Susceptible Lettuce Cultivars Research Articles

UV-B Induced Flavonoids Contribute to Reduced Biotrophic Disease Susceptibility in Lettuce Seedlings.

Biotrophic disease is one of the largest causes of decreased yield in agriculture. While exposure to ultraviolet B (UV-B) light (280–320 nm) has been previously observed to reduce plant susceptibility to disease, there is still a paucity of information regarding underlying biological mechanisms. In addition, recent advances in UV-LED technology raise the prospect of UV light treatments in agriculture which are practical and efficient. Here, we characterized the capability of UV-B LED pre-treatments to reduce susceptibility of a range of lettuce (Lactuca sativa) cultivars to downy mildew disease caused by the obligate biotroph Bremia lactucae. Innate cultivar susceptibility level did not seem to influence the benefit of a UV-B induced disease reduction with similar reductions as a percentage of the control observed (54–62% decrease in conidia count) across all susceptible cultivars. UV-B-induced reductions to conidia counts were sufficient to significantly reduce the infectivity of the diseased plant. Secondary infections caused by UV-B pre-treated plants exhibited yet further (67%) reduced disease severity. UV-B-induced flavonoids may in part mediate this reduced disease severity phenotype, as B. lactucae conidia counts of lettuce plants negatively correlated with flavonoid levels in a UV-B-dependent manner (r = −0.81). Liquid chromatography–mass spectrometry (LC-MS) was used to identify metabolic features which contribute to this correlation and, of these, quercetin 3-O-(6”-O-malonyl)-b-D-glucoside had the strongest negative correlation with B. lactucae conidia count (r = −0.68). When quercetin 3-O-(6”-O-malonyl)-b-D-glucoside was directly infiltrated into lettuce leaves, with those leaves subsequently infected, the B. lactucae conidia count was reduced (25–39%) in two susceptible lettuce cultivars. We conclude that UV-B induced phenolics, in particular quercetin flavonoids, may act as phytoanticipins to limit the establishment of biotrophic pathogens thus delaying or reducing their sporulation as measured by conidia count. These findings highlight the opportunity for UV-B morphogenesis to be exploited through the application of UV-LED technology, as part of the development of next-generation, sustainable disease control tools.

Effect of Planting Date and Inoculum Density on Severity of Fusarium Wilt of Lettuce in California.

Seven lettuce cultivars were transplanted into a field infested with Fusarium oxysporum f. sp. lactucae (causal agent of Fusarium wilt of lettuce) in August and September of 2014, 2015, and 2016. For moderately susceptible cultivars, 1- or 2-week differences in planting date had a significant effect on severity of Fusarium wilt. In growth chamber experiments, cultivars of moderate susceptibility were maintained in a growth chamber under cool conditions (23/18°C), and a subset of plants was transferred to a warm chamber (33/23°C) for 1 week, at weekly intervals after transplanting. Plants exposed to high temperatures at 2 and 3 weeks after transplanting (WAT) had more severe symptoms of Fusarium wilt than those exposed at 4 WAT. In October 2015, April 2016, August 2016, and August 2017, moderately susceptible cultivars were planted into field soil with an inoculum density gradient. Moderately susceptible cultivars were little affected by Fusarium wilt where inoculum densities of F. oxysporum f. sp. lactucae were <125 per gram of soil, even in warm planting windows. Adjusting planting dates to avoid high temperatures during a critical stage of growth and maintaining low inoculum density in soil can contribute to management of Fusarium wilt in moderately susceptible lettuce cultivars.

