Interspecific transfer of Brassica juncea-type high blackleg resistance to Brassica napus

Summary Complete resistance to Leptosphaeria maculans, the cause of blackleg of oilseed rape (Brassica napus), was transferred from B. juncea to B. napus through an interspecific cross. B. juncea-type complete resistance (JR) was recognized first in one F₃ progeny (Onap^JR) by the absence of leaf-lesions on seedlings and canker-free adult plants. The commercially important characters of B. napus were retained in advanced lines of Onap^JR, which combined JR with low erucic acid levels (<0.5%), high seed yield and variable maturity dates.JR appeared to be inherited as a major gene or genes. Segregation for resistance and susceptibility contintied to occur during later generations of selection of Onap^JR. JR was readily transferred from Onap^JR to other suitable B. napus cultivars or lines with partial resistance to blackleg and resulted in highly vigorous carly generation selections adapted to cold, wet situations along with complete resistance to blackleg.     

Strong heritability across years and sites for pod wall proportion and specific weight in Lupinus albus and genotypic correlation with other pod and seed attributes

Thinner pod walls might contribute to increased yield potential and adaptation of lupins. Fourteen autumn‐sown genotypes of Lupinus albus L. were evaluated over 2 years at four sites in southern Chile. Pod wall proportion (PWP), pod wall specific weight (WSW) and five other pod‐related characters were measured to estimate their genetic variation, heritability across years and sites, and genotypic and phenotypic correlations among the characters. Ranges for PWP and pod WSW were 22.9−43.1% and 16.4−37.4 mg/cm², respectively. PWP in cultivated L. albus was similar to that reported in L. angustifolius; however, specific weight was substantially lower. Highly significant genotypic effects were found for both characters. Broad sense heritabilities were high for PWP (0.63) and moderate for pod WSW (0.46). These characters were significantly correlated, more so genotypically (r_g = 0.67) than phenotypically (r_ph = 0.36). Selection for low PWP should lead to lower pod wall thickness or density, both of which contribute to pod WSW. Nevertheless, selection for pod WSW could achieve further reductions, particularly if practised among low PWP materials.     

Gene‐for‐gene resistance is expressed in cotyledons,leaves and pods,but not during late stages of stem colonization in the Leptosphaeria maculans–Brassica napus pathosystem

The blackleg fungus, Leptosphaeria maculans, interacts with canola (Brassica napus) in a gene‐for‐gene manner. These major resistance genes are well characterized in the seedling stage of development, but not in other plant organs. Cotyledons, leaves, pods and stems of plants of two cultivars of B. napus, each harbouring a different major resistance gene (Rlm1 and Rlm4), were inoculated with two individual L. maculans isolates with different alleles of the corresponding avirulence genes (AvrLm1, avrLm4 and avrLm1, AvrLm4), and the disease phenotype in terms of lesion development was determined. Major gene resistance was expressed in cotyledons, all leaves and during pod set, but not in the stems of the adult plant. This is the first time major gene resistance has been shown to be effective in B. napus pods.     

Different Behavior of Brassica juncea and B. carinata as Sources of Phoma lingam Resistance in Experiments of Interspecific Transfer to B. napus

With the aim to transfer Phoma lingam resistance into rape, successful interspecific crosses were made between three oilseed rape varieties (Brassica napus) and the resistant species B. carinata and B. carinata. Although both hybrid types B. napus×B. juncea and B. napus×B. carinata showed the same high level of resistance as the respective resistant parent, the resistance could be only transferred from juncea crosses. After three backcross generations, lines morphologically undistinguishable from rape, fertile, and with a high degree of resistance were obtained. The resistance of B. carinata was practically lost in the first backcross. A possible explanation of this different behavior could be a higher recombination between the genomes B and C (juncea crosses) than between B and A (carinata crosses). The: applied embryo culture increased the yield of hybrids and first backcross plants and reduced considerably the generation time.     

Biochemical markers for cabbage seedpod weevil (<Emphasis Type="Italic">Ceutorhynchus obstrictus</Emphasis> (Marsham)) resistance in canola (<Emphasis Type="Italic">Brassica napus</Emphasis> L.)

