Current status and future strategy in breeding chickpea for resistance to biotic and abiotic stresses

Chickpea (Cicer arietinum L.) production has remained static for the past two decades. One major limiting factor has been susceptibility of cultivars to several biotic and abiotic stresses that adversely affect yield. In recent years, cultivars resistant to Ascochyta blight (Ascochyta rabiei [Pass.] Lab.), Fusarium wilt (Fusarium oxysporum f. sp. ciceris), and cold have been bred and released in many countries. Some progress has been made in breeding for resistance to drought, insects, and cyst nematode, but not for viruses, heat, and salinity. Two or more stresses are of equal importance in most chickpea growing areas. Therefore, future efforts should be directed toward the development of cultivars with multiple-stress resistance. Proper understanding of important stresses in different countries and the genetics of resistance should lead to more systematic approaches to resistance breeding. Wild Cicer species hold promise and deserve attention in resistance breeding.     

Genetic variation for disease and nematode resistances and forage quality in perennial diploid and tetraploid lucerne populations (Medicago sativa L.)

Summary Twenty-five lucerne populations of the Medicago sativa complex, which were either diploid or tetraploid and wild or cultivated, were analysed for their resistance to four different fungal diseases and to stem nematode. Forage quality, including stem digestibility and saponin content, was also tested.Populations varied in susceptibility to the diseases caused by Colletotrichum trifolii, Verticillium albo-atrum, Sclerotinia trifoliorum and Pseudopezizza medicaginis, and to the nematode Ditylenchus dipsaci. Except for Sclerotinia rot, sativa and falcata subspecies differed in susceptibility, but this grouping of populations did not account for the full range of variation among them. However, the resistance to P. medicaginis was much lower in the sativa than in the falcata populations.Populations also varied significantly in stem fiber content and digestibility. Stem digestibility was negatively correlated to forage yield. Wild sativa and falcata populations had lower fiber content and higher digestibility than cultivated sativa populations. The medicagenic acid was the sapogenin responsible for the anti-nutritional effect of the lucerne measured by the yellow mealworm larvae Tenebrio molitor. The medicagenic acid content was lowest for the pure sativa populations, highest for the pure falcata populations, and intermediate for the French sativa varieties that have some traits originating from falcata germplasm. Some populations could be used in breeding programs to improve disease and nematode resistance, and forage quality.     

Comparison of sources of resistance to leaf aphids in lettuce (Lactuca sativa L.)

Summary Three lines of lettuce with resistance to Nasonovia ribisnigri, based on the dominant Nr-gene, and four lines selected for partial resistance to Myzus persicae were tested against three species of leaf aphid: N. ribisnigri, M. persicae and Macrosiphum euphorbiae. The effect of the Nr-gene was also studied in a segregating F2 population.In the material tested, resistance to N. ribisnigri was exclusively based on the Nr-gene, lines selected only for resistance to M. persicae showed no resistance to N. ribisnigri. The Nr-gene also induces partial resistance to M. persicae, but the level of this resistance is influenced by other genes, because the lines with Nr-gene differed significantly from each other for reproduction of M. persicae. The Nr-gene had no effect on the resistance of lettuce to M. euphorbiae.In lines with the Nr-gene, levels of resistance to M. persicae and to M. euphorbiae were correlated, suggesting that the resistance may be determined by the same genes. The Nr-line with highest resistance to M. persicae was comparable for this characteristic to the lines selected for resistance to M. persicae.The cultivars Taiwan and Ravel possess a resistance factor to M. euphorbiae that has no effect on M. persicae or N. ribisnigri. Lines selected for resistance to M. persicae also showed partial resistance to M. euphorbiae. Based on the present results no conclusions can be drawn whether this resistance is based on the same genes that provide resistance to M. persicae, or on a resistance factor comparable to that found in Taiwan and Ravel.     

