐bolting trait specific to the e‐type MAL was constantly expressed in the backcrossed progenies (BC₂, BC₃ and BC₄), whereas the reverted radish‐like plants (2n =18) were gradually converted to bolting as late as ‘Tokinashi’. The added e‐chromosome expressed an epistatic effect against the genome of Japanese radish. Its early‐bolting trait was dominant to the late‐bolting trait of ‘Tokinashi’ which may be under the control of a few genes. Moreover, e‐type specific RAPD markers detected in eight primers were invariably transmitted in the backcrossed progenies by ‘Tokinashi’. From the analysis of the characteristics to the e‐type MAL and e‐type specific RAPD markers, it is suggested that the e‐added chromosome of kale (B. oleracea) was transmitted from generation to generation without any recombination with the radish chromosome. The gene(s) for the early‐bolting trait detected in this study may be useful for breeding work in radish, especially in the tropical areas.     

Efficiency of PCR-RF-SSCP marker production in Brassica oleracea using Brassica EST sequences

Efficiencies of SCAR, CAPS and PCR-RF-SSCP marker production were investigated using two combinations of breeding lines in Brassica oleracea. Published EST sequences of B. oleracea, Brassica rapa, Brassica napus, and Arabidopsis thaliana and newly determined nucleotide sequences of anther cDNA clones from B. oleracea were used for designing primer pairs to amplify genes. The percentage of primer pairs yielding DNA amplification of a single gene was higher in primer pairs of B. oleracea (91%) than those of B. rapa (56%) and A. thaliana (17%). Single DNA fragments amplified by 9% of the primer pairs showed polymorphism as SCAR markers between a broccoli line and a Chinese kale line by agarose-gel electrophoresis. CAPS analysis showed different band patterns in 32% of the same-sized DNA fragments, and PCR-RF-SSCP analysis revealed DNA polymorphism in 52% of those showing no DNA polymorphism by CAPS. In total, 71% of the single DNA fragments were converted to DNA markers. The frequency of DNA polymorphism between parental lines of a cabbage F₁ hybrid was lower, 5% by SCAR and 12% by CAPS. However PCR-RF-SSCP analysis revealed DNA polymorphism in 21% of the DNA fragments showing no polymorphism by CAPS. These results suggest that PCR-RF-SSCP analysis enables highly efficient DNA marker production for mapping of genes in Brassica using progeny, even progeny of closely related parents. Analysis of selfed seeds of broccoli F₁ cultivars using PCR-RF-SSCP markers indicated that PCR-RF-SSCP analysis is also applicable to seed purity tests.     

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.     

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.     

Results of crossing Brassica pekinensis (Lour.) Rupr. with B. Oleracea L. Var. Acephala Dc.

Success in obtaining an interspecific hybrid between B. pekinensis and B. oleracea var. acephala was possible through the use of the Chinese cabbage as the seed parent and the use of the growth regulator, N-m-tolylphthalamic acid. The hybrids appeared vegetatively intermediate between the parents except for the inflorescence which appeared more like the kale parent. The F₁ was backcrossed with ease to the Chinese cabbage parent using the hybrid either as the pollen or seed parent. No backcross was made to the kale parent. Segregation in the F₂ was continuous for most characters studied.Michigan Agricultural Experiment Station, Journal Article Number 2588.     