The effect of co-infestation by conspecific and heterospecific aphids on the feeding behaviour of Nasonovia ribisnigri on resistant and susceptible lettuce cultivars

Aphid saliva can suppress the blocking of sieve elements, a reaction that plants employ to inhibit aphid feeding, but aphid saliva can also elicit plant defence responses. Such plant responses might affect interactions between different aphid species and intraspecifically, e.g. among different biotypes. The objectives of our study were to investigate if feeding behaviour and performance of two biotypes of the lettuce aphid Nasonovia ribisnigri are affected by (1) feeding by the other biotype and (2) feeding by the green peach aphid Myzus persicae or the potato aphid Macrosiphum euphorbiae. Additionally the effect of feeding in a group was studied. All experiments were performed on both a resistant and an isogenic susceptible lettuce cultivar. Feeding or probing by conspecific or heterospecific aphids had different effects on Nasonovia ribisnigri biotypes. Aphids were only slightly affected by feeding or probing of the same biotype on both susceptible and resistant lettuce. N. ribisnigri virulent biotype Nr:1 suppressed the resistance against Nr:0 in the resistant cultivar. In contrast, defence was induced by Nr:1 against Nr:0 in susceptible lettuce. Co-infestation by M. euphorbiae and M. persicae had minor effects on Nr:0. Defence against Nr:1 was induced on both susceptible lettuce and resistant lettuce by Nr:0 and M. euphorbiae. Additionally, M. persicae induced defence in resistant lettuce against Nr:1. Effectors in the saliva of Nr:1 aphids are likely responsible for the defence suppression in lettuce. Identification of these effectors could lead to a better understanding of the mechanism of virulence in N. ribisnigri.

A model for multiseasonal spread of verticillium wilt of lettuce.

Verticillium wilt, caused by Verticillium dahliae, is a destructive disease in lettuce, and the pathogen is seedborne. Even though maximum seed infestation rates of <5% have been detected in commercial lettuce seed lots, it is necessary to establish acceptable contamination thresholds to prevent introduction and establishment of the pathogen in lettuce production fields. However, introduction of inoculum into lettuce fields for experimental purposes to determine its long term effects is undesirable. Therefore, we constructed a simulation model to study the spread of Verticillium wilt following pathogen introduction from seed. The model consists of four components: the first for simulating infection of host plants, the second for simulating reproduction of microsclerotia on diseased plants, the third for simulating the survival of microsclerotia, and the fourth for simulating the dispersal of microsclerotia. The simulation results demonstrated that the inoculum density-disease incidence curve parameters and the dispersal gradients affect disease spread in the field. Although a steep dispersal gradient facilitated the establishment of the disease in a new field with a low inoculum density, a long-tail gradient allowed microsclerotia to be dispersed over greater distances, promoting the disease spread in fields with high inoculum density. The simulation results also revealed the importance of avoiding successive lettuce crops in the same field, reducing survival rate of microsclerotia between crops, and the need for breeding resistance against V. dahliae in lettuce cultivars to lower the number of microsclerotia formed on each diseased plant. The simulation results, however, suggested that, even with a low seed infestation rate, the pathogen would eventually become established if susceptible lettuce cultivars were grown consecutively in the same field for many years. A threshold for seed infestation can be established only when two of the three drivers of the disease-(i) low microsclerotia production per diseased plant, (ii) long-tail dispersal gradient, and (iii) low microsclerotia survival between lettuce crops-are present.

The Response of Lettuce Cultivars against TwoLettuce mosaic virusisolates, One Able to Systemically Infect the Resistant Cultivar Salinas 88

The response of seven lettuce cultivars to two geographically different Lettuce mosaic virus (LMV) isolates (LMV-A, LMV-T) was statistically evaluated based on infection rate, virus accumulation and symptom severity in different time trials. LMV-A is characterized by the ability to systemically infect cv. Salinas 88 (mo12-carrying resistant cultivar), and inducing mild mosaic symptoms. Among lettuce cultivars, Varamin (a native cultivar) similar to cv. Salinas showed the most susceptibility to both LMV isolates, whereas another native cultivar, Varesh, was tolerant to the virus with minimal viral accumulation and symptom scores, significantly different from other cultivars at P < 0.05. LMV-A systemically infects all susceptible lettuce cultivars more rapidly and at a higher rate than LMV-T. This isolate accumulated in lettuce cultivars at a significantly higher level, determined by semiquantitative ELISA and induced more severe symptoms than LMV-T isolate at 21 dpi. This is the first evidence for a LMV isolate with ability to systemically infect mo12-carrying resistant cultivar of lettuce from Iran. In this study, accumulation level of LMV showed statistically meaningful positive correlation with symptom severity on lettuce plants. Based on the results, three evaluated parameters differed considerably by lettuce cultivar and virus isolate.

