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.     

Sources and inheritance of resistance to stemphylium leaf spot of lettuce

Summary None of the tested cultivars of lettuce was found resistant to Stemphylium leaf spot, a common disease in Israel. Within a Lactuca saligna population collected in wild lettuce in Israel, resistance was traced. Interspecific crosses of L. saligna x L. sativa were made and the mode of inheritance of resistance to this disease was studied. Resistance is apparently controlled by two genes: one dominant (Sm₁) and one recessive (sm₂).Contribution No. 1176-E 1984 series, from the ARO.     

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.     

Evaluation of resistance to Pseudomonas solanacearum in potato under controlled conditions

Summary Potato plantlets derived from in vitro propagation of three cultivars known for their field resistance (Cruza 148 and BR-63.65) or susceptibility (Désirée) to Pseudomonas solanacearum E.F. Smith were artificially inoculated under controlled conditions. The aim of this work was to determine the optimal inoculum concentration and the best observation period in which the cultivars would show different reactions to bacterial infection as expected on the basis of their field performance.A suitable statistical analysis of disease indices is proposed to distinguish between resistant and susceptible responses, with a particular care for the applicative needs and a univocal interpretation of the results. In order to evaluate the significance of sources of variation related to the observed mean differences, the analysis of variance and a convenient clustering procedure of disease index means were applied.The statistical analysis revealed that, under our conditions, an inoculum concentration of 5×10⁶ cfu/plant was suitable for separating resistant from susceptible responses, in accordance with the reactions already observed in field experiments by other authors. Also, differences among the three cultivars were best observed nine to twelve days after inoculation with the pathogen.     

Nematode resistance in bananas: screening results on some wild and cultivated accessions of Musa spp.

Bananas cultivated for export all belong to Cavendish cultivars and are all recognized as very susceptible to nematodes, particularly to the burrowing nematode Radopholus similis and the lesion nematode Pratylenchus coffeae. Even if there have been many changes in the management of banana nematodes in large commercial banana plantations, chemical control still remains most often the last resort method to manage the nematodes, although the number of registered products is definitely declining. Therefore, nematode control though genetic improvement is gaining new interest worldwide. In this study, 55 banana accessions mostly diploids from the Musa acuminata genome group (AA) but including some triploid accessions (AAA), some diploids of the Musa balbisiana genome group (BB) and some interspecific hybrids (AAB, AB) were evaluated for resistance to four nematode species R. similis, P. coffeae, Meloidogyne incognita and M. arenaria. These experiments were conducted in a growth chamber under controlled conditions. All banana accessions were susceptible to nematode species, although many different levels of susceptibility were detected. This study confirmed the good resistance status to R. similis of some cultivars from the Pisang jari buaya and Pisang batuau subgroups and the partial resistance of 17 diploid accessions significantly different from the susceptible reference cv. Grande Naine. This study also showed that 12 diploid accessions exhibited a partial resistance to P. coffeae, including some usual or potential genitors belonging to the wild diploids subspecies burmannica (cvs. Long Tavoy 1 and 2) and burmannicoides (cv. Calcutta 4). No source of resistance to Meloidogyne spp. was found. These screening results, combining for the first time four nematode species, are discussed within the scope of banana breeding in order to produce parental diploid lines with single or combined nematode resistances and further develop triploids that can substitute existing susceptible commercial cultivars.     

Genotypical differences in nitrate content inLactuca sativa L. and related species and correlation with dry matter content

Summary Nitrate content was measured in 135 genotypes of cultivated lettuce (Lactuca sativa L.) and 21 genotypes of wildLactuca spp. grown in two experiments. In Experiment 1 plants were grown on nutrient film. In Experiment 2 plants were grown in large pots of potting soil. Within cultivated lettuce five plant types were distinguished and within each of them genotypes were found with low nitrate content. The coefficient of variation in the experiment with plants growing in large pots was not larger than in the experiment with plants growing on nutrient film. In butterhead genotypes nitrate content was negatively correlated with dry matter content and positively with plant fresh weight.     

Inheritance of resistance to angular leaf spot in Nicotiana tabacum L. cultivars burley 21 and kentucky 14

Summary The genetics of resistance to angular leaf spot caused by Pseudomonas syringae pv. tabaci in Nicotiana tabacum cultivars Burley 21 and Kentucky 14 was investigated by studying disease reactions to three isolates of parental, F₁, F₂ and backcross generations derived from crosses between the resistant cultivars and the susceptible cultivar Judy's Pride. Studies were conducted in the greenhouse and in field plant beds. Chi-square values were computed to determine whether the observed ratios for disease reactions deviated from expected Mendelian ratios for a single, dominant gene controlling resistance to angular leaf spot in tobacco. Based on the resistance of the F₁ and the backcross generation to the resistant parent (BC-R), a 3 resistant: 1 susceptible segregation ratio in the F₂, and a 1 resistant: 1 susceptible segregation ratio in the backcross generation to the susceptible parent (BC-S), it was concluded that resistance to three isolates of Pseudomonas syringae pv. tabaci is governed by a single, dominant gene.     

