Patterns of genetic diversity in landraces of common bean (Phaseolus vulgaris L.) from Galicia

Knowledge of patterns of genetic diversity among existing cultivars helps to broaden the genetic base of new cultivars and maximizes the use of available germplasm resources. This study examined the organization of diversity for morphological traits in 66 landraces of cultivated common bean (Phaseolus vulgaris) from Galicia and its relationship with phaseolin seed protein diversity. Data on growth habit, seed and pod traits obtained from field evaluations at two locations during the 1989—91 cropping seasons were subjected to multivariate statistical analysis. Cluster analysis based on 14 quantitative variables and five qualitative variables identified 11 groups. The landraces were also characterized by phaseolin electrophoresis. The results allowed separation of these landraces into Middle and Andean American groups, which could be further divided into at least eight groups within the Andean American cultivars and three within the Mesoamerican cultivars. These groups in turn corresponded to the previously described races Nueva Granada and Peru of South American origin, and races Durango, Jalisco and Mesoamerica from the Middle American domestication centre. These results confirm the existence of two major groups of germplasm in the cultivated common bean landraces from Galicia, Mesoamerican vs. Andean American.     

Variation in Phaseolus vulgaris in the Northwest of the Iberian Peninsula

A collection of 51 accessions of Phaseolus vulgaris L. from N.W. of the Iberian Peninsula was evaluated for phenological, morphological, and agronomic characters for two years to study patterns of variation. Data were subjected to cluster analysis by variables choice, and six groups could be identified. These in turn corresponded to each previously described races ‘Nueva Granada’ and ‘Chile’ of South American origin, and races ‘Durango’ and ‘Mesoamerica’ of Middle American domestication center. Accessions falling in race ‘Nueva Granada’ had two distinct gene pools.     

Improvement of large-seeded common bean cultivars under sustainable cropping systems in Spain

Approaches are needed to broaden the genetic base and improve earliness and yield potential of large-seeded beans under sustainable cropping systems. The objective of this research was to develop adapted dwarf bean populations having a commercial seed quality and yield suitable for the production in the South of Europe. The original base populations were produced from crosses between genotypes within each Mesoamerica, Nueva Granada and Peru races, and between Peru and Nueva Granada, and Mesoamerica and Nueva Granada races. Visual mass selection for plant performance was practised in the F₂ and F₃ generations. In the F₄ and F₅, single plants were harvested under two cropping systems (sole cropping and intercropping with maize). From F₄, selection was based on precocity, combined with seed yield and seed commercial type. The F_4:7 selected lines from each original population were compared with their parents and five checks at four environments and two cropping systems. Differences among environments, populations, parents and checks were observed for all traits. Under intercropping with maize, there was a 50% reduction in seed yield. Yield of parents and checks belonging to Andean South American races, intraracial (Nueva Granada × Nueva Granada) and interracial (Nueva Granada × Peru) populations, was higher than that of those of Middle American origin. Intraracial crosses within large-seeded Andean South American (Peru race) and Middle American gene pools (Mesoamerica race) did not produce lines yielding more than the highest yielding parent. Only two large-seeded lines selected from crosses between small- and large-seeded gene pools out-yielded the best parent and check cultivar.     

Selection for seed yield in Middle American versus Andean × Middle American interracial common-bean populations

The objective of this study was to compare the seed yield, seed weight, and maturity of random lines derived from two contrasting populations of common bean ( Phaseolus vulgaris L.): Middle American (TR 7790) and Andean × Middle American (TR 7791). TR 7790 involved crosses between common-bean races of Middle American origin only (Mesoamerica and Durango), whereas TR 7791 involved crosses between races of Middle American and Andean origin (Mesoamerica and Nueva Granada). A total of 85 random lines from each of these two double-cross, interracial populations, a bulk sample of both populations, and all parents were evaluated in a replications-in-set design in eight environments in Colombia between 1991 and 1993. The mean yields of parents, population bulk, and lines from TR 7790 were significantly higher (P < 0.05) than those of TR 7791. Six lines in TR 7790 and nine lines in TR 7791 yielded significantly higher (P < 0.05) than the mean of their parents. Four lines from TR 7790 out-yielded the highest-yielding line from TR 7791. Moreover, while three lines from TR 7790 out-yielded its best parent, none of the lines from TR 7791 out-yielded its highest yielding parent. Thus, substantial yield gains should be expected from interracial populations within the Middle American group of common beans, whereas selection for higher seed yield from Andean × Middle American populations would seem problematic.     