Evaluation of a DNA probe for the quantitative detection of common bacterial blight in common bean and its application in a breeding program

Summary Breeding of Phaseolus vulgaris L. for resistance to common bacterial blight (CBB) can be done with visual evaluations of symptoms to distinguish broad resistance classes, but a more quantitative measure was needed for genetic studies of resistance. A novel method of evaluation was developed by quantifying Xanthomonas campestris pv. phaseoli (XCP) in bean leaf tissue infected with CBB using a ³²P-labeled probe and densitometric analysis of hybridization signals. Quantification of bacterial populations using the probe was highly correlated (r=0.98) with the number of colony forming units (CFU) from plate counts of the same leaf samples. The probe was used to follow XCP population dynamics on susceptible (BAT 41) and resistant (OAC 88-1) bean genotypes. OAC 88-1 supported a maximum XCP population which was approximately tenfold less than BAT 41. The probe was also used to study an F₂/F₃ population segregating for resistance. Narrow sense heritability estimates were less for resistance measured on the basis of bacterial populations (0.18–0.26) than on visual scores of symptoms (0.29–0.38). The anticipated response to selection for CBB resistance would be less based on bacterial numbers than based on symptom expression in this population. In breeding for resistance to CBB, selection based on visual symptoms combined with measurements of XCP populations using a DNA probe can be used to develop bean genotypes that are both resistant to symptom development and bacterial multiplication.Abbreviations CBB common bacterial blight - CFU colony forming units - XCP Xanthomonas campestris pv. phaseoli     

Sources of resistance to diseases of sugar beet in related Beta germplasm: II. Soil-borne diseases

Between 580 and 700 accessions of related cultivated and wild species of the genus Beta were assessed for resistance to four soil-borne diseases of sugar beet: two seedling damping-off diseases caused by the fungi Aphanomyces cochlioides and Pythium ultimum and two diseases of more mature plants, Rhizoctonia root and crown rot, caused by the fungus R. solani, and Rhizomania, caused by Beet necrotic yellow vein virus (BNYVV), a furovirus transmitted by the plasmodiophorid Polymyxa betae. Analysis of resistance data (assessed on an international standardised 1–9 scale of Resistance Scores) indicated that the highest levels of resistance ({RS} 2) to A. cochlioides and P. ultimum were to be found amongst accessions of the more distantly related sections Corollinae (93% of accessions tested) and Procumbentes (10%), respectively; although useful levels could also be found in the more closely related, and sexually compatible, section Beta (1–6%). Resistance to Rhizoctonia was also found in section Beta (5–7%), depending on whether field or glasshouse tests were used, but there was little evidence of generally high levels of resistance to Rhizomania among accessions of this section. None of the accessions of sections Corollinae and Procumbentes exhibited any notable resistance to Rhizoctonia. However, all sections Procumbentes and some sections Corollinae (4%) accessions were highly resistant to Rhizomania. Individuals with high levels of resistance to Rhizomania were identified from within some section Beta and Corollinae accessions, in which there was evidence of segregation.     

Reaction of tritordeum to Fusarium culmorum and Septoria nodorum

Summary Hordeum chilense is a wild barley extensively used in wide crosses in the Triticeae. It could be a valuable source of resistance to Fusarium culmorum and Septoria nodorum. Some H. chilense x Triticum spp. amphiploids, named tritordeums, were more resistant than the parental wheat line to these diseases, others were not. Average contents of ergosterol and deoxynivalenol (DON) suggested that resistance to colonization by Fusarium was the highest for Hordeum chilense, followed by tritordeum and wheat in decreasing order. In particular, the H. chilense genotypes H7 and H17 enhanced the wheat resistance to F. culmorum in its tritordeum offsprings. Resistance to S. nodorum in tritordeum was not associated with tall plant height. There is sufficient genetic variation for resistance to F. culmorum and S. nodorum among tritordeum to allow the breeding of lines combining short straw and resistance to both diseases.     