In the last decade, the cabbage seedpod weevil (Ceutorhynchus obstrictus (Marsham)) has become a major insect pest of canola (Brassica napus L.) in Canada reducing seed yields up to 35%. Therefore, the benefits of developing weevil resistant germplasm to canola breeders and the industry would reduce input costs, pesticide use, environmental degradation and increase yield. Yellow mustard (Sinapis alba L.) is resistant to C. obstrictus (CSPW), although the exact mechanism is not known (McCaffrey et al. 1999). A unique canola population was generated at the University of Guelph from a cross between B. napus and S. alba through embryo rescue and backcrossed to canola several times prior to double haploid (DH) production. Approximately one-half of this DH population had canola quality glucosinolate concentration (<16 μmol/g) and was used for further breeding. The hypothesis was that some DH progeny from this cross inherited resistance to CSPW from S. alba. Weevil infestation levels were assessed for the B. napus × S. alba BC2 and BC3 DH populations in the field over 7 years in Alberta where weevil pressure is strong to establish the resistant or susceptible status of these lines. The basic objectives for this study were to confirm field resistance in the B. napus × S. alba germplasm in Ontario and to identify any biochemical markers associated with resistance/susceptibility. Canola doubled haploid lines derived from BC2 or BC3 families were field screened for resistance (R) followed by chemical analysis of glucosinolates to detect biochemical polymorphisms correlated with CSPW resistance using High Performance Liquid Chromatography (HPLC). Two polymorphic peaks were found, one each, from extracts of upper cauline leaves and Stage 3 pod seed, with retention times of ~23 and 19 min, respectively. These HPLC peaks consistently correlated with larval infestation data and the peak differences between R and S DH lines were significant. Therefore, these two peaks can be considered as biochemical markers in this breeding germplasm and may play a role in rapid and early detection of CSPW resistance.     

Development of molecular markers linked to the Leptosphaeria maculans resistance gene Rlm6 and inheritance of SCAR and CAPS markers in Brassica napus × Brassica juncea interspecific hybrids

Leptosphaeria maculans causes blackleg disease on Brassica napus, an economically important oilseed crop. Brassica juncea has high resistance to blackleg and is a source for the development of resistant B. napus varieties. To transfer the Rlm6 resistance gene from B. juncea into B. napus, an interspecific cross between B. napus “Topas DH16516” and B. juncea “Forge” was produced, followed by the development of F₂ and F₃ generations. Sequence characterized amplified region (SCAR) and cleaved amplified polymorphic sequence (CAPS) markers linked to the L. maculans resistance gene Rlm6 were developed. Segregation of SCAR and CAPS markers linked to Rlm6 were confirmed by genotyping of F₂ and F₃ progeny. Segregation of CAPS markers and phenotypes for blackleg disease severity in F₂ plants had a Mendelian ratio of 3:1 in resistant vs. susceptible plants, respectively, supporting the assumption that genetic control of resistance was by a single dominant gene. The molecular markers developed in this study, which show linkage with the L. maculans resistance gene Rlm6, would facilitate marker‐assisted backcross breeding in a variety development programme.     

Identification of Genes Controlling Pod Length in Spring Rapeseed, Brassica napus L., and their Utilization for Yield Improvement

Experiments were undertaken to determine the inheritance of pod length in a cross with spring rapesecd, Brassica napus, and to assess the value of pod length as a criterion of selection for high seed yield. Analyses of patterns of variation in F₂; and backcross populations derived from a cross between a short-pod line TB42 and long-pod line CA553 indicated that much of the variation in pod length could be attributed to two major genes interacting in a complementary manner. Short-pods were produced when cither one or both genes were homozygous for the recessive allele. Analyses of F₃ progenies of selected F₂ and inbred-backcross lines derived from the same cross supported the two-gene hypothesis but also indicated that the effects of the major genes on pod length were possibly modified by genes of minor effect. Field testing of families derived from random intermating between F₂, plants of the TB42 × CA553 cross showed that number of pods per plant varied independently of pod length, but seed weight per pod tended to increase with increasing pod length. As a result, families with the longest pods generally had significantly higher yields than those with short pods. It was concluded that yield improvement in B. napus could be achieved through introgression of long-pod genes into cultivars with an appropriate genetic background to ensure that selection for the long-pod character would be accompanied by an increase in seed weight per pod with little or no reduction in number of pods per plant.     

Identification of RAPD markers linked to recessive genes conferring siliqua shatter resistance in Brassica rapa

Shattering of siliquae causes significant seed loss in canola (Brassica napus) production worldwide. There is little genetic variation for resistance to shatter in canola and, hence, the trait has been studied in B. rapa. Previous studies have shown two randomly segregating recessive genes to be responsible for shatter resistance. Three random amplified polymorphic DNA markers were identified as being linked to shatter resistance using bulked segregant analysis in a F₃B. rapa population. The population was derived from a cross between a shatter‐susceptible Canadian cultivar and a shatter‐resistant Indian line. Of the three markers, RAC‐3₉₀₀ and RX‐7₁₀₀₀ were linked to recessive sh1 and sh2 alleles, and SAC‐20₁₃₀₀ was linked to both dominant Sh1 and Sh2 alleles. The common marker for the dominant wild‐type allele for the two loci was explained to have resulted from duplication of an original locus and the associated markers through chromosome duplication and rearrangements in the process of evolution of the modern B. rapa from its progenitor that had a lower number of chromosomes. Segregation data from double heterozygous F₃ families, although limited, indicated the markers were not linked to each other and provided further evidence for the duplication hypothesis.     

