Screening techniques and sources of resistance to parasitic angiosperms

Parasitic angiosperms cause great losses in many important crops under different climatic conditions and soil types. The most widespread and important parasitic angiosperms belong to the genera Orobanche, Striga, and Cuscuta. The most important economical hosts belong to the Poaceae, Asteraceae, Solanaceae, Cucurbitaceae, and Fabaceae. Although some resistant cultivars have been identified in several crops, great gaps exist in our knowledge of the parasites and the genetic basis of the resistance, as well as the availability of in vitro screening techniques. Screening techniques are based on reactions of the host root or foliage. In vitro or greenhouse screening methods based on the reaction of root and/or foliar tissues are usually superior to field screenings and can be used with many species. To utilize them in plant breeding, it is necessary to demonstrate a strong correlation between in vitro and field data. The correlation should be calculated for every environment in which selection is practiced. Using biochemical analysis as a screening technique has had limited success. The reason seems to be the complex host-parasite interactions which lead to germination, rhizotropism, infection, and growth of the parasite. Germination results from chemicals produced by the host. Resistance is only available in a small group of crops. Resistance has been found in cultivated, primitive and wild forms, depending on the specific host-parasite system. An additional problem is the existence of pathotypes in the parasites. Inheritance of host resistance is usually polygenic and its transfer is slow and tedious. Molecular techniques have yet to be used to locate resistance to parasitic angiosperms. While intensifying the search for genes that control resistance to specific parasitic angiosperms, the best strategy to screen for resistance is to improve the already existing in vitro or greenhouse screening techniques.     

Breeding for resistance to lentil Ascochyta blight

Ascochyta blight, caused by Ascochyta lentis, is one of the most globally important diseases of lentil. Breeding for host resistance has been suggested as an efficient means to control this disease. This paper summarizes existing studies of the characteristics and control of Ascochyta blight in lentil, genetics of resistance to Ascochyta blight and genetic variations among pathogen populations (isolates). Breeding methods for control of the disease are discussed. Six pathotypes of A. lentis have been reported. Many resistant cultivars/lines have been identified in both cultivated and wild lentil. Resistance to Ascochyta blight in lentil is mainly under the control of major genes, but minor genes also play a role. Current breeding programmes are based on crossing resistant and high‐yielding cultivars and multilocation testing. Gene pyramiding, exploring slow blighting and partial resistance, and using genes present in wild relatives will be the methods used in the future. Identification of more sources of resistance genes, good characterization of the host‐pathogen system, and identification of molecular markers tightly linked to resistance genes are suggested as the key areas for future study.     

Genes conferring low seedling reaction to Mexican pathotypes of Puccinia recondita f. sp. tritici,and adult-plant responses of recent wheat cultivars from the former USSR

Fifty-five spring bread wheat (Triticum aestivum L.) cultivars, mostly released between 1975 and 1991 in eight leaf rust-prone spring wheat growing regions of the former USSR, were tested in the seedling growth stage for reaction to 15 Mexican pathotypes of Puccinia recondita f. sp. tritici. In total, seven known and at least two unknown genes were identified, either singly or in combinations: Lr3 (7 cultivars), Lr10 (14), Lr13 (5), Lr14a (1), Lr16 (1), Lr23 (3); the unknown genes were identified in 14 cultivars. The first unknown gene could be either Lr9, Lr19, or Lr25; however, the second unknown gene in 9 cultivars was different from any named gene. Twelve of the 15 pathotypes are virulent for this gene, hence its use in breeding for resistance will be limited. The cultivars were also evaluated at two field locations in Mexico with two pathotypes in separate experiments. The area under the disease progress curve and the final disease rating of the cultivars indicated genetic diversity for genes conferring adult plant resistance. based on the symptoms of the leaf tip necrosis in adult plants, resistance gene Lr34 could be present in at least 20 cultivars. More than half of the cultivars carry high to moderate levels of adult plant resistance and were distributed in each region.     