A single dominant gene for downy mildew resistance in broccoli

Downy mildew, incited by Peronospora parasitica (Pers.: Fr.) Fr., is a destructive disease of broccoli (Brassica oleraceaL., Italica Group). Resistant cultivars represent a desirable control method to provide a practical, environmentally benign, and long-term means of limiting damage from this disease. Doubled-haploid (DH) lines developed by us exhibit a high level of downy mildew resistance at the cotyledon stage. To determine the mode of inheritance for this resistance, a resistant DH line was crossed to a susceptible DH line to make an F₁, from which F₂ and backcross (BC) populations were developed. All populations were evaluated for response to artificial inoculation with P. parasitica at the cotyledon stage. All F₁ plants (including reciprocals) were as resistant as the resistant parent, indicating no maternal effect for this trait. F₂ populations segregated approximately 3resistant to 1 susceptible, BC populations using the resistant parent as the recurrent parent contained all resistant plants, and the BC to the susceptible parent segregated 1 resistant to 1 susceptible. These results indicate that resistance is controlled by a single dominant gene. This gene should be easily incorporated into F₁ hybrids and used commercially to prevent downy mildew at the cotyledon stage. This revised version was published online in August 2006 with corrections to the Cover Date.     

Inheritance of internode length,plant form,and annual habit in a cross of cabbage and broccoli (Brassica oleracea var. capitata L. and var. italica Plenck.)

Summary When an inbred line of cabbage, Brassica oleracea L. var. capitata L., was crossed with an inbred line of broccoli B. oleraceae var. italica, the F₁ progeny were vigorous late annuals. All F₁ × broccoli backcross plants and 92% of the 3260 F₂ plants were annuals, while 40% of the F₁ × cabbage backcross plants were biennials. Annual habit is thus dominant and controlled by more than a single gene. Number of days to bud appearance in annuals varied continuously, and was primarily additive in inheritance. F₁ data suggested partial dominance for lateness but this was not supported by the F₂. Internode length was also continuous in distribution and primarily additive in inheritance, but with some dominance for short internodes in the F₁. Cabbage head forming ability was recessive and multigenic, with 2% of the F₂ plants forming heads, of which none were of commercial type and about half bolted as annuals. There was a significant chi square association between biennial habit and tendency for cabbage head formation. Clasping habit of terminal leaves was recessive to open leaves, multigenic, and associated with both cabbage heading and biennial habit.Technical Paper 4836, Oregon Agricultural Experiment Station; from an M.S. thesis by the senior author.     

Genetic analysis of resistance to clubroot (Plasmodiophora brassicae Woron) in two Brassica oleracea groups (ssp. acephala and ssp. botrytis) through diallel analysis

Clubroot, caused by Plasmodiophora brassicae Woron, is a major disease of cruciferous crops and few sources of resistance have been detected and genetically studied in the Brassica oleracea species. In this study, two diallel mating systems using resistant kale lines and susceptible cauliflower lines were performed. Resistance was assessed from a disease index evaluated on young plants artificially inoculated with local isolates of the parasite. Both general and specific combining ability effects (GCA and SCA) and partly reciprocal effects were demonstrated. Resistance inheritance was, however, characterized by a predominance of additive genetic effects (GCA). Three groups of parent lines could be separated; kale lines with very high resistance, kale lines with intermediate resistance and susceptible cauliflower lines. In the two kale groups, two genetically different resistance types were suggested and various recurrent selection procedures are proposed following the diallel results.     

Production and characterization of intergeneric somatic hybrids between Moricandia arvensis and Brassica oleracea

Somatic hybrids were produced between Moricandia arvensis (MaMa, 2n= 28) and Brassica oleracea (CC, 2n= 18) through cell fusion and then characterized by analysing their morphology, cytology, DNA constitution, leaf anatomy and seed fertility. Cell fusion was carried out between greenish protoplasts isolated from the mesophyll of M. arvensis and colourless ones from hypocotyls of B. oleracea. Three plants were generated from one shoot via cuttings and acclimatized in vivo. They closely resembled each other in morphology, exhibiting traits intermediate between the parental species. They were confirmed to be amphidiploids by mitotic and meiotic analyses, being 2n= 46 (MaMaCC), with pollen fertility of about 50%, which was enough to develop the subsequent progenies. Anatomical analysis of the for leaf tissue showed that the bundle sheath cells of the somatic hybrids contained some centripetally arranged organelles, like those of M. arvensis. The hybridity was also confirmed by random amplified polymorphic DNA analysis. Both chloroplast DNA and mitochondrial DNA of the somatic hybrids were estimated to be derived from M. arvensis. In leaf anatomy, the somatic hybrid showed the C₃‐C₄ intermediate trait as in M. arvensis. Many progenies resulted from backcrossing with parental species. The somatic hybrids are expected to be used as bridging plant material to introduce the C₃‐C₄ intermediate trait into Brassica crop species.     

