The location and characteristics of novel sources of resistance to Bremia Lactucae Regel (downy mildew) in wild Lactuca L. Species

Summary Twenty-one accessions of 3 wild Lactuca species which could be hybridised with L. sativa, the cultivated lettuce, were inoculated at different stages of plant development with 3 multivirulent isolates of Bremia lactucae. Nineteen sources of resistance to B. lactucae, not attributable to the previously recognised resistance factors 1–11 were identified. Two lines of L. serriola showed similar resistance patterns as lines carrying R11. The resistance of some accession was incomplete particularly at the seedling stage and this phenomenon may be race specific.Tests on segregating F₂ populations of crosses between 2 different L. serriola accessions and L. sativa cultivars showed that the resistance in one line (LSE/18) appears to be inherited as a single dominant gene, which is sometimes incomplete in expression and allelic to either Dm6 or R7. The segregation patterns for resistance in PI 281876 did not give readily interpretable ratios.To assess the frequency of occurrence in B. lactucae populations of virulence factors to overcome this novel resistance, 11 of the novel sources of resistance were inoculated with numerous collections of the pathogen from the UK, Czechoslovakia and elsewhere and found to show a high level of resistance.     

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.     

New sources of major gene resistance in Lactuca to Bremia lactucae

Summary A total of 1789 accessions of several lettuce collections was screened to find new major gene resistance to the downy mildew fungus Bremia lactucae Regel. The accessions belonged to the species Lactuca sativa (N=1288), L. serriola (N=399), L. saligna (N=52) and L. virosa (N=50). A total of 20 races of B. lactucae were used, 14 of which were NL-races, isolated from cultivated lettuce in the Netherlands. The other six races were isolated from wild L. serriola in Czechoslovakia. The accessions were initially screened with two races: NL1 and NL3. Accessions with resistance to one or both of these races were tested with the other races. Phenotypes with new resistance were found in accessions of all four Lactuca species. Of L. sativa, four accessions were found with resistance phenotypes that could not be explained by combinations of known major genes. Many accessions of L. serriola had resistance phenotypes that indicated the presence of unknown resistance genes. All interactions between accessions of L. saligna and races of B. lactucae were incompatible in leaf disc tests, except for four accessions, which showed some sporulation with race NL6. Several accessions of L. virosa were resistant to all races used. Other accessions of L. virosa gave a race-specific interaction with B. lactucae.     

An association between resistance to root aphid (Pemphigus bursarius L.) and downy mildew (Bremia lactucae Regel) in lettuce

Summary Many lettuce cultivars (Lactuca sativa L.) with high resistance to lettuce root aphid (Pemphigus bursarius L.) also carried the gene Dm-6 for specific resistance to downy mildew (Bremia lactucae Regel). This suggests the possibility of linkage between this gene and root aphid resistance. The origin of this association is discussed.     

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.     

Evaluation of seedling and adult plant resistance in European wheat cultivars to Australian isolates of <Emphasis Type="Italic">Puccinia striiformis</Emphasis> f. sp. <Emphasis Type="Italic">tritici</Emphasis>

A set of 105 European wheat cultivars was assessed for seedling resistance and adult plant resistance (APR) to stripe (yellow) rust in greenhouse and field tests with selected Australian isolates of Puccinia striiformis f. sp. tritici (Pst). Twelve cultivars were susceptible to all pathotypes, and among the remainder, 11 designated seedling genes (Yr1, Yr3, Yr4, Yr6, Yr7, Yr9, Yr17, Yr27, Yr32, YrHVII and YrSP) and a range of unidentified seedling resistances were detected either singly or in combination. The identity of seedling resistance in 43 cultivars could not be determined with the available Pst pathotypes, and it is considered possible that at least some of these may carry uncharacterised seedling resistance genes. The gene Yr9 occurred with the highest frequency, present in 19 cultivars (18%), followed by Yr17, present in 10 cultivars (10%). Twenty four cultivars lacked seedling resistance that was effective against the pathotype used in field nurseries, and all but two of these displayed very high levels of APR. While the genetic identity of this APR is currently unknown, it is potentially a very useful source of resistance to Pst. Genetic studies are now needed to characterise this resistance to expedite its use in efforts to breed for resistance to stripe rust. Colin R. Wellings seconded from NSW Department of Primary Industries.     

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.     