Characterization of the anthracnose resistance gene present in Ouro Negro (Honduras 35) common bean cultivar

Ouro Negro (Honduras 35) is a highly productive Mesoamerican black seeded bean cultivar that possesses a major dominant gene conferring resistance to anthracnose (causal organism Colletotrichum lindemuthianum). In this work the anthracnose resistance gene present in Ouro Negro was characterized by studying allelic relationships to the following previously characterized anthracnose resistance genes (cultivars): Co-1 (MDRK), Co-1 ² (Kaboon), Co-1 ³ (Perry Marrow), Co-2 (Cornell 49-242), Co-3 (Mexico 222), Co-4 (TO), Co-4 ² (SEL 1308), Co-5 (SEL1360), Co-6 (AB 136), and the resistance genes present in PI 207262 and Widusa. In addition, we determined the resistance spectrum of Ouro Negro in relation to 19 pathotypes of C. lindemuthianum. The allelism tests confirmed that the dominant anthracnose resistance gene present in Ouro Negro is positioned at a locus distinct from those with which it was compared. We propose that this new gene be named Co-10. The inoculation of Ouro Negro with the 19 pathotypes of C. lindemuthianum demonstrated that Co-10 confers resistance to pathotypes 23, 64, 67, 73, 81, 83, 87, 89, 95, 102, 117, 119, 343, 453, 1033, 1545 and 1600. The identification of Co-10 is an important contribution to bean breeding programs that are in constant need of new sources of resistance to anthracnose. This revised version was published online in July 2006 with corrections to the Cover Date.     

Evaluation of seedling and adult plant resistance to leaf rust in European wheat cultivars

Summary A set of 105 European wheat cultivars, comprising 68 cultivars with known seedling resistance genes and 37 cultivars that had not been tested previously, was tested for resistance to selected Australian pathotypes of P. triticina in seedling greenhouse tests and adult plant field tests. Only 4% of the cultivars were susceptible at all growth stages. Twelve cultivars lacked detectable seedling resistance to leaf rust, and among the remaining cultivars, 10 designated genes were present either singly or in combination. Lr13 was the most frequently detected gene, present in 67 cultivars, followed by the rye-derived gene Lr26, present in 19 cultivars. Other genes present were Lr1, Lr3a, Lr3ka, Lr10, Lr14a, Lr17b, Lr20 and Lr37. There was evidence for unidentified seedling resistance in addition to known resistance genes in 11 cultivars. Field tests with known pathotypes of P. triticina demonstrated that 57% of the cultivars carried adult plant resistance (APR) to P. triticina. The genetic identity of the APR is largely unknown. Genetic studies on selected cultivars with unidentified seedling resistances as well as all of those identified to carry APR are required to determine the number and inheritance of the genes involved, to determine their relationships with previously designated rust resistance genes, and to assess their potential value in breeding for resistance to leaf rust.     

黄瓜霜霉病研究进展

为给黄瓜霜霉病抗病育种及病害防治提供系统、完善的理论依据,笔者对黄瓜霜霉病发生危害、病原菌生理特性、寄主范围、生理分化以及寄主抗性遗传、综合防治等多个方面国内外研究进展进行归纳总结。分析发现,尽管国内外研究者对病原菌生理分化、抗药性和寄主抗性基因鉴定等方面进行了大量富有成效的研究,但由于鉴定方法不统一及病原菌的高度变异,病原菌生理分化及寄主遗传规律等研究尚存争议,抗性基因功能验证的研究相对匮乏,因而在这些方面还有待进一步深入系统的研究。本研究同时也探讨了今后黄瓜霜霉病的研究方向。     

Breeding for resistance to cereal cyst nematode in wheat

Summary The progress of a backcross breeding programme to introduce resistance against the cereal cyst nematode into wheat is described. Methods of resistance screening and criteria for selection are detailed and the results discussed with reference to alternative procedures for the introduction of new resistance genes into major breeding programmes.     

Detection and inheritance of leaf rust resistance in common wheat lines Agra Local and IWP94

Genetic studies were undertaken to determine the number and identities of leaf rust resistance genes in common wheat lines Agra Local and IWP94. The infection type arrays of the two lines with eight pathotypes (pt.) of P. triticina were different from those of lines possessing known leaf rust resistance (Lr) genes. Agra Local possessed two recessive resistance genes, one conditioning resistance to pathotype 4R9-7, and the other, a temperature-sensitive factor, gave resistance to pt. 121R127 at high temperature (27°C). IWP94 was previously demonstrated to carry Lr23. From the present study IWP94 was determined to have at least four leaf rust resistance genes. The first of these was the same recessive gene conferring resistance to pathotype 4R9-7 which was found in Agra Local. A second partially dominant gene conferred resistance to pathotype 121R127 at high temperature and two additional recessive genes governed resistance to pathotype 93R15. When present together, these two recessive genes complemented each other and provided resistance to pathotype 69R13 as well. One of the two recessive genes conferring resistance to pathotypes 93R15 and 69R13 was Lr23.     