Assessment of Gossypium sturtianum and G. australe as potential sources of Fusarium wilt resistance to cotton

When challenged with Fusarium oxysporum f. sp. vasinfectum (Fov) from vegetative compatibility groups (VCGs) 01111 and 01112 in glasshouse tests, Gossypium australe Mueller and Gossypium sturtianum Willis accessions showed a variety of disease responses ranging from highly resistant to highly susceptible. Under high disease pressure G. sturtianum accession Gos-5275 was significantly more resistant than the commercial G. hirsutum cultivars that are designated standards for Fusarium resistance by Australian cotton breeders. Under low disease pressure G. sturtianum accession Gos-5250 was more susceptible than a highly susceptible commercial cultivar. A series of glasshouse tests was performed at two locations (Indooroopilly, QLD. and Canberra, ACT), and under low and high disease pressure. In these tests, a hexaploid cross (Gos-5271) generated from a Fusarium-resistant G. sturtianum (Gos-5275) and a Fusarium-susceptible G. hirsutum L. (CPI-138969) was significantly more resistant to Fusarium wilt than its G. hirsutum parent. Thus G. sturtianum, with a diploid genome and a range of responses to Fov challenge, has the potential to provide the basis for the elucidation of the genetic basis of resistance to Fusarium wilt in cotton species. In addition, resistant accessions of G. sturtianum are identified as a potential source of Fusarium wilt resistance genes for cotton breeding. In the glasshouse tests used to assess the resistance of various Gossypium accessions to Fusarium wilt disease, the scoring of vascular browning was found to give a more reliable indication of disease severity than the scoring of foliar symptoms. This revised version was published online in August 2006 with corrections to the Cover Date.     

Assessment of drought resistance in 20 accessions of Triticum aestivum and related species I. Total dry matter and grain yield stability

Summary Interspecific hybridisation has been suggested to improve the resistance of crops to biotic and abiotic stresses. To identify sources of resistance to drought which could be used in a breeding programme, an assessment of a large number of wheat species was carried out. The evaluation was based on the estimation of a drought susceptibility index and stability in total dry matter and grain yields of various species grown across a range of soil moisture conditions. On the basis of stability in total dry matter yield, T. aestivum was relatively more drought resistant. However, on the basis of grain yield stability, T. sphaerococcum, T. vavilovii and T. aestivum cv. C 306 exhibited better yield stability and drought resistance than the other species tested. It is suggested that the above species could be utilized as parental material in a hybridisation programme aimed at improving the drought resistance of wheat.     

Sources of resistance to Pyrenophora tritici-repentis in bread wheats

Summary No complete resistance to Pyrenophora tritici-repentis has been located in more than 1400 bread wheats examined. Incomplete resistance, however, occurs at different levels in many spring and winter types and data are presented for the strongest sources of resistance detected. In particular, there is a high frequency of Brazilian spring wheats with appreciable levels of resistance to this pathogen. Recent international nurseries from CIMMYT, Mexico, also contain numerous potentially valuable sources of resistance and these wheats may be shorter and higher yielding in Australia than the Brazilian wheats. The resistances in many Brazilian cultivars may be largely common because the cultivars are often strongly interrelated. Some of the Brazilian wheats resistant to P. tritici-repentis are also resistant to one or more of the septoria diseases and/or possess tolerance to aluminium toxicity.     

Assessment of genetic diversity in synthetic hexaploid wheats and their Triticum dicoccum and Aegilops tauschii parents using AFLPs and agronomic traits