Colonization of resistant and susceptible lettuce cultivars by a green fluorescent protein-tagged isolate of Verticillium dahliae.

Interactions between lettuce and a green fluorescent protein (GFP)-expressing, race 1 isolate of Verticillium dahliae, were studied to determine infection and colonization of lettuce cultivars resistant and susceptible to Verticillium wilt. The roots of lettuce seedlings were inoculated with a conidial suspension of the GFP-expressing isolate. Colonization was studied with the aid of laser scanning confocal and epi-fluorescence microscopes. Few differences in the initial infection and colonization of lateral roots were observed between resistant and susceptible cultivars. Hyphal colonies formed on root tips and within the root elongation zones by 5 days, leading to the colonization of cortical tissues and penetration of vascular elements regardless of the lettuce cultivar by 2 weeks. By 8 to 10 weeks after inoculation, vascular discoloration developed within the taproot and crown regions of susceptible cultivars well in advance of V. dahliae colonization. Actual foliar wilt coincided with the colonization of the taproot and crown areas and the eruption of mycelia into surrounding cortical tissues. Advance colonization of stems, pedicels, and inflorescence, including developing capitula and mature achenes was observed. Seedborne infection was limited to the maternal tissues of the achene, including the pappus, pericarp, integument, and endosperm; but the embryo was never compromised. Resistant lettuce cultivars remained free of disease symptoms. Furthermore, V. dahliae colonization never progressed beyond infected lateral roots of resistant cultivars. Results indicated that resistance in lettuce may lie with the plant's ability to shed infected lateral roots or to inhibit the systemic progress of the fungus through vascular tissues into the taproot.

Prevalence of Lettuce mosaic virus - common strain on three lettuce producing areas from São Paulo State

LMV is one of the most important pathogens of lettuce worldwide. Based on their ability to overcome the resistance genes mo1¹ and mo1² in lettuce, isolates can be divided in two types: LMV-Most, which can infect and are seed-borne in cultivars containing the mo1 gene and LMV-Common, which do not cause symptoms on these cultivars and are seed transmitted only in susceptible cultivars. To evaluate the occurrence of these two types of LMV isolates, a survey was carried out during 2002-2005 in three lettuce production areas from São Paulo State. Total RNA was used for the diagnosis of LMV isolates by RT-PCR using universal primers for the variable N-terminus of the capsid protein, in the 3' end of the genome. Positives samples were analyzed by a second RT-PCR using specifics primers for LMV-Most isolates designed to amplify a fragment from the central region (CI-VPg) of the genome. A total of 1362 samples showing mosaic symptoms were collected and 504 (37.29 %) were positives for LMV. On susceptible lettuce cultivars, LMV-Common was prevalent (77.3%). LMV-Most was found frequently associated with tolerant (mo1¹) lettuce cultivars. Susceptible cultivars correspond today for most of the area of lettuce production. So, despite the ability of LMV-Most isolates to overcome the resistance provided by the recessive mo1¹ gene, they are not prevalent in the conditions of São Paulo State.

Lettuce mosaic virus: from pathogen diversity to host interactors.