DNA marker-assisted evaluation of potato genotypes for potential resistance to potato cyst nematode pathotypes not yet invading into Japan

One of major objectives of crop breeding is conferring resistance to diseases and pests. However, large-scale phenotypic evaluation for many diseases and pests is difficult because strict controls are required to prevent their spread. Detection of disease resistance genes by using DNA markers may be an alternative approach to select potentially resistant accessions. Potato (Solanum tuberosum L.) breeders in Japan extensively use resistance gene H1, which confers nearly absolute resistance to potato cyst nematode (Globodera rostochiensis) pathotype Ro1, the only pathotype found in Japan. However, considering the possibility of accidental introduction of the other pathotypes, breeding of resistant varieties is an important strategy to prevent infestation by non-invading pathotypes in Japan. In this study, to evaluate the prevalence of resistance genes in Japanese genetic resources, we developed a multiplex PCR method that simultaneously detects 3 resistance genes, H1, Gpa2 and Gro1-4. We revealed that many Japanese varieties possess not only H1 but Gpa2, which are potentially resistant to other pathotypes of potato cyst nematode. On the other hand, no genotype was found to have the Gro1-4, indicating importance of introduction of varieties having Gro1-4. Our results demonstrate the applicability of DNA-marker assisted evaluation of resistant potato genotypes without phenotypic evaluation.     

Past,present and future opportunities in breeding for disease resistance,with examples from wheat

Summary This introductory chapter contains some general comments about plant breeding and breeding for disease resistance. The use of disease resistant crop plants is an environmentally favourable method of controlling disease but the process of breeding for disease resistance is subject to several constraints. Among them is the variability of pathogens in relation to host resistance. Some parts of this variation can be resolved into gene-for-gene interactions, but the boundaries within which such interactions can be detected are not sharp. The discussion of this variation is illustrated by reference to some important diseases of wheat, especially yellow rust, septoria and eyespot. The objective of obtaining durable resistance is discussed and some contributions of new genetical and molecular techniques to breeding for resistance are considered. It is suggested that new technology will enhance breeding for disease resistance but that established techniques of plant breeding will remain relevant and important.     

Elucidation of virus-host interactions to enhance resistance breeding for control of virus diseases in potato

Potato virus Y (PVY) and Potato mop-top virus (PMTV) are viruses whose geographical distribution is expanding and economic losses are increasing, in contrast to most of other viruses infecting potato crops. Most potato cultivars lack broad-spectrum resistance to the new, genetically complex strains of PVY, and no efficient resistance to PMTV is known in potato. Control of the vectors of these viruses is not an efficient or possible strategy to prevent infections. Studies on molecular virus-host interactions can discover plant genes that are important to viral infection or antiviral defence. Both types of genes may be utilized in resistance breeding, which is discussed in this paper. The advanced gene technologies provide means to fortify potato cultivars with effective virus resistance genes or mutated, non-functional host factors that interfere with virus infection.     

Characterization of resistance genes against scald (Rhynchosporium secalis (Oudem.) J.J. Davis) in barley (Hordeum vulgare L.) lines from central Norway, by means of genetic markers and pathotype tests

Rhynchosporium secalis is a serious pathogen of barley (Hordeum vulgare L.) in central Norway. A breeding effort was initiated in 1977 to introduce resistance from different sources into adapted genotypes, and the first cultivar from the program was recently released. However, little is known about the resistance genes introgressed in this cultivar or in advanced breeding lines. An effort was made to address this issue through a set of isolates and available molecular markers. Fourteen breeding lines and their resistance donors were investigated by evaluating their reactions to 11 R. secalis isolates. Bulked segregant analysis was used to identify molecular markers linked to resistance genes in 12 of the breeding lines. The isolates were found to be of less discriminating value than the markers. Useful information has been obtained as to the nature of several of the resistance genes introgressed. Eight of the 12 breeding lines contained introgressed genes that were located at the `complex Rh' locus on chromosome 3H and hence may not easily be pyramided into the same genotype. Previous information about the nature of the resistance in `Jet' is questioned. Neither of the resistance genes Rh or Rh2 seems to have been incorporated into Norwegian breeding material.     