A core collection of common bean from the Iberian peninsula

Characterization of crop germplasm from specific regions helps understand the patterns of genetic variation that facilitates further germplasm collection, characterization, management and their more efficient utilization in genetics, breeding and other studies. Common bean (Phaseolus vulgaris L.) is a traditional crop in the Iberian Peninsula (Spain and Portugal) where subsistence farmers have been growing and maintaining their own cultivars since their introductions from the Americas in the sixteenth century. Our objectives were to: (i) characterize diversity in the landraces collected from the Iberian Peninsula and (ii) form a core collection. Of 388 landraces from the major production regions characterized for 34quantitative and 13 qualitative characters, including morphological, agronomic and biochemical traits, 74.7% had an Andean origin, 16.8% a Mesoamerican origin and 8.4% had seed mixtures or were recombinants between the two gene pools. Landraces of indeterminate climbing growth habit Type IV(47.2%) and bush determinate Type I(26.4%) with large (52.9%) and medium(27.4%) seeds of white (38.8%) and cream(25.9%) colour were predominant. Similarly, the ‘T’ phaseolin pattern and common bean race Nueva Granada were the most frequent(51%). Some exceptionally large-seeded landraces of Andean (e.g., PHA-0917 with119 g 100-seed weight^-1) and Mesoamerican (e.g., PHA-0399 with 66 g100-seed weight^-1) were found. These and other possible recombinants between the two gene pools merit further investigation. Fifty two landraces (13%) were chosen to form a core collection representing the genetic diversity in the Iberian Peninsula. This revised version was published online in July 2006 with corrections to the Cover Date.     

Selection potential for seed yield from intra- and inter-racial populations in common bean

Common bean populations from crosses between lines of different races are thought to be more promising for selection of high yield potential than those from intra-racial crosses. Three distinct diallel crosses were made to test this hypothesis and to determine the possibility of substituting diallel crosses for multivariate techniques that estimate genetic divergence. The crosses were between races Mesoamerica × Mesoamerica, Mesoamerica × Durango and Jalisco, and Mesoamerica × Nueva Granada. The parents and the resulting F₄ populations were evaluated at Lavras-MG, Lambari-MG and Patos de Minas-MG, Brazil. The diallel analysis of seed yield was done and the genetic divergence estimated by Mahalanobis distance. Estimates of general and specific combining ability indicated that some inter-racial populations were more promising for selection to increase seed yield than intra-racial populations. However, due to their undesirable seed color and size, and growth habit, especially in a short term breeding program, the chances of obtaining high yielding lines with an acceptable bean is reduced. Genetic divergence was not a good measure to choose the parents because usually, the most divergent groups included were not adapted lines. This revised version was published online in July 2006 with corrections to the Cover Date.     

Selection for seed yield in Andean intra-gene pool and Andean × Middle American inter-gene pool populations of common bean