Inheritance of resistance to angular leaf spot in common bean and validation of the utility of resistance linked markers for marker assisted selection out side the mapping population

Inheritance of resistance to angular leaf spot (ALS) disease caused by Phaeoisariopsis griseola (Sacc.) Ferr was investigated in two common bean cultivars, Mexico 54 and BAT 332. Both Andean and Mesoamerican backgrounds were used to determine the stability of the resistance gene in each of the two cultivars. Resistance to P. griseola was phenotypically evaluated by artificial inoculation with one of the most widely distributed pathotypes, 63–39. Evaluation of the parental genotypes, F₁, F₂ and backcross populations revealed that the resistance to angular leaf spot in the cultivars Mexico 54 and BAT 332 to pathotype 63–39 is controlled by a single dominant gene, when both the Andean and Mesoamerican backgrounds were used. Allelism test showed that ALS resistance in Mexico 54 and BAT 332 to pathotype 63–39 was conditioned by the same resistance locus. Resistant and susceptible segregating populations generated using Mexico 54 resistant parent were selected for DNA extraction and amplification to check for the presence /absence of the SCAR OPN02 and RAPD OPE04 markers linked to the Phg-2 resistance gene. The results indicated that the SCAR OPN02 was not polymorphic in the study populations and therefore of limited application in selecting resistant genotypes in such populations. On the other hand, the RAPD OPE04 marker was observed in all resistant individuals and was absent in those scored susceptible based on virulence data. Use of the RAPD OPE04 marker in marker-assisted selection is underway.     

The Level of Field Resistance to Bremia lactucae in Lettuce (Lactuca sativa) Cultivars Carrying the Resistance Gene Dm11

Forty-one lettuce cultivars (butterhead type), carrying race-specific resistance gene Dm11 in combination with some others, were studied for field resistance to Bremia lactucae. Significant differences in the level of field resistance were revealed. Most of the cultivars under study were highly susceptible under field conditions. There are evident relationships between combinations of race-specific Dm genes and field resistance. The cultivars with gene combinations Dm2, Dm3 and Dm11 are susceptible. A very high level of field resistance was detected in the cultivars with combination of genes Dm5/8, Dm6 and Dm16 (cvs. ‘Celia’, ‘Katanga’ and ‘Musette’). This is in close link with their Lactuca serriola origin and a high level of field resistance in progenitors.     

The identification and characteristics of field resistance to lettuce downy mildew (Bremia lactucae Regel)

Summary Eighty-one accessions of three Lactuca species which showed no recognisable race specific resistance to Bremia lactucae when tested in the laboratory as seedlings, exhibited different degrees of susceptibility when exposed to natural field infection. As a group, crisp genotypes had less mildew and a slower rate of disease development than other types of lettuce. Wild forms of Lactuca sativa and Lactuca serriola were particularly susceptible. In a further trial, the low field susceptibility of three lettuce cultivars (Iceberg, Batavia blonde de Paris and Grand Rapids) was confirmed. Disease development on cv. Iceberg was compared to that on the highly susceptible cv. Hilde in experiments where the two cultivars were grown either in close proximity or in isolation. The absolute level of attack on cv. Iceberg depended upon disease pressure and differences between the two types only became apparent approximately 8 wk after sowing.     

Inheritance of resistance to Zucchini yellow mosaic virus in the interspecific cross Cucurbita maxima × C. ecuadorensis

Summary More than 200 accessions of three wildLactuca species were screened in the laboratory for race-specific resistance toBremia lactucae. Only a fewLactuca entries showed resistance both as seedlings and in a leaf disc test. Accessions ofL. serriola andL. sativa had a low degree of resistance. The best entries were also compared under field conditions. TheL. saligna entries were totally free from disease in the field test. High resistance was also recorded in a fewL. serriola entries and in lettuce cultivars such as Saffier and Mariska.     

Disease resistance in protected crops and mushrooms

Summary Cultivars of tomatoes, cucumbers, lettuce and peppers have been bred for resistance to one or more pathogens. Some tomato and cucumber cultivars have resistance to a wide range of diseases. Resistance has been transient in many cases and a succession of cultivars with new genes or new combinations of resistance genes has been necessary to maintain control. There has been a number of notable exceptions and these have included durable resistance to such pathogens asFulvia fulva and tomato mosaic virus. With lettuce the resistance situation is complicated by the occurrence of fungicide resistant pathotypes. There are no strains ofAgaricus bisporus purposely bred for disease resistance.In protected flower crops only resistance to Fusarium wilt in carnations has been purposely bred but differences in disease resistance are apparent in cultivars of many ornamental crops. This is particularly so in chrysanthemums where there are cultivars with resistance to many of the major pathogens. Similar situations occur with other flower crops and pot plants. Cultivars of some species have not been systematically investigated for resistance.The need for genetic resistance will increase with the further reduction, in the limits on pesticide use and an increasing public awareness and importance of pesticide pollution.ADAS is an executive agency of the Ministry of Agiculture, Fisheries and Food and the Welsh Office.     