Synthetic hexaploid wheats are of interest to wheat breeding programs, especially for introducing new genes that confer resistance to biotic and abiotic stresses. A group of 54 synthetic hexaploid wheats derived from crosses between emmer wheat(Triticum dicoccum, source of the A and B genomes) and goat grass (Aegilops tauschii, D genome donor) were investigated for genetic diversity. Using the AFLP technique, dendrograms revealed clear grouping according to geographical origin for the T. dicoccum parents but no clear groups for the Ae. tauschii parents. The geographical clustering of the T. dicoccum parents was also reflected in the dendrogram of their derived synthetic hexaploids. Diversity of the T. dicoccum parents and their derived synthetic hexaploids was further evaluated by measuring 18morphological and agronomic traits on the plants. Clustering based on morphological and agronomic data also reflected geographical origin. However, comparison of genetic distances obtained from AFLP and agronomic data showed no correlation between the two diversity measurements. Nevertheless, similarities among major clusters with the two systems could be identified. Based on percentage of polymorphic markers, the synthetic hexaploids had a considerably higher level of AFLP diversity (39%) than normally observed in cultivated hexaploid wheat (12–21%). This suggests that synthetic hexaploid wheats can be used to introduce new genetic diversity into the bread wheat gene pool. This revised version was published online in July 2006 with corrections to the Cover Date.     

Chromosome location of resistance to septoria leaf blotch and common bunt in wheat-barley addition lines

Septoria leaf blotch and common bunt are important diseases of wheat to which Hordeum vulgare is resistant. Addition lines of H. vulgare in wheat were utilized to determine which H. vulgare chromosomes carry resistance genes. Resistance to septoria leaf blotch was conferred by gene(s) present all over the barley genome, but more strongly by those located on chromosomes 7 and 4. Almost complete resistance to common bunt was conferred by gene(s) present in chromosomes 6 and a slight but significant level of resistance was conferred by chromosome 7. This revised version was published online in August 2006 with corrections to the Cover Date.     

Screening relatives of wheat for snow mold resistance and freeing tolerance

Four hundred and eighty one accessions of species and subspecies of genera Triticum and Aegilops were evaluated for resistance to speckled snow mold caused by Typhula ishikariensis and for freezing tolerance. All Triticum and seven of Aegilops species were severely affected by snow mold. Among surviving Aegilops spp., only the Ae. cylindrica accessions exhibited resistance similar to that of the most resistant winter wheat cultivar, PI 173438. After repeated screening of accessions of Ae. cylindrica, 12 accessions were identified as having resistance similar to that of PI 173438; eleven of those were considered more freezing tolerant than PI 173438, but less than wheat cv. Valjevskaya, the freezing tolerant check. Accessions of Ae. cylindrica with snow mold resistance exceeding that of PI 173438, and with freezing tolerance, are currently being screened. This revised version was published online in August 2006 with corrections to the Cover Date.     

Resistance of cowpea and cereals to the parasitic angiospermStriga

Summary Striga species are parasitic angiosperms that attack many crops grown by subsistence farmers in sub-Saharan Africa and India. Control of the parasite is difficult and genetically resistant crops are the most feasible and appropriate solution. In cowpea, complete resistance toStriga gesnerioides has been identified. Breeding for resistance in sorghum has identified varieties with good resistance toS. asiatica in Africa and India. One variety was also resistant toS. hermonthica in W. Africa. No such resistance toStriga has been found in maize or millets.Resistant varieties have usually been sought by screening germplasm in fields naturally infested withStriga. However, laboratory techniques have also been developed, including anin vitro growth system used to screen cowpeas for resistance toS. gesnerioides. Two new sources of resistance in cowpea have been identified using the system. The technique has also been used to investigate the mechanisms of resistance in this crop. Two mechanisms have been characterised, both were expressed after penetration of cowpea roots by the parasite.The resistance of some sorghum varieties toStriga is controlled by recessive genes. In cowpea, resistance toStriga is controlled by single dominant genes. The genes for resistance are currently being transferred to cowpea varieties which are high yielding or adapted to local agronomic conditions. OneStriga resistant cowpea variety, Suvita-2, is already being grown widely by farmers in Mali. Reports of breakdown of resistance in cowpea toStriga have not yet been confirmed, but a wider genetic base to the resistance is essential to ensure durability ofStriga resistance.Abbreviations ICRISAT International Crops Research Institute for the Semi-Arid Tropics - IITA International Institute of Tropical Agriculture - LARS Long Ashton Research Station - SAFGRAD Semi-Arid Food Grain Research and Development     