Yields of large-seeded Andean (A) common bean (Phaseolus vulgaris L.) cultivars of Chile and Nueva Granada races are 40 to 60% lower compared to their Middle American (M) counterparts of small-seeded Mesoamerica and medium-seeded Durango races. Our objective was to use the concept of congruity backcrossing between Andean x Middle American inter-gene pool [AM 11833 = A 483 (A)///// A 686 (M) //// PVA 800A(A)/// ‘Carioca’ (M)// Carioca (M)/ G 19833 (A)] and between races within Andean gene pool [i.e., intra-gene pool Andean, AA 11834 = A483 (A)//// ‘Cardinal’(A) /// ‘Blanco Español’(A) // BlancoEspañol (A) / ‘Taylor’ (A)] to compare selection for seed yield improvement of large-seeded Andean beans. Seven hundred sixty seven F₂-derived F₃ (F_2:3) families were produced for each population. Visual appraisal for total plant performance, combined with seed yield from non-replicated plots was used for selection of 551 families in F_2:3, 182 families in F_2:4, and 91 families in F_2:5 in each population. Eight hundred twenty three F_5:6 lines were developed from the 91 F_2:5 families in each population. Visual selection, combined with seed yield in non-replicated plots was again used to select 294 lines in F_5:6 in each population. Similarly, 44 highest yielding F_5:7 lines were selected in AM 11833 and 39 F_5:7 lines in AA 11834. Thus, single plant selections were made in the F₂ and F₅, and plants within each plot were harvested in bulk in F₃, F₄, F₆, and F₇. Thirty nine F_5:8 lines from AA 11834 and 44 lines from AM 11833, parents, and checks were evaluated at Popayán and Quilichao, Colombia in 1998 and 1999. Selected lines in both populations, on average, out-yielded the mean of their large-seeded Andean parents. Mean yield of the lines selected from AM 11833 was 50% higher than AA 11834 lines. Twelve F_5:8 lines out-yielded the highest yielding Andean parents G 19833 and A 483 in AM 11833, whereas only one line yielded significantly higher (p < 0.05) than the highest yielding parent A 483 in AA 11834. However, none of selected lines out-yielded small-seeded Middle American parents used in AM 11833 (A 686 and ‘Carioca’). The mean 100 seed-weight of AA 11834 was 36 g compared to 28 G for AM 11833 F_5:8 lines. Selected lines had similar days to maturity as parents in AM 11833, and matured 3 d later in AA11834. Correlation coefficients between yield and 100 seed-weight were negative in both populations. Yield and days to maturity were positively correlated in AA 11834.     

Pathogenic variation in, sources of, and breeding for resistance to Phaeoisariopsis griseola causing angular leaf spot in common bean

If we are to breed common bean (Phaseolus vulgaris L.) for durable resistance to diseases, we must understand pathogenic variation and find sources of resistance. Our first objective was to determine the patterns of pathogenic variation found among isolates of Phaeoisariopsis griseola (PG), the fungus that causes angular leaf spot (ALS) in common bean. We characterized 433 PG isolates from 11 Latin American and 10 African countries, using differential cultivars, isozymes, and/or random amplified polymorphic DNA (RAPD) markers. We also systematically screened, for ALS resistance, common bean accessions from the world collection held at CIAT, and assessed the progress so far made in breeding for resistance to ALS. Despite their great diversity within and between countries on both continents, the PG isolates were classified into two major groups: Andean, and Middle American. Although each group had internal differences for virulence, and biochemical and molecular characteristics, the ‘Andean’ PG isolates were more virulent on common beans of Andean origin, than on those of Middle American origin, thus, suggesting a host-pathogen co-evolution. The ‘Middle American’ PG isolates, although more virulent on common beans from Middle America, also attacked Andean beans, thus, exhibiting a much broader virulence spectrum. To find sources of resistance, we tested 22,832 common bean accessions against naturally occurring PG isolates in the field at CIAT's Experiment Station, Quilichao, Colombia, between 1985 and 1992. The resulting 123 intermediate (scores of 4 to 6) and resistant (scores of 1 to 3) accessions were then tested in the greenhouse against selected 14 PG isolates of diverse origins. Nineteen accessions were intermediate or resistant to at least 13 of 14 PG isolates. Similarly, of 13,219 bred lines tested in the field between 1978 and 1996, 89 were intermediate or resistant. Of these, 33 bred lines proved intermediate or resistant to at least eight of nine PG isolates to which they were challenged in the greenhouse. We suggest that, to breed for durable resistance to ALS, common bean populations should be developed from crosses between Andean and Middle American gene pools. The populations should then be systematically evaluated and selected against the broadest range of the most virulent PG isolates of diverse evolutionary origins. This revised version was published online in August 2006 with corrections to the Cover Date.     

