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Plants can resist herbivore damage through three broad mechanisms: antixenosis, antibiosis and tolerance(1). Antixenosis is the degree to which the plant is avoided when the herbivore is able to select other plants(2). Antibiosis is the degree to which the plant affects the fitness of the herbivore feeding on it(1).Tolerance is the degree to which the plant can withstand or repair damage caused by the herbivore, without compromising the herbivore's growth and reproduction(1). The durability of herbivore resistance in an agricultural setting depends to a great extent on the resistance mechanism favored during crop breeding efforts(3). 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 brachiariagrases(5).

Both nymphal and adult spittlebugs (Hemiptera: Cercopidae) cause serious economic damage to susceptible brachiariagrass [genus Brachiaria (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.

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.

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.

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.

Prosapia simulans (Walker) is an important spittlebug species that attacks forage grasses of the genus Brachiaria (Trin.) Griseb. from Mexico to Colombia. This, and several other species of spittlebugs, cause important economic losses to the livestock production industry. Development of resistant cultivars is regarded as the best method of control. In the present study we used taxonomic keys, dissection of male genitalia and RAPD-PCR patterns to reconfirm the identity of P. simulans specimens collected in Colombia and Mexico. We were able to reconfirm that P. simulans occurs as a pest of Brachiaria from Mexico to Colombia. We also studied the levels and mechanisms of resistance present in 34 Brachiaria hybrids developed by CIAT. Infestations were made with six eggs per plant. We used 10 replications (plants) per genotype in a completely randomized design. Seven hybrids were found to be susceptible, 16 showed intermediate resistance and 11 were resistant. Antibiosis was the mechanism of resistance expressed in resistant hybrids as well as in the resistant checks CIAT 6294 and CIAT 36062. Tolerance was absent. The genotypes BRX 4402 and CIAT 0606 were classified as highly susceptible.

Several genera and species of spittlebugs (Homoptera: Cercopidae) are economic pests of Brachiaria spp. grasses in tropical America. To support current breeding programs aimed at obtaining multiple spittlebug resistance, we undertook a series of studies on antibiosis and tolerance as possible mechanisms of resistance to five major spittlebug species affecting Brachiaria spp. in Colombia: Aeneolamia varia (F.), Aeneolamia reducta (Lallemand), Zulia carbonaria (Lallemand), Zulia pubescens (F.), and Mahanarva trifissa (Jacobi). Four host genotypes, well known for their reaction to A. varia attack, were used to compare their resistance to other spittlebug species: CIAT 0654 and CIAT 0606 (susceptible) and CIAT 6294 and CIAT 36062 (resistant). CIAT 0654 and CIAT 36062 were used in antibiosis studies. Tolerance studies were conducted with CIAT 0654, CIAT 6294, and CIAT 36062. Sixty-five hybrid-derived clones were used to identify levels of multiple resistance to three spittlebug species. The levels of antibiosis resistance in CIAT 36062 clearly differed by spittlebug species and were classified as follows: very high for M. trifissa, high for A. varia and A. reducta, moderate for Z. pubescens, and absent for Z. carbonaria. Our results suggest the presence of true tolerance to Z. carbonaria in CIAT 6294 and CIAT 36062, true tolerance to Z. pubescens in CIAT 6294 and a combination of tolerance and antibiosis as mechanisms of resistance to Z. pubescens in CIAT 36062. Of the 65 hybrid clones tested with A. varia, A. reducta, and Z. carbonaria, 15 combined resistance to two species and three showed antibiosis resistance to all three spittlebug species.