The effect of the cytoplasms of Brassica napus and B. juncea on some characteristics of B. carinata, including flower morphology

This study was conducted to assess the cytoplasm effects of Brassica napus and B. juncea on the some characteristics of B. carinata, as well as the phylogenetic distances separating the three species. Alloplasmic lines of B. carinata were developed from B. napus × B. carinata and B. juncea × B. carinata hybrids by recurrent backcrossing to the BC₇ generation. Sixteen populations from three generations were compared for a number of characteristics. Plants with the cytoplasm of B. napus flowered later, had shorter filaments and longer pistils, lower pollen amount, lower seed set, lower petal length and width and different petal color; plants with the cytoplasm of B. juncea had shorter pistils and filaments, and lower petal length and width than their corresponding euplasmic sibs, respectively. The results suggest that the cytoplasm is involved in the development of flower organs. The natural species, B. carinata showed a balance between the nucleus and cytoplasm. The cytoplasm from B. napus showed a stronger disturbing effect than that of B. juncea, suggesting that B. carinata might be genetically closer to B. juncea than to B. napus. The significant difference in the alloplasmic effect of the cytoplasms of B. napus and B. juncea also suggests that in B. carinata the B genome may play a greater role than the C genome. An erratum to this article can be found at     

A study on disease variation in the populations of an interspecific cross of Brassica juncea L. x B. napus L.

Summary F₁ behaviour and F₂ variation in disease reaction were studied in the interspecific cross Brassica juncea x B. napus. Gene(s) for adult resistance to blackleg (Leptosphaeria maculans) were found to be present in the A genome of B. juncea and could be transferred to B. napus. Gene(s) for complete (seedling plus adult) resistance in B. juncea appeared to be located in the B genome. The chance of their transfer to the oilseed rapes (B. napus or B. campestris) would therefore seem to be remote.     

Foliar fungal disease‐resistant introgression lines of groundnut (Arachis hypogaea L.) record higher pod and haulm yield in multilocation testing

Introgression lines (ILs) of groundnut with enhanced resistance to rust and late leaf spot (LLS) recorded increased pod and haulm yield in multilocation testing. Marker‐assisted backcrossing (MABC) approach was used to introgress a genomic region containing a major QTL that explains >80% of phenotypic variance (PV) for rust resistance and 67.98% PV for LLS resistance. ILs in the genetic background of TAG 24, ICGV 91114 and JL 24 were evaluated for two seasons to select 20 best ILs based on resistance, productivity parameters and maturity duration. Multilocation evaluation of the selected ILs was conducted in three locations including disease hot spots. Background genotype, environment and genotype × environment interactions are important for expression of resistance governed by the QTL region. Six best ILs namely ICGV 13192, ICGV 13193, ICGV 13200, ICGV 13206, ICGV 13228 and ICGV 13229 were selected with 39–79% higher mean pod yield and 25–89% higher mean haulm yield over their respective recurrent parents. Pod yield increase was contributed by increase in seed mass and number of pods per plant.     

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.     

Identification of QTL underlying the resistance of soybean to pod borer, <Emphasis Type="Italic">Leguminivora glycinivorella</Emphasis> (Mats.) obraztsov,and correlations with plant,pod and seed traits

Soybean (Glycine max L. Merr.) pod borer (Leguminivora glycinivorella (Mats.) Obraztsov) (SPB) results in severe loss in soybean yield and quality in certain regions of the world, especially in Northeastern China, Japan and Russia. The aim here was to evaluate the inheritance of pod borer resistance and to identify quantitative trait loci (QTL) underlying SPB resistance for the acceleration of the control of this pest. Used were the 129 recombinant inbred lines (RILs) of the F_5:6 derived population from ‘Dong Nong 1068’ × ‘Dong Nong 8004’ and 131 SSR markers. Correlations between the percentage of damaged seeds (PDS) by pod borer and plant, pod and seed traits that were potentially related to SPB resistance were analyzed. The results showed highly significant correlations between PDS by pod borer and plant height (PH), maturity date (MA), pod color (PC), pubescence density (PB), 100-seed weight (SW) and protein content existed. Soybeans with dwarf stem, light color of pod coat, small seeds, lower density of pubescence, early maturity and low content of protein seemed to have higher resistance to SPB. The correlated traits had potential to inhibit egg deposition and thereby to decrease the damage by SPB. Three QTL directly associated with the resistance to SPB judged by PDS at harvest were identified. qRspb-1 (Satt541–Satt253) and qRspb-2 (Satt253–Satt314) were both on linkage group (LG) H and qRspb-3 (Satt288–Satt199) on LG G. The three QTL explained 10.96, 9.73 and 11.59% of the phenotypic variation for PDS, respectively. In addition, 12 QTL that underlay 10 of 13 traits potentially related with SPB resistance were found. These QTL detected jointly provide potential for marker assisted selection to improve cultivar resistance to SPB. Guiyun Zhao, Jian Wang, and Yingpeng Han have equal contribution to the paper.     