Resistance in Cucumis sativus L. to Tetranychus urticae Koch. 1. The role of plant breeding in integrated control

Summary The role of plant breeding and particularly of host plant resistance in integrated control is discussed. Host plant resistance to insects and mites, especially to Tetranychus urticae is reviewed. A standard terminology for disease and pest resistance is recommended.     

Partial resistance to a Fusarium root disease in Egyptian white lupin landraces

A Fusarium sp. root pathogen of lupin is the causal agent of the most important disease that affects the cultivation of white lupin (Lupinus albus L.) in Egypt. The aim of our research was to investigate whether host resistance to Fusariumroot disease was available in Egyptian landraces ofLupinus albus. Five Fusarium isolates collected from white lupin samples in Egypt were tested with an Egyptian landrace and a French cultivar. The most aggressive isolate was used to screen an additional 15 Egyptian landraces, two cultivars released in Egypt after selection among landraces, one Polish cultivar, and two French cultivars. The assessment of host response to Fusarium was performed in a field, and under controlled conditions in a greenhouse pot experiment. Most landraces and the two Egyptian cultivars showed better resistance with good accordance between field and pot experiment. This experiment showed that Egyptian genetic resources of white lupin possess partial resistance to Fusarium root rot. Egyptian germplasm may be an alternative genetic source for incorporating partial resistance to Fusarium root rot into the breeding pools. This revised version was published online in July 2006 with corrections to the Cover Date.     

Pathotype-specificity in potato cyst nematodes,a reconsideration

Summary Within the two potato cyst nematode (PCN) species, Globodera rostochiensis and G. pallida, eight pathotypes are presently discerned by seven differential Solanum clones, whereby resistance is defined as a Pf/Pi ratio <1.0, and susceptibility as a ratio above 1.0. This pathotyping is felt to be unsatisfactory. Reconsidering the original data and data published by others resulted in the conclusion that there are only three reliably recognizable Ro-pathotypes Ro1/Ro4, Ro2/Ro3 and Ro5. The differential clones 60.21.19 and 65.346.19 differentiate only between the two species. Within G. pallida it does not seem possible to identify pathotypes reliably. The resistance-virulence pattern is of a quantitative nature fitting a polygenic system. There appear to be pathotype-specific effects, which are not large enough to identify and classify pathotypes reliably. It is therefore suggested to rename the Ro-pathotypes into Ro1 (old Ro1 and Ro4), Ro3 (old Ro2 and Ro3) and Ro5 (old Ro5) and to speak of more and less virulent Pa-populations.The rigorous requirements for resistance in various countries present great problems to the breeders and result in the removal of much valuable partial resistance. Especially with G. pallida these rigorous requirements are probably an obstacle rather than an encouragement to solve the pallida problem.     

Transgenic potato plants resistant to viruses

Summary Traditional potato breeding is a laborious process in which, by intercrossing, valuable traits from different parental clones are combined in a progeny genotype. Depending on the availability of genes, molecular techniques can be used to add specific genes to existing cultivars that, although otherwise satisfactory, lack a lew commercially important traits. For virus resistance the gene for the coat protein of a given virus transplanted into the genome of the plant renders the plant resistant to that virus. In conferring such resistance to potato varieties it proved to be possible to preserve their intrinsic properties.     