Andean beans (Phaseolus vulgaris L.) with resistance to the angular leaf spot pathogen (Phaeoisariopsis griseola) in southern and eastern Africa

Common beans (Phaseolus vulgaris) are separated into two distinct groups: Andean and Middle American. We identified CAL 143 as the first Andean bean with resistance to angular leaf spot disease caused by Phaeoisariopsis griseola. Angular leaf spot is the most widespread and economically important bean disease in southern and eastern Africa, and it is especially severe on the extensively grown Andean beans. Cal 143 was resistant in Malawi, South Africa, Tanzania, and Zambia, but it was susceptible in Uganda. This was attributed to the presence of races of P. griseola in Uganda not present in the other countries. We identified two additional Andean bean lines, AND 277 and AND 279, with resistance to angular leaf spot in Malawi. We also characterized the virulence diversity of 15 isolates of P. griseola from southern and eastern Africa into nine different races. Five of six isolates from Malawi and two of seven from Uganda, obtained from large-seeded Andean beans, were characterized into four different races considered Andean. These were compatible only or mostly with large-seeded Andean cultivars. The other eight isolates from Uganda, Malawi, and the Democratic Republic of Congo, obtained from a small- or medium-seeded Middle American beans, were characterized into five different Middle American races. These were compatible with Middle American and Andean cultivars. CAL 143 was resistant or intermediate under greenhouse conditions to all but one of the same 15 isolates from southern and eastern Africa, but it was susceptible to an isolate from Uganda obtained from a medium-seeded Middle American bean. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diversity in seed quality traits of common bean populations from Basilicata (Southern Italy)

The diversity of nutritional and culinary traits in 21 populations of common bean from the Basilicata region was studied for three consecutive harvests (1995‐97). The results were compared with the six commercial cultivars widely grown in the region. Some populations deserve particular attention since they had large seeds, high protein content, low trypsin inhibitor levels and short cooking time. The cluster analysis showed that 18 populations clustered in two main groups and that the bush populations were clearly distinguishable from the climbing ones. When phaseolin type and some plant morphological traits were added to the nutritional and culinary traits to classify these populations, it was observed that they resembled the race Peru (Andean gene pool).     

Sources of resistance to angular leaf spot (Phaeoisariopsis griseola) in common bean core collection, wild Phaseolus vulgaris and secondary gene pool

Angular leaf spot (ALS) is one of the most devastating diseases of common bean (Phaseolus vulgaris L.) in tropical and subtropical countries. The causal fungus, Phaeoisariopsis griseola(Sacc.) Ferr. is highly variable and a diverse source of resistance genes is required to manage this disease. We evaluated a common bean core collection,primary and secondary gene pools and lines derived from inter-specific crosses of P. vulgaris and P. coccineus or P. polyanthus (secondary gene pool) for resistance to angular leaf spot. Of the 1441 accessiones in the core collection, only 2.2% were resistant to both Andean and Mesoamerican races of P. griseola, 28% were resistant only to Andean and 9% to Mesoamerican races. Of the 32 resistant accessions, 68%originated from Bolivia, Colombia,Guatemala and Mexico. More accessions from these countries should be examined for P. griseola reaction. Very few wild P. vulgaris accessions (4%), were resistant to ALS. In contrast, high levels of resistance (62%) were found in the secondary gene pool. Among the 1010 lines from inter-specific crosses, 109 lines were highly resistant. These genotypes from the primary and secondary common bean gene pools resistant to Andean and Mesoamerican races of P. griseola offer a potential for developing broad and durable ALS resistance. This revised version was published online in August 2006 with corrections to the Cover Date.     

The kappa race of Colletotrichum lindemuthianum and sources of resistance to anthracnose in Phaseolus beans

Summary The utilization of American and European bean cultivars as host differentials for distinction of races of Colletotrichum lindemuthianum has been discussed. The new race occurring at Ebnet. Germany, since 1973 is named kappa. It broke down resistance derived from the Are gene originating from Cornell 49–242. Resistance to this kappa race appeared to be present in some European and Asiatic bean cultivars as well as in some American bean accessions.     