Susceptibility to leaf silvering in the cultivar groups of summer squash

Summary Summer squash (Cucurbita pepo) cultivars, three from each of the six cultivar groups, were grown under field and greenhouse conditions at Fort Pierce, Florida, U.S.A. in the presence of sweetpotato whiteflies (Bemisia tabaci) and observed for severity of leaf silvering. All cultivars were affected by leaf silvering, but some had less severe symptoms than others. The cocozelle group was the least susceptible to silvering. There appeared to be an association between geographic origin of cultivars and their susceptibility to silvering.Florida Agricultural Experiment Station Journal Series no. R-02844.     

Variation in virulence and resistance in the bean-bean rust pathosystem in Ecuador

Twenty two Uromyces appendiculatus isolates were tested on 20 differential and 25 Ecuadorian Phaseolus vulgaris cultivars in the seedling stage. Based on the infection types 20 races could be discerned. The Ecuadorian cultivars differed greatly in their reaction to the isolates, from resistant to only one isolate (`Red Small Garden') to resistant to all isolates (’G2333‘). The isolates showed a wide range of virulence to the Ecuadorian cultivars, from virulent to only two cultivars (isolate 13) to virulent to 21 cultivars (isolates 5 and 23). Seven cultivars with a basically susceptible infection type appeared to differ greatly in quantitative resistance when tested in three consecutive crop cycles. The disease severities in percentage leaf area affected averaged over the three cycles ranged between 83.9% for ‘Red Small Garden’ and 13.1% for ‘INIAP-414’. Race-specific resistance does not seem an advisable breeding strategy, but the quantitative resistance offers a good alternative.     

A demonstration of the inheritance of field resistance to lettuce downy mildew (Bremia lactucae Regel.) in progeny derived from cv. Grand Rapids

Summary The inheritance of field resistance to downy mildew in lettuce derived from cv. Grand Rapids was studied in F₂, F₃ and F₄ progeny from a cross with the more susceptible cv. Lobjoits Green Cos. The results for 32 randomly produced F₃ families and the F₂ population corresponded closely although the absolute level of resistance varied from season to season according to disease pressure and environmental conditions. The field resistance character showed a quantitative pattern of inheritance but the rapid response to selection at F₃ implied that only a few genes may be involved. F₄ families raised from families selected from the two extremes of the F₃ distribution fell into distinct categories with resistance levels corresponding to that of their F₃ parents. An association between resistance and morphological features of the resistant parent was present at F₂ but not in subsequent generations when individuals and families which were as resistant as cv. Grand Rapids and yet bore little morphological resemblance to this cultivar were readily recovered.     

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.     

Interactions between plant and aphid genotypes in resistance of lettuce to Myzus persicae and Macrosiphum euphorbiae

Summary Six lettuce lines, representing two types of resistance to the green peach aphid, Myzus persicae, and a control line with high susceptibility to M. persicae were tested for resistance to six different clones of Myzus persicae and two clones of the potato aphid, Macrosiphum euphorbiae.The clones of M. persicae showed very different levels of aggressiveness on lettuce: two had a high level of reproduction, two had an intermediate level and two were poorly adapted to lettuce as a host. Differences between lettuce lines in aphid reproduction increased with increasing aggressiveness of the aphid clone, which means that aggressive clones are most effective for selection purposes. No evidence was found for clone-specific plant genotype reactions, meaning that lines resistant to one clone will also be resistant to other clones of M. persicae, allthough not neccessarily at the same level. The lettuce lines selected for partial resistance to the aggressive clone WMp1 were completely or almost completely resistant to less aggresive clones.No differences in level of reproduction were found between the two clones of M. euphorbiae and no relation was observed between resistance to M. persicae and M. euphorbiae, indicating the species-specific character of resistance to leaf aphids in lettuce.     