The breeding of the cultivated potato species solanum x juzepczukii and S. x curtilobum II. The resynthesis of S. x juzepczukii and S. x curtilobum

Summary The possibility of combining anew the genomes of wild and cultivated progenitors of triploid S. x juzepczukii and pentaploid S. x curtilobum by following the known evolutionary pathway of these species was investigated.The resynthesis of S. x juzepczukii was easy, and a wide range of synthetic forms was bred. Among these were forms with higher frost resistance (-5°C) than has been found in natural S. x juzepczukii. The total tuber glycoalkaloid content of several synthetic hybrids was lower than or as low as that of natural clones. Most synthetic hybrids were more vigorous than natural S. x juzepczukii and produced about the same types of tubers as are found in the natural range of variation. The best diploid parents were found in the species S. goniocalyx.The attempt to resynthesize pentaploid S. x curtilobum has not been successful but tetraploid plants were obtained in the process. An explanation for the occurrence of tetraploids resulting from triploid x tetraploid and/or diploid crosses is offered.The newly bred tetraploids contain at least one genome from S. acaule (possibly two) and hybridize easily with ssp. andigena. They thus provide a means for the transfer of S. acaule germ plasm into the tetraploid cultivated gene pool which would profit from the frost resistance of S. acaule.     

Screening techniques and sources of resistance against parasitic weeds in grain legumes

Summary A number of parasitic plants have become weeds, posing severe constraints to major crops including grain legumes. Breeding for resistance is acknowledged as the major component of an integrated control strategy. However, resistance against most parasitic weeds is difficult to access, scarce, of complex nature and of low heritability, making breeding for resistance a difficult task. As an exception, resistance against Striga gesnerioides based on a single gene has been identified in cowpea and widely exploited in breeding. In other crops, only moderate to low levels of incomplete resistance of complex inheritance against Orobanche species has been identified. This has made selection more difficult and has slowed down the breeding process, but the quantitative resistance resulting from tedious selection procedures has resulted in the release of cultivars with useful levels of incomplete resistance. Resistance is a multicomponent event, being the result of a battery of escape factors or resistance mechanisms acting at different levels of the infection process. Understanding these will help to detect existing genetic diversity for mechanisms that hamper infection. The combination of different resistance mechanisms into a single cultivar will provide durable resistance in the field. This can be facilitated by the use of in vitro screening methods that allow highly heritable resistance components to be identified, together with adoption of marker-assisted selection techniques.     

Resistance to cane spot (Elsinoë veneta) in the red raspberry and its relationship to resistance to yellow rust (Phragmidium rubi-idaei)

Summary Varying degrees of resistance to cane spot were recorded among red raspberries in two cultivar trials and in 15 segregating families. The inheritance of resistance was studied, and the effect of gene H which determines the presence of cane hairs was assessed in eight of the families. The 0 to 5 scale used to record disease incidence was found to approximate to a logarithmic scale for the range of 0 to 93 cane spots per cane. Gene H and h phenotypes averaged scores of 3.05 and 2.42 respectively, but other genes independent of these had more influence on resistance. The latter genes appeared to be largely dominant. The possibility is discussed that one of them was a major gene with a large effect, but the evidence was equivocal in the absence of discontinuity in the expression of resistance. Resistance to yellow rust was studied in five of the families and was highly correlated with resistance to cane spot. Gene H had more influence on this disease, the mean counts of telia per unit leaf area for H and h segregates being 17.1 and 4.1 respectively.     