Phenotypic variation in progenies from somatic hybrids between Brassica napus and Sinapis alba

A number of variant phenotypes, such as long siliques, high density of pods, increased seed number per pod and yellow seed color resembling Sinapis alba, were selected from intergeneric somatic hybrids between Brassica napus and Sinapis alba through successive backcrosses. Resistance to Sclerotinia sclerotiorum among 24 BCF₄ lines was also tested by inoculation with mycelia, and four lines showed stronger resistance than the resistant rapeseed variety ‘Zhongshuang 9’. Comparative anatomy studies on mature seed revealed that the seed coat pigments were mainly distributed in the palisade layer, which is considerably thinner in S. alba compared to B. napus. The area index of protein bodies in cotyledon cells was highest in S. alba, lowest in B. napus and intermediate in the intergeneric progenies. This study demonstrates that wide hybridization can enable exploitation of valuable trait diversity from S. alba for broadening the genetic basis for rapeseed breeding.     

Relationship between hybridization frequency of Brassica juncea × B. napus and distance from pollen source (B. napus) to recipient (B. juncea) under field conditions in Japan

Several imported transgenic canola (Brassica napus) seeds have been spilled and have grown along roadsides around import ports. B. juncea, a relative of B. napus with which it has high interspecific crossability, is widely distributed throughout Japan. There is public concern about the harmful impacts of feral B. napus plants on biodiversity, but spontaneous hybridization between spilled B. napus and weedy B. juncea populations is hardly revealed. We evaluated the relationship between the hybridization frequency of B. juncea × B. napus and their planting distance in field experiments using the mutagenic herbicide-tolerant B. napus cv. Bn0861 as a pollen source for hybrid screening. The recipient B. juncea cv. Kikarashina was planted in an experimental field with Bn0861 planted in the center. No hybrids were detected under natural flowering conditions in 2009. However, the flowering period was artificially kept overlapping in 2010, leading to a hybridization frequency of 1.62% in the mixed planting area. The hybridization frequency decreased drastically with distance from the pollen source, and was lower under field conditions than estimated from the high crossability, implying that spontaneous hybridization between spilled B. napus and weedy B. juncea is unlikely in the natural environment.     

Genetic basis of pod borer (Helicoverpa armigera) resistance and grain yield in desi and Kabuli Chickpea (Cicer arietinum L.) under unprotected conditions

A series of half-diallel crosses involving early, medium and late maturity desi and kabuli type chickpea (Cicer arietinum L.) genotypes with stable resistance to Helicoverpa pod borer, along with the parents, were evaluated at two locations in India to understand the inheritance of pod borer resistance and grain yield. Inheritance of resistance to pod borer and grain yield was different in desi and kabuli types. In desi type chickpea, the additive component of genetic variance was important in early maturity and dominance component was predominant in medium maturity group, while in the late maturity group, additive as well as dominance components were equally important in the inheritance of pod borer resistance. Both dominant and recessive genes conferring pod borer resistance seemed equally frequent in the desi type parental lines of medium maturity group. However, dominant genes were in overall excess in the parents of early and late maturity groups. In the kabuli medium maturity group, parents appeared to be genetically similar, possibly due to dispersion of genes conferring pod borer resistance and susceptibility, while their F₁s were significantly different for pod borer damage. The association of genes conferring pod borer resistance and susceptibility in the parents could be attributed to the similarity of parents as well as their F₁s for pod borer damage in kabuli early and late maturity groups. Grain yield was predominantly under the control of dominant gene action irrespective of the maturity groups in desi chickpea. In all the maturity groups, dominant and recessive genes were in equal frequency among the desi parental lines. Dominant genes, which tend to increase or decrease grain yield are more or less present in equal frequency in parents of the early maturity group, while in medium and late maturity groups, they were comparatively in unequal frequency in desi type. Unlike in desi chickpea, differential patterns of genetic components were observed in kabuli chickpea. While the dominant genetic component was important in early and late maturity group, additive gene action was involved in the inheritance of grain yield in medium duration group in kabuli chickpea. The dominant and recessive genes controlling grain yield are asymmetrically distributed in early and medium maturity groups in kabuli chickpea. The implications of the inheritance pattern of pod borer resistance and grain yield are discussed in the context of strategies to enhance pod borer resistance and grain yield in desi and kabuli chickpea cultivars.     