Comparison of the resistance and susceptibility to downy mildew [<Emphasis Type="Italic">Hyaloperonospora parasitica</Emphasis>, constant. (Pers. Ex Fr)] of nine <Emphasis Type="Italic">Brassica olearacea</Emphasis> accessions in laboratory,seedbed, and field screens

The relationship between resistance in seedlings, young and adult plants is studied for the pathosystem Brassica oleracea–Hyaloperonospora parasitica. Genotypes identified in the laboratory as resistant or susceptible or exhibiting a differential reaction to a selection of H. parasitica isolates were tested in 1997–1999 in seedbeds and fields under natural infestation. Isolates tested in the laboratory were grouped in five pathotypes, of which four were presented by isolates from Brittany, France. Genotypes susceptible to all pathotypes in the laboratory were also susceptible in the seedbed and field tests, while genotypes expressing a differential response to pathotypes were either resistant or susceptible. Accessions Everest, DEGC, ESPG and RS1105 exhibiting resistance to all pathotypes except I, were resistant in all environments and remained resistant in 2000–2002. Pathotype I was not prevailing in the field and results support the hypothesis that accessions resistant under laboratory conditions will be resistant under field conditions, provided the same pathotypes are present under the laboratory and field conditions.     

Novel pathotypes of lettuce mosaic virus — breakdown of a durable resistance?

Summary Resistance to lettuce mosaic virus (LMV) is derived either from cv. Gallega (g gene) or the wild accession PI251245 (mo gene). Previous studies indicated that these two genes were identical. Breeders in Europe produced numerous resistant cultivars utilisingg while in the USAmo was used. The resistance has been effective for over 20 years. However, recently there have been reports of LMV isolates causing unusually severe and sometimes necrotic symptoms on cultivars with these resistance genes. Investigations of these severe isolates have distinguished three new pathotypes in addition to the common pathotype (II) and identified a novel dominant gene for resistance. Themo/g genes confer resistance to pathotypes I and II but pathotype III possesses virulence for cultivars withg but not for those withmo. These two genes are therefore not identical but are probably either closely linked genes or alleles. Pathotype IV possesses virulence for all lettuce lines so far tested. Some isolates of this pathotype are seed transmitted in cultivars possessingmo org and have caused severe crop losses in southern France. The durability of the resistance conditioned by these two genes is discussed.     

Analysis of durable resistance to stem rust in barley

Summary Since the mid-1940's, barley cultivars grown in the northern Great Plains of the USA and Canada have been resistant to stem rust caused byPuccinia graminis f. sp.tritici. This durable resistance is largely conferred by a single gene,Rpg1, derived from a single plant selection of the cultivar Wisconsin 37 and an unimproved Swiss cultivar. At the seedling stage, barley genotypes withRpg1 generally exhibit low mesothetic reactions at 16–20° C and slightly higher mesothetic reactions at 24–28° C to many stem rust pathotypes. This resistance is manifested by a low level of rust infection and mostly incompatible type uredia on adult plants.Rpg1 reacts in a pathotype-specific manner since some genotypes ofP. g. f. sp.tritici are virulent on cultivars carrying this gene in the field. Several factors may have contributed to the longevity of stem rust resistance in barley, a) since barley is planted early and matures early, it can sometimes escape damage from stem rust inoculum carried from the south; b) one or more minor genes may augment the level of resistance already provided byRpg1; c) the cultivation of resistant wheat cultivars and eradication of barberry have reduced the effective population size and number of potential new pathotypes ofP. g. f. sp.tritici, respectively; and d) virulent pathotypes ofP. g. f. sp.tritici andP. g. f. sp.secalis have not become established. This situation changed in 1989 when a virulent pathotype (Pgt-QCC) ofP. g. f. sp.tritici became widely distributed over the Great Plains. However,Rpg1 may still confer some degree of resistance to pathotype QCC because stem rust severities have been low to moderate and yield losses light on barley cultivars carrying the gene during the last four seasons (1989–1992). Several sources of incomplete resistance to pathotype QCC have been identified in barley. To facilitate the transfer of resistance genes from these sources into advanced breeding lines, molecular marker assisted selection is being employed.     

Small differential interactions for partial resistance in rice cultivars to virulent isolates of the blast pathogen

Summary Six rice genotypes, differing in partial resistance, were exposed to three isolates of the blast pathogen. Of the variance due to host and pathogen genotypes, 39% was due to host genotype effects, 60% was due to isolate effects, and only 1% was due to host genotype × isolate interactions. Although small, this interaction variance was highly significant and mainly due to the IR50 × W6-1 and IR37704 × JMB8401-1 combinations. Although behaving largely as race-non-specific (large main effects only), the partial resistance cannot be classified as race-non-specific. The results suggest that minor genes for partial resistance operate in a gene for gene relationship with minor genes in the pathogen.