Guillermo Sotelo

Screening for Resistance to Adult Spittlebugs (Hemiptera: Cercopidae) in Brachiaria spp.: Methods and Categories of Resistance

ABSTRACT Both nymphal and adult stages of several species of spittlebugs (Hemiptera: Cercopidae) are key economic pests of brachiariagrasses (Brachiaria spp.) in tropical America. Progress has been made in the characterization and development of antibiosis resistance to nymphs in brachiariagrasses. Essentially no attention has been given to screening germplasm for resistance to adults. To support current breeding programs, a series of experiments was conducted to develop a methodology to screen for adult damage and to study categories of resistance to adult feeding damage. Six host brachiariagrass genotypes were used: two susceptible checks (CIAT 0606 and CIAT 0654) and four nymph-resistant genotypes (CIAT 6294, CIAT 36062, CIAT 36087, and SX01NO/0102). Test insects were Aeneolamia varia (F.) and Zulia carbonaria (Lallemand). None of the nymph-resistant genotypes was antibiotic to adults. All four nymph-resistant genotypes showed tolerance to A. varia and Z. carbonaria feeding damage. The levels of tolerance to adults of Z. carbonaria, a larger, more aggressive species, were lower. Of the four nymph-resistant genotypes, only CIAT 6294 and CIAT 36087 showed some tolerance to Z. carbonaria expressed as lower leaf damage scores, less chlorophyll loss, and lower functional plant loss indices. The fact that a genotype like SX01NO/0102, which is highly antibiotic to nymphs, is susceptible to adult damage suggests that mechanisms of resistance to the two spittlebug life stages may be independent. Results of these studies suggest a need to incorporate routine screening for tolerance to adult feeding damage as an additional selection criterion in the breeding scheme.

Independence of Resistance in Brachiaria spp. to Nymphs or to Adult Spittlebugs (Hemiptera: Cercopidae): Implications for Breeding for Resistance

ABSTRACT Both nymphal and adult spittlebugs (Hemiptera: Cercopidae) cause serious economic damage to susceptible brachiariagrass [genus Biachiaria (Trin.) Griseb], pastures in tropical America. Both life stages are xylem feeders: nymphs feed primarily on roots and stems, whereas the adults feed mainly on foliage. Numerous interspecific brachiariagrass hybrids with high levels of antibiosis resistance to nymphs of several important spittlebug species have been obtained. Recent studies revealed major inconsistencies between reaction to nymphs and reaction to adults on the same host genotype. Because both insect life stages can cause severe economic damage on susceptible brachiariagrass pastures, a cultivar development strategy must take into account resistance to both life stages. To assess the degree of association between resistance to spittlebug nymphs and to adult feeding, we tested 164 hybrids and six check genotypes for resistance to both life stages of three spittlebug species: Aeneolamia varia (F.), Aeneolamia reducta (Lallemand), and Zulia carbonaria (Lallemand). Most hybrids tested were classified as resistant to nymphs. On the contrary, for all three species, the overall mean damage score of the 164 hybrids did not differ from the mean score of the susceptible checks. None of the hybrids was classified as resistant to adult feeding damage. Correlations between percentage nymph survival and adult damage scores were consistently low (r = 0.0104–0.0191). Correlations between nymphal and adult damage scores were also low (0.109–0.271), suggesting that resistances to the different life stages are largely independent. Chi-square analyses comparing frequency distributions of responses of the 164 breeding hybrids to nymphs or adults confirmed essential genetic independence of these two traits. We conclude that attention to improving genetic resistance specifically to adult feeding damage is warranted.

Response of Resistant and Susceptible Brachiaria spp. Genotypes to Simultaneous Infestation with Multiple Species of Spittlebugs (Hemiptera: Cercopidae)

Abstract The response of one susceptible and three resistant Brachiaria spp. (Hemiptera: Cercopidae) genotypes to individual or combined attacks by nymphs of Aeneolamia varia (F.), Aeneolamia reducta (Lallemand), Zulia carbonaria (Lallemand), and Zulia pubescens (F.) was studied. We assessed the effect of infesting plants of the susceptible check BRX 44–02 and of the A. varia-resistant genotypes CIAT 6294 and CIAT 36062 with A. varia, Z. carbonaria, or Z. pubescens either alone or in two-species combinations. In a second trial, we studied the performance of BRX 44-02, CIAT 6294, and the multiple resistant clone SX01NO/0102 exposed to individual or combined attack by A. reducta and Z. carbonaria. In a third trial, we compared the response of BRX 44-02, CIAT 6294, and CIAT 36062 to individual A. varia, Z. carbonaria, or Z. pubescens attack as opposed to a combined three-species attack. Plant damage scores and percentage of nymphal survival were recorded in all three trials. Data on percentage of survival indicated that competition between and among spittlebug species occurs. However, we found no evidence of interaction between species competition and different levels of resistance to spittlebug. Rather, host genotype reactions conformed to previously known categories of resistance regardless of the presence of more than one spittlebug species. Resistance rather than competition seems to have been the overriding factor determining nymph survival and resistance expression (damage scores) in these experiments. Our results corroborate the need to develop brachiariagrass genotypes with multiple resistance to spittlebugs.

Sublethal effects of antibiosis resistance on the reproductive biology of two spittlebug (Hemiptera: Cercopidae) species affecting Brachiaria spp.

