Abstract
This thesis examines the three-way interaction between yeast-like symbionts, an insect herbivore [Nilaparvata lugens (Stal)] and its rice (Oryza sativa L.) host, during adaptation of the herbivore to resistant rice varieties. A long-term selection study (20 generations of continuous rearing, ca. 24 months) was conducted with N. lugens populations on four rice varieties (IR22, a susceptible variety and IR65482, IR62, and PTB33, three resistant varieties). Planthopper performance and the abundance of yeast-like symbionts (YLS) were monitored throughout the selection process. N. lugens populations adapted to the resistant varieties as noted by increasing body size and increased egglaying. Xylem feeding was observed as a possible behavioural adaptation of N. lugens: planthoppers on resistant plants had relatively high levels of xylem feeding compared with planthoppers on susceptible plants. Planthoppers selected on resistant varieties, had clear differences in YLS densities that were not related to fitness on the varieties and, therefore, did not support a YLS density-mediated adaptation hypothesis. Furthermore, this study examined whether YLS density affected the capacity of planthoppers to switch between hosts on which they have been selected for several generations (natal plant) to new varieties (exposed plants) under normal YLS densities (symbiotic) and after reduction of YLS densities by heat treatment (aposymbiotic). The results suggested that YLS do not mediate host plant switching in planthoppers as removal of symbionts influenced body weight but not the relative capacity of nymphs to feed on different plants. This study also tested if virulence is acquired by shared feeding sites with virulent and avirulent planthoppers. In the study, planthoppers with varying levels of virulence affected the host plants differently: The most virulent hoppers appeared to suppress rice defences to a greater extent than non-virulent planthoppers. Planthoppers attained highest weights on those plants on which virulent planthoppers had previously fed which suggests that feeding by the virulent planthoppers facilitated subsequent planthopper feeding on the same plant. Our preliminary results indicate that feeding by mixed virulent-avirulent populations could potentially accelerate adaptation by N. lugens to resistant rice varieties. The capacity of virulent and avirulent planthoppers to feed on a range of 24 resistant rice varieties was examined using a series of bioassays. Planthoppers were observed to feed and lay eggs on all the varieties tested, many of which have never been widely deployed in the field. Furthermore, planthoppers selected on resistant varieties often had increased fitness on other resistant varieties, even when these possess different resistance genes. However, there was no strong evidence that once planthoppers have adapted to a resistant variety, they will exhibit fitness costs on other varieties with dissimilar genes. The mechanisms underlying insect virulence are complex and further research on planthopper adaptation is necessary to help conserve genetic resources and prolong the durability of available resistant varieties.
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