Lettuce mosaic virus (LMV) belongs to the genus Potyvirus (type species Potato virus Y) in the family Potyviridae. The virion is filamentous, flexuous with a length of 750 nm and a width of 15 nm. The particles are made of a genomic RNA of 10 080 nucleotides, covalently linked to a viral-encoded protein (the VPg) at the 5' end and with a 3' poly A tail, and encapsidated in a single type of capsid protein. The molecular weight of the capsid protein subunit has been estimated electrophoretically to be 34 kDa and estimated from the amino acid sequence to be 31 kDa. The genome is expressed as a polyprotein of 3255 amino-acid residues, processed by three virus-specific proteinases into ten mature proteins. LMV has a worldwide distribution and a relatively broad host range among several families. Weeds and ornamentals can act as local reservoirs for lettuce crops. In particular, many species within the family Asteraceae are susceptible to LMV, including cultivated and ornamental species such as common (Lactuca sativa), prickly (L. serriola) or wild (L. virosa) lettuce, endive/escarole (Cichorium endiva), safflower (Carthamus tinctorius), starthistle (Centaurea solstitialis), Cape daisy (Osteospermum spp.) and gazania (Gazania rigens). In addition, several species within the families Brassicaceae, Cucurbitaceae, Fabaceae, Solanaceae and Chenopodiaceae are natural or experimental hosts of LMV. Genetic control of resistance to LMV: The only resistance genes currently used to protect lettuce crops worldwide are the recessive genes mo1(1) and mo1(2) corresponding to mutant alleles of the gene encoding the translation initiation factor eIF4E in lettuce. It is believed that at least one intact copy of eIF4E must be present to ensure virus accumulation. LMV is transmitted in a non-persistent manner by a high number of aphid species. Myzus persicae and Macrosiphum euphorbiae are particularly active in disseminating this virus in the fields. LMV is also seedborne in lettuce. The effectiveness of LMV transmission depends on the cultivar and the age of the seed carrier at the inoculation time. The characteristic symptoms on susceptible lettuce cultivars are dwarfism, mosaic, distortion and yellowing of the leaves with sometimes a much reduced heart of lettuce (failure to form heads). The differences in virus strains, cultivars and the physiological stage of the host at the moment of the attack cause different symptom severity: from a very slight discoloration of the veins to severe necrosis leading to the death of the plant.

Effects of green fluorescent protein or beta-glucuronidase tagging on the accumulation and pathogenicity of a resistance-breaking Lettuce mosaic virus isolate in susceptible and resistant lettuce cultivars.

The RNA genome of a resistance-breaking isolate of Lettuce mosaic virus (LMV-E) was engineered to express the jellyfish green fluorescent protein (GFP) or beta-glucuronidase (GUS) fused to the helper-component proteinase (HC-Pro) to study LMV invasion and spread in susceptible and resistant lettuce cultivars. Virus accumulation and movement were monitored by either histochemical GUS assays or detection of GFP fluorescence under UV light. The GFP- and GUS-tagged viruses spread systemically in the susceptible lettuce cultivars Trocadero and Vanguard, where they induced attenuated symptoms, compared with the wild-type virus. Accumulation of the GFP-tagged virus was reduced but less affected than in the case of the GUS-tagged virus. Systemic movement of both recombinant viruses was very severely affected in Vanguard 75, a lettuce cultivar nearly isogenic to Vanguard but carrying the resistance gene mo1(2). Accumulation of the recombinant viruses in systemically infected leaves was either undetectable (GUS-tag) or erratic, strongly delayed, and inhibited by as much as 90% (GFP-tag). As a consequence, and contrary to the parental virus, the recombinant viruses were not able to overcome the protection afforded by the mo1(2) gene. Taken together, these results indicate that GUS or GFP tagging of the HC-Pro of LMV has significant negative effects on the biology of the virus, abolishing its resistance-breaking properties and reducing its pathogenicity in susceptible cultivars.

