Polyploidy and host specificity genetics in Nasonia parasitoid wasps Leung, Kelley
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10.33612/diss.134432017
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Publication date: 2020
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Leung, K. (2020). Polyploidy and host specificity genetics in Nasonia parasitoid wasps. University of Groningen. https://doi.org/10.33612/diss.134432017
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Summary
This PhD project was part of Breeding Invertebrates for Next Generation Biological Control (BINGO), a Marie-Skłodowska Curie Initial Training Network to improve biological control. Biological control is the practice of using whole organisms, mostly natural enemies of the target species, to control agricultural pests. It is increasing in importance as a means of crop protection, as governing bodies also strive to decrease pesticide use. There are many aspects to the improvement of biological control, one being breeding better biological control agents, many of which are parasitic wasps. The genetics of parasitic wasps includes the phenomenon of polyploidy, which will be explained below, that opens up interesting possibilities for improving biocontrol stocks. Another target for improvement is host specificity that may be optimized for the target species. These objectives originated from the goal of improving biological control and expanded to include elements of evolutionary theory. Parasitoid wasps are within the insect order Hymenoptera, which comprise the wasps, bees, sawflies, and ants. All hymenopterans have haplodiploid sex determination: unfertilized (haploid) eggs, with only one chromosome set (from the mother) develop into males and fertilized (diploid) eggs, with two sets chromosomes, develop into females. However, in many species, polyploids (individuals with a higher number than one or two chromosome sets) appear. Polyploidization often occurs for species with the common hymenopteran sex determination system, Complementary Sex Determination (CSD). In this system, only diploid eggs with two different variants (alleles) of a sex-determining gene develop as females. In CSD species, inbreeding causes the production of diploid eggs with similar alleles for this gene, leading to sterile, diploid males, which increase in incidence until the population goes extinct. This phenomenon, called “the diploid male vortex,” has resulted in polyploidy’s reputation as being problematic to biological control, because maintaining control stocks will eventually lead to inbreeding without costly and time-consuming breeding regimes. Ironically, most parasitoid wasps, the most prominent class in arthropod biological control, do not have CSD, and inbreeding does not cause a “diploid male vortex”. Therefore, despite the assumption that polyploidy is disadvantageous, how polyploidy actually impacts life history traits and downstream biocontrol performance of non-CSD parasitoids has not been known. The only non-CSD parasitoids that have a well-characterized sex determination system are the blowfly parasitoids of Nasonia, which have Maternal Effect Genomic Imprinting Sex Determination (MEGISD). Within MEGISD, inbreeding does not cause polyploidy, but polyploids (diploid males, triploid females) appeared spontaneously in lab stocks of N. vitripennis and were used to derive the Whiting Polyploid Line (WPL). The WPL has been maintained for many years now and is used as a resource for genetic research for decades. Through these studies using the WPL it is known that the diploid males have high fertility (atypical for diploid male hymenopterans) and a high degree of failed chromosome distribution (aneuploidy, inviablegamete production) in triploid females. However, not much else was known about the WPL’s life history, including traits most relevant to biocontrol performance. The WPL served as a starting point for assessing the effects on polyploidy on the life history non-CSD parasitoids (Chapter 2). Both the inbred line itself and individuals outcrossed for a single generation were studied. Lifespan under starvation and fed conditions were not found to be significantly different for polyploids and non-polyploids, with the exception of outbred females (although triploids lived longer under starvation conditions and diploids lived longer under fed conditions). Haploid males and diploid males have similar reproductive capabilities. They produce the same number of offspring and have the same ability to compete for single and multiple female mates. The latter result is consistent with the trend across Hymenoptera; indeed, the inability of females to reject diploid males, which are sterile in almost all other species, is why polyploidy has been considered disadvantageous for biological control. Both inbred and outbred polyploid females had highly reduced parasitization rates, although wasps with the outbred background had higher success. This is a negative result for biological control because it directly translates to a lower rate of pest control. Cumulatively, these results indicate that polyploidy might be not as much of an impediment for non-CSD species as it is for CSD species, although traits must be carefully screened for their effect in a biocontrol context. The higher parasitization rate and longer lifespan (under starvation conditions) of outbred triploid females over inbred counterparts led to the hypothesis that negative polyploid phenotypes could be modulated with outcrossing. Therefore a new study was conducted in the genetically variable lab population, HVRremix (HVRx) (Chapters 3 and 4). It was established in previous studies that Nasonia polyploids can be generated in any background using RNAi knockdown of the sex determination gene transformer (tra). Deactivation of this maternally provided feminization factor results in diploid offspring that would have normally developed into females becoming diploid males instead, whose triploid daughters are used to found new polyploid lines. We generated a tra KD line (tKDL) in the HVRx background and similar to the WPL study, the life history traits of non-polyploid and polyploid individuals were compared. This led to an unexpected divergence from the results in the WPL. Chapter 3 focuses on broader themes in polyploid evolution. Whole genome duplication (polyploidization) is deleterious because it causes immediate problems by changing gene expression dosage, disrupting cellular interactions and causing gigantism scaling up to the organismal level, and inducing sterility. Nevertheless polyploidy is a major driving force in evolution. Evolutionary advantage emerged with polyploidization events across Eukaryota because additional gene copies allow for diversification of gene function, promoting mass speciation, increased stress resistance, and expanded geographic range. A primary topic in the field of polyploid research is resolving the paradox of how polyploidization, initially so harmful, can ultimately become highly beneficial. Among questions prompted by this poorly understood transition are, what are the immediate effects of polyploidization, what are the mechanisms by