The influence of disease complexes involving Leptosphaeria (Septoria) nodorum on detection of resistance to three leaf spot diseases in wheat

Summary In controlled inoculation studies with Septoria nodorum and Pyrenophora tritici-repentis, estimates of the relative proportion of each pathogen demonstrated differences in the responses of cultivars to pathogen mixtures that were not apparent from measurements of diseased leaf areas. Under field conditions estimates of the relative proportion of S. nodorum, P. tritici-repentis and S. tritici varied between field screening locations in Western Australian but also between lines within locations. Lines with known resistance to P. tritici-repentis and S. tritici, but susceptible to S. nodorum, could not be distinguished from susceptible lines on the basis of leaf area diseased or grain weight depression when S. nodorum was present in the disease complex. Such conditions, while suitable for the selection of combined resistance to these pathogens, were unsuitable for identifying resistance to individual pathogens. As symptoms were similar, the proportion of diseased leaf area sporulating with each pathogen provided a means of measuring the variation in disease development induced on lines varying in resistance. Knowledge of the components of disease and their relative importance were essential in understanding varietal response information under mixed infections of these leaf spot pathogens.     

The exploitation of genetic resources of Brassica oleracea in breeding for resistance to clubroot (Plasmodiophora brassicae)

Summary About 1000 Brassica oleracea accessions were evaluated in glasshouse tests for response to Plasmodiophora brassicae (clubroot). Resistance was confirmed in some north and west European kales and cabbage. A new source of resistance in cabbage, from Eire, is reported. Most other accessions were highly susceptible but lower levels of susceptibility were observed in open pollinated Brussels sprouts and forms of south European cabbage, cauliflower and broccoli. Modern breeding (as in the production of hybrid cultivars) appears to have resulted in increased susceptibility in several crop types. The implications of these results for the exploitation of germplasm are discussed.     

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.     

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.     

Genetic variation in Zantedeschia spp. (Araceae) for resistance to soft rot caused by Erwinia carotovora subsp. carotovora

Bacterial soft rot caused by Erwiniacarotovora subsp. carotovora is amajor disease in Zantedeschia spp.,particularly in cultivars from the sectionAestivae. The disease can partiallybe controlled by cultivation measures, so acombination with resistant cultivars couldeffectively protect the crop. However,resistant commercial Aestivaecultivars are not available yet. By meansof a recently developed non-destructiveresistance test, variation inaggressiveness was observed among fiveisolates of Erwinia carotovora subsp.carotovora without interactionsbetween the isolates and three Zantedeschia accessions. Within elevenaccessions of Z. aethiopica,variation was observed from almost completeto moderate resistance, while theZ. odorata accession was susceptible.All 21 Aestivae cultivars weresusceptible. Within the Aestivaespecies, Z. elliotiana and Z.pentlandii were also susceptible, butwithin twelve accessions of Z.albomaculata, as well as in six accessionsof Z. rehmannii, variation was foundfrom susceptible to moderately resistant.Hence, new sources of resistance wereidentified that show good potentials forresistance breeding.     

Triforine sensitivity in lettuce: A potentially useful genetic marker

Summary A commercial formulation of triforine¹, a systemic fungicide not normally used on lettuce, induced marked phytotoxicity on certain cultivars. The phytotoxic agent was found to be triforine and not one of the other components of the formulation.Test crosses between resistant and susceptible cultivars have shown that resistance is recessive and determined by a single major gene for which the symbol tr is proposed. The character is unlikely to be of much significance in lettuce crop protection but is potentially useful in genetic studies and for the production of hybrid lettuce seed.     

Inheritance of the resistant reaction of the lettuce cultivar `Grand Rapids' to the southern root-knot nematode Meloidogyne incognita (Kofoid & White) Chitwood

Resistance to the southern root-knot nematode Meloidogyne incognita Chitwood would be an important attribute of lettuce Lactuca sativa L. cultivars adapted to both protected and field cultivation in tropical regions. `Grand Rapids' and a few other cultivars are reported to be resistant to this nematode. In this paper, we studied the inheritance of the resistant reaction of `Grand Rapids' (P₂) in a cross with a standard nematode-susceptible cultivar Regma-71 (P₁). F₁(Regina-71 × Grand Rapids) and F₂ seed were obtained, and inoculated along with the parental cultivars with different races of M. incognita to evaluate nematode resistance. Broad sense heritability estimates for the number of galls and of egg masses per root system, gall size and gall index were generally in the order of 0.5 or higher. Class distributions of these variables over generations P₁, P₂, F₁ and F₂ were in agreement with simulated theoretical distributions based on monogenic inheritance models. F₃ families were obtained from randomly sampled F₂ plants and tested for reaction to the nematode. The frequency ratio of homozygous resistant, segregating and homozygous susceptible F₃ families did not differ from the 1:2:1 ratio expected from monogenic inheritance. M. incognita resistance appears to be under control of a single gene locus. The Grand Rapids allele (for which the symbol Me is proposed) is responsible for the resistant reaction, and shows high (though incomplete) penetrance, variable expressivity and predominantly additive gene action. This revised version was published online in July 2006 with corrections to the Cover Date.