Possibilities of direct introgression from Brassica napus to B. juncea and indirect introgression from B. napus to related Brassicaceae through B. juncea

The impact of genetically modified canola (Brassica napus) on biodiversity has been examined since its initial stage of commercialization. Various research groups have extensively investigated crossability and introgression among species of Brassicaceae. B. rapa and B. juncea are ranked first and second as the recipients of cross-pollination and introgression from B. napus, respectively. Crossability between B. napus and B. rapa has been examined, specifically in terms of introgression from B. napus to B. rapa, which is mainly considered a weed in America and European countries. On the other hand, knowledge on introgression from B. napus to B. juncea is insufficient, although B. juncea is recognized as the main Brassicaceae weed species in Asia. It is therefore essential to gather information regarding the direct introgression of B. napus into B. juncea and indirect introgression of B. napus into other species of Brassicaceae through B. juncea to evaluate the influence of genetically modified canola on biodiversity. We review information on crossability and introgression between B. juncea and other related Brassicaseae in this report.     

Functional Growth Model for Pod Development in Greengram and Blackgram

Identification of readily measured traits that are associated with higher productivity of field crops is important to breeders. The objective of this study was to find out whether pod filling rate or duration of pod filling greatly decide final pod weight in greengram and blackgram. The effective pod filling duration is only 16 days in both the crops. Pod filling rate is more important than duration of pod development. Richards function is found to more closely fit the data on pod development from anthesis to maturity in both the species.     

Brassilexin accumulation and resistance to Leptosphaeria maculans in Brassica spp. and progeny of an interspecific cross B. juncea × B. napus

Summary Resistance to Leptosphaeria maculans was assessed in Brassica napus, B. juncea, B. carinata, B. nigra and progeny issuing from an interspecific cross B. napus × B. juncea, using a cotyledon-inoculation test. In these individual plants, brassilexin accumulation was determined following an abiotic, non-specific, elicitation. All the tested B. napus cultivars were highly susceptible to the parasite and weakly accumulated brassilexin. In contrast, B. juncea, B. carinata, and B. nigra usually displayed a hypersensitive response to the inoculation and accumulated more brassilexin than B. napus. The same correlation between resistance to L. maculans and phytoalexin accumulation was observed in the interspecific hybrid progeny. The cotyledon-inoculation test allowed the discrimination of plants displaying a hypersensitive response to the inoculation from those highly sensitive to the parasite, but intermediate disease severity classes were not usually representative of resistance or susceptibility. In this respect, brassilexin determination allowed differentiation, within a set of plants presenting an intermediate response to the pathogen, of plants with a high (B. juncea-like), and with a weak (B. napus-like) ability to accumulate brassilexin.Abbreviations IHP interspecific hybrid progeny - JR B. juncea-type complete resistance to blackleg (Roy, 1984) - W&D test cotyledon-inoculation test as described by Williams & Delwiche (1979)     

Geneticanalysis of drought stress response in rapeseed (Brassica campestris and B. Napus). II. Yield improvement and the application of selection indices

Summary The expected improvement in grain yield if selection was made for plant characters measured between flowering and maturity was evaluated in populations of rapeseed (Brassica campestris and B. napus) grown in a droughted environment. Drought was commenced at flowering in each species and measurements were made on plants at the commencement of the drought stress, during the stress treatment and at crop maturity.Substantial genetic and phenotypic variation was observed in yield as well as the different morphophysiological determinants of yield. In B. campestris no single parameter was found to be a suitable alternative selection criterion to yield since the correlated responses in yield if selection was for another character was lower than if selection was for yield alone. By the use of a selection index however, joint selection for yield, as well as harvest index, 1000 seed weight and seeds per pod, was expected to be 20% more effective than direct selection for yield under drought. In the B. napus population direct selection for flowering time or for harvest index was predicted to result in a genetic advance in yield equal to or greater than that obtained by direct selection for yield, whereas joint selection for flowering time and yield should result in a 16% greater yield increase. Selection for increased yield in these populations grown in droughted conditions is discussed.