Comparative genetic analysis of field resistance to downy mildew in the lettuce cultivars ‘Grand Rapids’ and ‘Iceberg’

Downy mildew on lettuce is currently controlled using host resistance genes (Dm genes) that confer race-specific resistance in seedlings. Field resistance (FR) that is active in adult plants but not seedlings was identified in the cvs. Grand Rapids and Iceberg. The goal of our study was to evaluate the utility of Grand Rapids as a source of novel Bremia resistance alleles, particularly in comparison with Iceberg. To measure FR, downy mildew symptoms were evaluated following natural infection in field experiments. The responses of Grand Rapids and Iceberg were similar in many respects. Although both cultivars had a small percentage of plants exhibiting disease symptoms, the average disease ratings were as low as for cultivars with effective Dm genes. We observed no evidence for race specificity. FR was effective over 3 years of our study, despite documented variation within pathogen populations. Both cultivars lacked all known seedling resistance genes except Dm13, which was not responsible for the resistance observed in field experiments. Similar segregation of FR was observed in F₂ populations for both Grand Rapids and Iceberg. The presence of highly susceptible families within Grand Rapids × Iceberg populations suggested the presence of at least one unique resistance allele in each cultivar. Preliminary genetic analysis of FR from Grand Rapids revealed a high estimate of narrow-sense heritability that suggested simple inheritance, but single gene models did not fit the observed data. Our results suggest that Grand Rapids may represent an underutilized resource for controlling downy mildew in lettuce.     

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.     

Development and characterization of a new 1BL.1RS translocation line with resistance to stripe rust and powdery mildew of wheat

The 1BL.1RS wheat-rye translocation from Petkus rye has contributed substantially to the world wheat production. However, following the breakdown of disease resistance genes in 1RS, its importance for wheat improvement decreased. We have developed a new 1BL.1RS line, R14, by means of crossing rye inbred line L155, selected from Petkus rye to several wheat cultivars. One new gene each, for stripe rust and powdery mildew resistance, located on 1RS of the line R14, are tentatively named YrCn17 and PmCn17. YrCn17 and PmCn17 confer resistance to Puccinia striiformis f. sp. tritici pathotypes that are virulent on Yr9, and Blumeria graminis f. sp. tritici pathotypes virulent on Pm8. These two new resistances, YrCn17 and PmCn17, are now available for wheat improvement programs. The present study indicates that rye cultivars may carry yet untapped variations as potential sources of resistance.     

Characterization of the resistance to <Emphasis Type="Italic">Phytophthora infestans</Emphasis> in local potato cultivars in Bolivia

This experiment was carried out to investigate whether and how much field resistance to late blight, caused by Phytophthora infestans, is present in the local cultivated potato germplasm. In total 36 entries were compared in a field experiment in an area highly conducive to late blight development. Of the 36 cultivars 32 were local cultivars belonging to five Solanum species, S. tuberosum (1 accession), S. andigena (18), S. juzepczukii (2), S. stenotomum (9) and S. ajanhuiri (2). The other four cultivars were derived from breeding programmes, one being the Dutch cultivar Alpha used as a highly susceptible control. The 36 cultivars were planted according to a simple 6 × 6 lattice design with three replicates. Each replicate was divided in six incomplete blocks each with six cultivars. The disease severity was assessed weekly during 9 weeks starting 48 days after planting. The area under the disease progress curve (AUDPC) was used as a measure of the field resistance. Nine isolates from surrounding potato fields were tested for their virulence to the resistance genes R1–R11 using 22 differential cultivars. The components of the field resistance of 19 of these cultivars were compared in the greenhouse using a local isolate with virulence to all known R-genes, except to R9. The nine isolates represented seven races with a race complexity varying from 7 to 10 virulence factors. All isolates carried virulence against R1, R2, R3, R7, R10 and R11, while virulence against R9 was absent. The AUDPC among the 32 local cultivars ranged from very large, significantly larger than that of ‘Alpha’ to very small. The AUDPC from S. stenotomum accessions ranged from very large to intermediate, those from S. andigena accessions from large to very small. Especially among the S. andigena accessions interesting levels of field resistance were found. Four components of field resistance were assessed, latency period (LP), lesion size (LS), lesion growth rate (LGR) and relative sporulation area (RSA). All four showed a considerable variation among the cultivars. The LP ranged from 3½ to 6 days. The LS ranged from 225 mm² to 20 mm². The LGR varied about six-fold, the RSA more than 10-fold. The components tended to vary in association with one another. LP and LGR were well associated with each other and had a significant correlation with the AUDPC.     