Evaluation of resistance sources and inheritance of resistance in kidney bean to Indian virulences of Colletotrichum lindemuthianum

Summary Forty nine common bean lines comprising of exotic accessions and locally grown cultivars evaluated against Colletotrichum lindemuthianum exhibited differential resistance to its races in Himachal Pradesh, a north-western Himalayan state of India. Some exotic accessions like G 2333, Cornell 49242, PI 207262, Mexique 222, TO, Perry Marrow, Kaboon and Widusa were resistant to more than five Indian races, whereas two Indian accessions KRC-5 and Hans showed resistance to six and four races, respectively. However, nine accessions KRC-8, KR-40, KR-43, KR-81, KR-62-2, KR-90, KR-142, KR-148, and KR-216 were resistant to three races. Race specific resistance has been observed in different bean cultivars. Studies on inheritance of resistance in exotic accession G 2333 and Indian accession, KRC-5 showed that two independent dominant genes conferred resistance in G 2333 to race 3 and 515 and a single dominant gene controlled resistance in KRC-5 to race 775, indicating resistance from these sources is easily transferable to the locally adapted susceptible cultivars.     

Identification of resistance to rust (Uromyces appendiculatus) and powdery mildew (Erysiphe diffusa) in Portuguese common bean germplasm

Portugal has a diverse common bean germplasm, which is still grown in farmers' fields. In this work, we searched for resistance to rust (Uromyces appendiculatus) and powdery mildew (Erysiphe diffusa) in a representative collection of the Portuguese common bean germplasm. Despite many accessions depicting intermediate levels of resistance when compared to the susceptible check, 24 and 13 accessions showed low levels of infection, in spite of a compatible interaction (disease severity (DS) values lower than 5% and infection type (IT) of 4), to rust and powdery mildew, respectively, indicative of partial resistance. Moreover, a resistant reaction was observed in 11 accessions when inoculated with powdery mildew (IT = 0–1) and in additional 11 accessions (one in common) when inoculated with rust (IT = 0–2). The levels of resistance found in this report anticipate great potential of the Portuguese national germplasm, recently reported as genetically closer to the Andean common bean gene pool, for disease resistance breeding of this important crop.     

Combining ability for nodulation in common bean (Phaseolus vulgaris L.) genotypes from Andean and Middle American gene pools

Ten F₁'s obtained from crosses among five common bean genotypes of Andean (WAF 15, Mineiro Precoce and Batatinha) and Middle American (BAT 304 and Ouro) gene pools were assessed for their combining abilities for root nodulation with Rhizobium tropici strain CIAT 899. The plants were grown under controlled conditions and evaluated for number of nodules per plant (NN), nodule dry weight (NDW), mean nodule weight (MNW) and plant fresh weight (PFW). The subdivision of the treatment effects on the general (GCA) and specific combining effects (SCA) were performed according to Griffing's diallel analysis method 2, model 1. The analyses of variance and estimates of quadratic components showed that non-additive gene effects were more important in the expression of NN and PFW, whereas additive gene effect was predominant for NDW and MNW. A close association was observed between high number of nodules and GCA. Generally, crosses involving parents of different gene pools yielded hybrids with high positive estimates of SCA for all assessed traits. The genotypes of Andean origin WAF 15 and Mineiro Precoce are the most promising parents for breeding programs to increase NN and NDW in common beans. This revised version was published online in July 2006 with corrections to the Cover Date.     

An allelic series at the Co-1 locus conditioning resistance to anthracnose in common bean of Andean origin

In this study, we characterized the genetic resistance of the Andean bean cultivars Kaboon and Perry Marrow and their relation to other sources of anthracnose resistance in common bean. Based on the segregation ratio (3R:1S) observed in two F₂ populations we demonstrated that Kaboon carries one major dominant gene conferring resistance to races 7 and 73 of Colletotrichum lindemuthianum. This gene in Kaboon is independent from the Co-2 gene and is an allele of the Co-1 gene present in Michigan Dark Red Kidney (MDRK) cultivar. Therefore, we propose the symbol CO-1 ² for the major dominant gene in Kaboon. The Co-1 is the only gene of Andean origin among the Co anthracnose resistance genes characterized in common bean. When inoculated with the less virulent Andean race 5, the segregation ratio in the F₂ progeny of Cardinal and Kaboon was 57R:7S (p = 0.38). These data indicate that Kaboon must possess other weaker dominant resistance genes with a complementary mode of action, since Cardinal is not known to possess genes for anthracnose resistance. Perry Marrow, a second Andean cultivar with resistance to a different group of races, was shown to possess another resistant allele at the Co-1 locus and the gene symbol Co-1 ³ was assigned. In R × R crosses between Perry Marrow and MDRK or Kaboon, no susceptible F₂ plants were found when inoculated with race 73. These findings support the presence of a multiple allelic series at the Andean Co-1 locus, and have major implications in breeding for durable anthracnose resistance in common bean. This revised version was published online in July 2006 with corrections to the Cover Date.     