Plasmodiophora brassicae: aspects of pathogenesis and resistance inBrassica oleracea

Summary Clubroot is one of the most damaging diseases inBrassica oleracea crops world-wide. The pathogenicity ofPlasmodiophora brassicae is highly variable between as well as within field populations. Several sources of resistance to clubroot have been identified inB. oleracea. Generally, resistance tends to inherit partly as a recessive, partly as an additive trait, and appears to be controlled by few major genes. Progress in the understanding of the inheritance of resistance is being made through the use of single-spore isolates of the pathogen, and the use of molecular markers for resistance genes.Abbreviations cv cultivar - DH doubled haploid - ECD European Clubroot Differential set - RFLP Restriction Fragment Length Polymorphism     

Association of RFLP markers with trait loci affecting clubroot resistance and morphological characters in Brassica oleracea L.

Summary Resistance to Plasmodiophora brassicae Wor. race 7, the causal agent of the disease clubroot, was examined in an F₂ population of a cross between a clubroot resistant broccoli (Brassica oleracea var. italica) and a susceptible cauliflower (B. oleracea var. botrytis). A genetic linkage map was constructed in the same population based on the segregation of 58 dispersed restriction fragment length polymorphism (RFLP) markers. Associations between the inheritance of RFLP marker genotypes and segregation for disease resistance, morphological and maturity characteristics were examined. For each triat examined, several chromosomal regions marked by RFLP probes appeared to contain trait loci, suggesting that each trait was under polygenic control. RFLP marker linkage to a major factor imparting dominance for clubroot resistance from the broccoli parent was observed in this population. Additionally, RFLP marker linkage to an independently segregating factor contributing clubroot resistance from the cauliflower parent was observed, indicating that it should be possible to use RFLP markers to facilitate selection of transgressive segregants having the combined resistance from both parental sources. In some instances, RFLP markers from the same or closely linked chromosomal regions were associated with both clubroot resistance and morphological traits. Analysis of RFLP marker genotypes at linked loci should facilitate the selection of desired disease resistant morphotypes.     

Quantitative trait loci for scald resistance in barley localized by a non-interval mapping procedure

Three quantitative trait loci (QTL) for scald resistance in barley were identified and mapped in relation to molecular markers using a population of chromosome doubled‐haploid lines produced from the F₁ generation of a cross between the spring barley varieties ‘Alexis’ and ‘Regatta’. Two field experiments were conducted in Denmark and two in Norway to assess disease resistance. The percentage leaf area covered with scald (Rhynchosporium secalis) ranged from 0 to 40% in the 189 doubled‐haploid (DH) lines analysed. One quantitative trait locus was localized in the centromeric region of chromosome 3H, Qryn3, using the MAPQTL program. MAPQTL was unable to provide proper localization of the other two resistance genes and so a non‐interval QTL mapping method was used. One was found to be located distally to markers on chromosome 4H (Qryn4) and the other, Qryn6, was located distally to markers on chromosome 6H. The effects of differences between the Qryn3, Qryn4 and Qryn6 alleles in two barley genotypes for the QTL were estimated to be 8.8%, 7.3% and 7.0%, respectively, of leaf covered by scald. No interactions between the QTLs were found.     