Yield Losses to Iceberg Lettuce Due to Corky Root Caused byRhizomonas suberifaciens

Plant growth and yield variables of closely related resistant and susceptible lettuce cultivars (440-8 and Salinas, respectively) were compared in microplots at Davis, CA, in soil infested with Rhizomonas suberifaciens, the causal agent of corky root of lettuce. To validate the results for large scale lettuce production, the same variables were compared for 440-8 and Salinas in fumigated soil and soil naturally infested with R. subenfaciens at Salinas, CA. Most plant growth variables for 440-8 in infested microplots were not significantly different from those for 440-8 or Salinas in uninfested microplots (...)

An Association Between High Peroxidase Activity in Lettuce (Lactuca sativa) and Field Resistance to Downy Mildew (Bremia lactucae)

AbstractThe association between variation for pre‐infection peroxidase activity and levels of field resistance‐susceptibility to downy mildew (Bremia lactucae) was investigated in lettuce (Lactuca sativa) cultivars, accessions of L. serriola (prickly lettuce), segregating F2 populations and selected F3 families from a cross between field resistant and susceptible lettuce cultivars. A trend was apparent in this series of experiments indicating that one component of field resistance could be related to a high level of peroxidase activity prior to infection. The data suggest that in breeding programmes there could be merit in imposing primary selection for high peroxidase activity prior to field selection for resistance.

Further work on the genetics of race specific resistance in lettuce (Lactuca sativa) to downy mildew (Bremia lactucae)

SUMMARYData are presented on the segregation of resistance to four races of Bremia lactucae in the F2 progenies of crosses involving 15 resistant and various susceptible lettuce cultivars. Most of these data and those recently published by other workers fit the systematic model for the genetics of race specific resistance to B. lactucae proposed by Crute &amp; Johnson(1976).Seven different dominant resistance genes of major effect were found. There was also evidence of a pair of dominant genes with complementary effect, one of which may sometimes be effective on its own. Two of the genes may be linked and another may have two different alleles for resistance at the same locus. The resistance conferred by these genes is specified in relation to two British, two Dutch and four French races of the fungus. Resistance genotypes are proposed for 16 cultivars.

The genetics of race specific resistance in lettuce (Lactuca sativa) to downy mildew (Bremia lactucae)

SUMMARYData are presented on the segregation of resistance to five British races and two Dutch races of Bremia lactucae in the F2 progenies of crosses involving seven resistant and several susceptible lettuce cultivars. These data and also those previously published by other workers are considered in relation to the systematic model proposed by Crute &amp; Johnson (1976) to explain the genetics of race specific resistance to B. lactucae in lettuce. It is shown that, with minor modifications, the model accommodates almost all of the previously published data and correctly predicts the new data, except for one set which cannot at present be interpreted.It is concluded that genetic evidence exists for the presence, among various cultivars of lettuce, of at least four and possibly five different dominant resistance genes of major effect designated Dm2, Dm3, Dm4, Dm6 and Dm8; and of a pair of dominant genes with complementary effect designated Dm7/1 and Dm7/2. The resistance conferred by these genes is specified in relation to five British races, five Dutch, three Israeli and one United States race of the fungus. Resistance genotypes are proposed for cultivars Avoncrisp, Avondefiance, Calmar, Great Lakes 659, Kares, Meikoningen, Mildura, Proeftuins Blackpool, Solito, Valverde, Ventura and the USDA line PI 164937.

The influence of seedling age and development on the infection of lettuce by Bremia lactucae

SUMMARYLaboratory methods are described for the culture and observation of Bremia lactucae on lettuce.Mycelial growth of the fungus within the cotyledons of the cultivar Cheshunt Early Giant progressed to a minimum density at which sporulation occurred, and then continued to a maximum density coinciding with peak sporulation. The timing and intensity of sporulation was influenced by the concentration of the inoculum. Seedlings often became systemically infected and this, together with intensity of sporulation, was a good indicator of fungal growth and hence host susceptibility.The severity of disease development on four susceptible lettuce cultivars (Cheshunt Early Giant, Paris White, Trocadero Improved and Crackling Ice) decreased with age and development of the seedlings at inoculation. The reasons for this are discussed.