Yield and fruit quality losses caused by ToMV in pepino (Solanum muricatum L.) and search for sources of resistance

Tomato mosaic virus (ToMV) is a limiting factor for the success of pepino (Solarium muricatum) as a new crop. The effects of ToMV infection on total and marketable yield, fruit weight, length/width ratio and soluble solids content (SSC) have been studied in two commercial clones (`Sweet Long' and `Sweet Round'). ToMV infection depressed total yield in infected plants of `Sweet Long' (43.1%), while it had no effect on this trait in `Sweet Round' . Marketable yield was dramatically reduced by ToMV infection in both clones, 94% in `Sweet Long' and 100% in `Sweet Round'. Infected plants of clone `Sweet Long' had a lower weight than healthy plants. Although no differences in fruit weight were detected in `Sweet Round' between ToMV infected and healthy plants, many fruits from infected plants showed deformities. Changes in fruit length/width ratio and SSC as a result of ToMV infection were not relevant, but fruit quality was lower in infected fruits, most of which had corky-like flesh. Forty-two clones from cultivated (S. muricatum), wild (S. caripense and S. tabanoense) and interspecific hybrids were tested for ToMV resistance. All but seven clones (four from S. muricatum and three from interspecific hybrids S. muricatum × S. caripense) were susceptible. Non-susceptible clones showed variable degrees of resistance and developed hypersensitive local lesions. Among these clones the most promising as sources of variation for resistance to ToMV are those belonging to the cultivated species. Although no immunity was found, plants from these clones remained asymptomatic and absorbance values resulting from the DAS-ELISA tests in these plants were always lower than those of the susceptible control (cv. `Sweet Round'). These sources of resistance may be of great utility in developing commercial clones resistant to this severe disease affecting pepino. This revised version was published online in July 2006 with corrections to the Cover Date.     

Resistance to stripe rust in Triticum turgidum,T. tauschii and their synthetic hexaploids

Resistance to stripe rust (caused by Puccinia striiformis Westend.) of 34 Triticum turgidum L. var.durum, 278 T. tauschii, and 267 synthetic hexaploid wheats (T. turgidum x T. tauschii) was evaluated at the seedling stage in the greenhouse and at the adult-plant stage at two field locations. Mexican pathotype 14E14 was used in all studies. Seedling resistance, expressed as low infection type, was present in all three species. One hundred and twenty-eight (46%) accessions of T. tauschii, 8 (23%) of T. turgidum and 31 (12%) of synthetic hexaploid wheats were highly resistant as seedlings. In the field tests, resistance was evaluated by estimating area under disease progress curve (AUDPC). Synthetic hexaploid wheats showed a wide range of variability for disease responses in both greenhouse and field tests, indicating the presence of a number of genes for resistance. In general, genotypes with seedling resistance were also found to be resistant as adult plants. Genotypes, which were susceptible or intermediate as seedlings but resistant as adult plants, were present in both T. turgidum and the synthetic hexaploids. Resistances from either T. turgidum or T. tauschii or both were identified in the synthetic hexaploids in this study. These new sources of resistance could be incorporated into cultivated hexaploid wheats to increase the existing gene pool of resistance to stripe rust.     

Assessment of quantitative resistance of rice cultivars to Xanthomonas campestris pv. Oryzae: a comparison of inoculation methods

Summary Lesion size and lesion number were measured on cultivars of rice inoculated by clipping or spraying with virulent isolates of Xanthomonas campestris pv. oryzae. Mean percentage diseased leaf area (%DLA) gave a similar ranking for the two inoculation methods but differences in lesion size among cultivars were much more evident after clip than after spray inoculation. Correlation between the methods was high (r=0.82^**), but some cultivars responded differently with the two techniques. Cultivars which had low scores following spray inoculation showed low disease progress during the first nine weeks after transplanting into a screen-house experiment. Assessment after clip inoculation measures resistance due to spread of bacteria within the leaf xylem system, an important component of quantitative resistance. Assessment after spray inoculation measures all resistance, including resistance to entrance of bacteria into the leaf. In order to select rice entries with improved quantitative resistance to X. c. oryzae based on both components, a screening based on lesion length after clip inoculation, followed by a test for lesion number after spray inoculation, is advised.     

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)