The inheritance of days to flowering in broccoli (Brassica oleracea var. italica)

Summary Crosses between an early flowering inbred broccoli (Brassica oleracea L. var. italica) HS140 and four later maturing inbred lines, S301, S310, s318, and S258, were studied to determine the inheritance of earliness as expressed by days to first open flower. Mean days to first flower for F₁ and F₂ were almost identical, and were close to the mean of the two parents in three crosses, indicating additive inheritance. In the cross HS140×S258, for which no F₁ plants were available, the F₂ mean was closer to the mean of the early parent, but this was likely due to a distortion of the data caused by the very late maturation of S258, the latest maturing parent in the study. Frequency distribution for parents and progenies supported the conclusions from parent and progeny mean values and indicated that days to maturity is a quantitative character, with mostly additive inheritance.Oregon Agricultural Experiment Station Technical Paper No. 7578.     

The transfer of triazine resistance from Brassica napus L. to B. oleracea L. III. First backcross to parental species

Summary Atrazine resistant Brassica napus × B. oleracea F₁ hybrids were backcrossed to both parental species. The backcrosses to B. napus produced seeds in both directions but results were much better when the F₁ hybrid was the pollen parent. Backcrosses to B. oleracea failed completely but BC₁s were rescued by embryo culture both from a tetraploid hybrid (2n = 4x = 37; A₁C₁CC) and sesquidiploid hybrids (2n = 3x = 8; A₁C₁C). Progeny of crosses between the tetraploid hybrid and B. oleracea had between 25 and 28 chromosomes. That of crosses between the sesquidiploid hybrid and B. oleracea had between 21 and 27. A few plants that had chromosome counts outside the expected range may have originated from either diploid parthenogenesis, unreduced gametes or spontaneous chromosome doubling during in vitro culture. Pollen stainability of the BC₁s ranged from 0% to 91.5%. All the BC₁s to B. oleracea were resistant to atrazine.     

Segregation for heading date and stem enlargement in kohlrabi x broccoli crosses

Summary F₁ and F₂ progenies from crosses of an early broccoli (Brassica oleracea var. italica L.) line with kohlrabi (B. oleracea var. gongyloides) indicated that days to flower bud maturity in annual segregates is quantitative and additive in inheritance. It appeared that the biennial kohlrabi parent strongly contributed genetic factors for late maturity. F₁ plants of both crosses, 98% of F₂ plants from early broccoli × kohlrabi, and 81% of the F₂ plants from late broccoli × kohlrabi were annual in 1980. In 1986, 96% of early broccoli × kohlrabi F₂ and 87% of late broccoli × kohlrabi F₂ plants were annual. Kohlrabi stem enlargement showed a continuous range of expression in the F₂ with some dominance of broccoli type. Few F₂ plants were close to the kohlrabi parent in degree of stem enlargement.     

Comparative responses of botanical varieties ofBrassica oleracea L. (brussels sprouts,sprouting broccoli and marrow stem kale) to inbreeding and hybridisation

Summary The effects of inbreeding and hybridisation inBrassica oleracea L. were studied in diallel crosses within four varieties of Brussels sprouts three varieties of sprouting broccoli and one kale variety. In addition to each intra-varietal diallel within the two horticultural brassicas, plants of one of the other varieties were also included so that the first diallel analysis could be compared with an intra-plus-intervarietal analysis.In general it was found that the total harvest yield of cultivars of the two horticultural types was controlled by an additive genetic system while whole plant weight of all three brassicas was subject to gene interaction. Selfing for one generation reduced the mean harvest yield of Brussels sprout and sprouting broccoli by 26 and 3%, respectively, compared with their intravarietal F₁'s, while whole plant weight of Brussels sprouts, sprouting broccoli and marrow-stem kale was reduced by 21, 26 and 44%, respectively.Kale plants exhibited significant correlations of plant height with plant yield in 2 years and it is therefore possible to achieve some improvement in yield by selecting for height alone.It was postulated that in kale the greatest yield increases should be obtained by the production of hybrids, but hybridisation with subsequent selection is probably sufficient to increase the yield of the two horticultural brassicas still further.