Abstract Several greenhouse experiments were used to measure how high levels of antibiosis resistance to nymphs in two interspecific Brachiaria (brachiariagrass) hybrids affect life history parameters of the spittlebugs Aeneolamia varia (F.) and Zulia carbonaria (Lallemand), two of the most important spittlebug (Hemiptera: Cercopidae) species affecting Brachiaria production in Colombia. The A. varia-resistant hybrid CIAT 36062, the Z. carbonaria-resistant hybrid SX01NO/0102, and the susceptible accession CIAT 0654 were used to compare the effect of all possible combinations of food sources for nymphs and adults. Calculation of growth indexes showed a significant impact of antibiosis resistance on the biology of immature stages of both species. Median survival times of adults feeding on resistant genotypes did not differ from those recorded on the susceptible genotype, suggesting that factors responsible for high mortality of nymphs in the resistant hybrids did not affect adult survival. Rearing nymphs of A. varia on CIAT 36062 and of Z. carbonaria on SX01NO/0102 had deleterious sublethal effects on the reproductive biology of resulting adult females. It is concluded that high nymphal mortality and subsequent sublethal effects of nymphal antibiosis on adults should have a major impact on the demography of the two spittlebug species studied.

Characterization of Resistance to Adult Spittlebugs (Hemiptera: Cercopidae) in Brachiaria spp.

ABSTRACT Nymphs and adults of several spittlebug (Hemiptera: Cercopidae) species are key pests of forage brachiariagrasses (Brachiaria spp.) in tropical America. To support current breeding programs, a series of experiments aimed at characterizing the mechanisms of resistance to adult feeding damage were carried out. Five genotypes were used: two susceptible checks (CIAT 0606 and CIAT 0654) and three nymph-resistant genotypes (CIAT 36087, CIAT 6294, and SX01NO/0102). Test insects were Aeneolamia varia (F.), A. reducta (Lallemand), and Zulia carbonaria (Lallemand). The nymphresistant genotypes showed tolerance to all spittlebug species tested. Tolerance in these genotypes can be classified as only moderate given the extent of losses (60–80%) caused by both female and male adults. None of the nymph-resistant genotypes had antibiotic effects on adults feeding on foliage. The results also indicated that antixenosis for feeding is not a plausible explanation for lower damage scores and less biomass losses in resistant genotypes. The fact that adult longevity (usually 8 d) was not affected when the adults were forced to feed on roots of a genotype with strong antibiotic resistance to nymphs is regarded as additional evidence that resistances to nymphs and to adults in Brachiaria are largely independent.

Characterizing herbivore resistance mechanisms: spittlebugs on Brachiaria spp. as an example.

Plants can resist herbivore damage through three broad mechanisms: antixenosis, antibiosis and tolerance1. Antixenosis is the degree to which the plant is avoided when the herbivore is able to select other plants2. Antibiosis is the degree to which the plant affects the fitness of the herbivore feeding on it1.Tolerance is the degree to which the plant can withstand or repair damage caused by the herbivore, without compromising the herbivores growth and reproduction1. The durability of herbivore resistance in an agricultural setting depends to a great extent on the resistance mechanism favored during crop breeding efforts3. We demonstrate a no-choice experiment designed to estimate the relative contributions of antibiosis and tolerance to spittlebug resistance in Brachiaria spp. Several species of African grasses of the genus Brachiaria are valuable forage and pasture plants in the Neotropics, but they can be severely challenged by several native species of spittlebugs (Hemiptera: Cercopidae)4.To assess their resistance to spittlebugs, plants are vegetatively-propagated by stem cuttings and allowed to grow for approximately one month, allowing the growth of superficial roots on which spittlebugs can feed. At that point, each test plant is individually challenged with six spittlebug eggs near hatching. Infestations are allowed to progress for one month before evaluating plant damage and insect survival. Scoring plant damage provides an estimate of tolerance while scoring insect survival provides an estimate of antibiosis. This protocol has facilitated our plant breeding objective to enhance spittlebug resistance in commercial brachiariagrases5.

Phenotyping Brachiaria Genotypes to Assess Rhizoctonia Resistance by Comparing Three Inoculum Types

Rhizoctonia foliar blight, caused by Rhizoctonia solani, is an important disease of Brachiaria spp. in tropical America. Host-plant resistance is an attractive option for disease management. In this study, we evaluated three inoculum types (mycelium-infected agar disc, microdiscs suspensions, and microencapsulated-mycelium suspensions) in order to identify a rapid and accurate method for mass screening of Brachiaria genotypes for resistance to Rhizoctonia spp. in greenhouse trials. Visual damage score, area under the disease progress curve, and percent chlorophyll loss were estimated to determine the most accurate and precise method for evaluating Rhizoctonia resistance. The microencapsulated-mycelium solution (0.75 g/ml in potato dextrose broth sprayed on plants 30 days after planting) caused greater foliar damage than the other inoculum types and allowed effective discrimination between resistant and susceptible genotypes. The effectiveness of spray-applied, microencapsulated-mycelium was further corroborated by the evaluation of 350 genotypes not previously selected for resistance to Rhizoctonia spp., which varied significantly in their reaction to R. solani. The microencapsulated-mycelium methodology has several advantages over existing methods, including its high-throughput capacity, efficient use of time and space, ease of quantification of inoculum, and consistent results over replicate trials. This methodology could be applied to assess resistance to Rhizoctonia spp. in other crops.