Geographical distribution of the DL1 and DL2 genes causing hybrid dwarfism in Phaseolus vulgaris L., their association with seed size,and their significance to breeding

Summary Dwarlism in F₁ hybrids has been observed in over 100 crosses of dry beans (Phaseolus vulgaris L.) at the Centro Internacional de Agricultura Tropical (CIAT), Cali, Colombia. In each cross, one parent always had small seeds and the other parent either medium or la ge ones. This apparent incompatibility between the two groups of germplasm was controlled by two complementary, dominant genes: DL₁ and DL₂. Smallseeded bean lines carried gene DL₁ and originated in Brazil, Colombia, Guatemala, and Mexico; medium for large-seeded bean lines carried gene DL₂ and were from Bolivia, Brazil, Chile, Colombia, Turkey, The United States, and West Germany. Thes two genes have probably played an important role in the evolution of dry bean forms of different seed sizes by serving as a genetic barrier or isolating mechanism, thus limiting free genetic recombination between the two germplasm groups.Apparent differences in the adaptiveness and yielding ability of the two groups of bean germplasm, smallys, medium- and large-seeded, and some breeding implications for manipulation of the genes causing F₁ hybrid dwarfism were also discovered.     

New sources of resistance to anthracnose and angular leaf spot of beans (Phaseolus vulgaris L.)

Summary over 13000 CIAT bean accessions were evaluated for their reactions to the anthracnose (Colletotrichum lindemuthianum) and angular leaf spot (Isariopsis griseola) pathogens over a 3 yr period. Among these accessions, 156 were resistant to all races of the anthracnose pathogen collected from Popayán, Colombia. Thirty were resistant to numerous races obtained from other parts of the world, including Europe. Although many of these new resistant sources originated in Mexico and Central America, they are quite diverse for geographic origin, plant type, seed color and seed size. In addition, more than 50 of the 156 lines were also resistant to isolates of I. griseola with diverse sources of origin throughout Colombia.     

Inter-and intraracial hybridization and selection for seed yield in early generations of common bean,Phaseolus vulgaris L.

Summary Five parents from each of four race groups of common bean (Phaseolus vulgaris L.) were hybridized to produce five crosses within each group. Also, five crosses were made for each of the six possible combinations among four groups. Parents, F₁ and F₂, and parents, F₂ and F₃ were evaluated for seed yield in 1990 and 1991, respecitively, at two locations in Colombia.Yield of parents belonging to Middle American races and crosses among them was higher than that of races of Andean origin. Positive correlations were found among the mid-parent value, F₁, F₂ and F₃. Also, the mid-parent value predicted the mean seed yield of all possible lines that could be derived from the F-generation in 42 out of 47 crosses. Four crosses, all between common bean races of Andean and Middle American origin, indicated a possible loss of favorable epistatic parental alleles. On average, mean yield of interracial F₁ hybrids was higher than that of intraracial ones.Positive heteroris (26.4%–123.8%) over the mid-parent in 31 crosses, and F₁s yielding higher (23.7%–91.8%) than the high parent in 20 crosses and yielding higher (22.1%–53.2%) than the highest control among all parents (MAM 13) in 12 crosses, were found. Heritability, estimated by the parent-offspring regression, ranged from 0.42± 0.07 to 0.49±0.04. Expected and realized gains from selection (at 20% selection pressure) ranged from 10.3% to 21.0% over the mean of F₁ hybrids and F₂ and F₃ population bulks.     

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.