Regional phenotypic diversity of Puccinia triticina and wheat host resistance in western Europe, 1995

Pathogenicity data from surveys of Puccinia triticina (formerly P. recondita f. sp. tritici) conducted in western Europe in 1995 were analysed to compare the structure of regional populations of the pathogen. Many of the populations differed in phenotypic diversity and pathotypic composition, even though they occurred within a single epidemiological unit, suggesting that local factors may influence the establishment and propagation of individual pathotypes in the regional populations. Neighbouring regions were more similar than distant regions, and all regions shared at least one pathotype, except populations in northern Italy and Scotland. A high degree of similarity was found between populations in northern France and Great Britain, providing strong evidence of free movement of inoculum between these regions. Resistance genes were postulated for a selection of 91 wheat cultivars, representing those most commonly grown in western Europe in 1995. Thirteen cultivars lacked detectable seedling resistance genes and the remaining 78 possessed from one to three resistance genes; those detected were Lr1, Lr3a, Lr10, Lr13, Lr14a, Lr17b, Lr20, Lr26 and Lr37. The most commonly detected resistance gene was Lr13, which was present singly or in combination with other resistance genes in 48 cultivars (53%). The gene Lr14a was detected in 18 cultivars, Lr26 was present in 16 cultivars. The role of host selection in the composition of the regional populations of P. triticina in western Europe in 1995 was difficult to assess on the basis of the results obtained, since virulence data were not available for Lr13 and Lr14a. This revised version was published online in August 2006 with corrections to the Cover Date.     

Response of Lettuce Cultivars Carrying the Resistance Gene Dm11 to Isolates of Bremia lactucae from Lactuca serriola

Seven Bremia lactucae isolates from Lactuca serriola were tested for pathogenicity on 41 Lactuca sativa cultivars carrying a race specific resistance gene Dm11 which originated from L. serriola. Some of these cultivars also carried other Dm genes. Sporulation was observed in 34 % of the 287 cultivar/isolate combinations, but it was limited in most cases. A compatible response was recorded in 7.3% of cultivar/isolate. Cultivars with Dm 11 alone, or in combination with Dm 2, exhibited a higher decree of susceptibility then those also carrying Dm 3. There were differences in specific virulence among the isolates under study. Nearly all the isolates studied were virulent on newly described resistance genes Dm 15, Dm 16, and Dm 7 +? which originate from L. serriola.     

Pathogenicity of the Rhynchosporium secalis population in the Western Cape province of South Africa

The virulence spectra of 50 Rhynchosporium secalis isolates from a population in the Western Cape province of South Africa were determined, and 21 races were detected when evaluated against 17 differential cultivars. The virulence spectrum of the R. secalis population shows considerable variation, and carries unnecessary virulence genes which is quite unexpected, since chiefly susceptible barley cultivars are grown in the south Western Cape. The two most prevalent races, namely races 4 and 7 had three and four virulence genes respectively. Both race 4 and 7 were virulent on the most susceptible cultivars, West China, Steudelli, C.I.8618 and C.I.2226. Considering the resistance genes reported for the cultivars Atlas 46, Turk, and C.I.3515 which showed no susceptible cultivar-pathogen interaction, it would appear that the Rh-Rh3-Rh4 complex is primarily involved in conferring resistance to the local R. secalis isolates This revised version was published online in July 2006 with corrections to the Cover Date.     

Partial resistance in lettuce to downy mildew (Bremia lactucae). 2. Differential interactions between plant genotypes and fungus races and the relationship of latent period,infection frequency and number of infected leaves with the level of partial resistance

Summary In three experiments differential interactions between lettuce genotypes and Bremia lactucae races were investigated and also the influence of latent period, infection frequency and number of infected leaves on the level of resistance.In experiment 1 very significant differential interactions were observed mainly caused by two plant genotypes (PIVT 82 and PIVT 315) and race NL 7. In experiments 2 and 3 differential interactions were absent or of minor importance. Results suggest that partial resistance often was of a race-non-specific character. Latent period appeared to be a major component of resistance in the lettuce-Bremia lactucae relationship. It determines the onset of the next cycle of spore production and is also indicative of fungus growth in host tissue. Compared with infection frequency and number of infected leaves, latent period shows a small error variation and can efficiently be measured. The closeness of the relationship of the number of primary infections and the number of infected leaves with resistance level varied, depending on the experiment. The above three components appeared to be associated but the degree of the association varied. The variable behaviour of the three components suggests that they are governed by different genes.For the level as well as for the complexity and durability of the resistance, accumulation of as many components of resistance as possible is desirable. Therefore, in parents all such components should be analysed before crosses are carried out to improve resistance level. In the final products of crossing and selection the relative importance of components should also be investigated to determine whether an increased resistance level results from additional effects of the parents on one component (e.g. latent period) or from accumulation of different components in a plant genotype.Students at the Agricultural University Wageningen