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ABSTRACT OPPENHEIM, SARA JANE. the Genetic Basis of Host

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ABSTRACT OPPENHEIM, SARA JANE. the Genetic Basis of Host ABSTRACT OPPENHEIM, SARA JANE. The Genetic Basis of Host Plant Use in a Specialist and a Generalist Moth. (Under the direction of Fred L Gould.) Differences in host plant use between Heliothis subflexa and H. virescens provide a compelling example of closely-related species with strikingly divergent ecological adaptations. I used a QTL mapping approach to investigate the genetic basis of interspecific differences between H. subflexa and H. virescens in use of Physalis angulata, a host of H. subflexa but not H. virescens. I introgressed H. subflexa genes into the H. virescens background by backcrossing interspecific hybrids to H. virescens, and examined the effects of QTL from H. subflexa on use of P. angulata. Many QTL had different size effects, and sometimes even different direction of effects, in males and females. Many QTL differed among lineages, but this variation probably reflects a lack of power to detect QTL rather than true differences in QTL effects. Size, significance, and direction of effects of some QTL varied among families within a lineage. The same Hs-like phenotype could be produced by introgressing different H. subflexa chromosomes into the H. virescens background. Sample size was increased almost four-fold from 2001 to 2007, and the increase in the number of QTL detected almost exactly matched the increase in sample size. QTL effect sizes were small, when measured as percent of backcross variation explained, but were larger when considered as the percent of interspecific difference explained. QTL effect sizes were consistent across traits. For most traits, individual QTL accounted for 10 to 40 percent of the interspecific difference, while backcross variation explained ranged from 0.5 to 6 percent. Most of the QTL affected multiple traits, and these effects were usually in the same direction. I did experiments on the response to selection within H. virescens for use of P. angulata and on mapped QTL in backcrosses of H. virescens control and selection lines. Selection resulted not only in increased survival on Physalis, but also in greater tendency to feed, greater larval weight gain, and greater assimilation efficiency on Physalis. The same phenotype could be produced by different QTL. Many QTL distributed across about half of the chromosomes affected the traits measured, and some QTL affected more than one trait. The effects of QTL appeared to act additively or dominantly so that trait values for backcross progeny, which had at most one copy of a given selection-line allele, were intermediate between in phenotype. Most QTL explained 5-10 percent of variation among backcross progeny, and phenotypes that resembled selection line H. virescens or H. subflexa were produced when several selection line QTL were inherited together. Each QTL explained a much greater amount of the difference between control H. virescens and selection line H. virescens or H. subflexa because the differences in the means between species and between control and selections lines were less than the variation among backcross progeny. For assimilation efficiency, the number of chromosomes harboring QTL associated with phenotypic variation was remarkable similar within versus between species: I found 11 chromosomes associated with interspecific differences between H. subflexa and H. virescens, and 12 chromosomes associated with intraspecific differences within H. virescens. Some QTL had effects in the “wrong” direction: 3 interspecific and 6 intraspecific chromosomes had QTL associated with phenotypes that were Hv-like rather than Hs-like. For proportion of larvae feeding on Physalis, the number of chromosomes with QTL was again similar, but the pattern of effects was quite different. In the interspecific crosses, most chromosomes that harbored QTL had Hs-like effects, but in the intraspecific crosses, half of the chromosomes had QTL with Hv-like effects. Selection for adaptation to Physalis in H. virescens may have been accompanied by an overall decrease in willingness to feed that is unrelated to the particular host plant but reflects a typical specialist phenotype. The Genetic Basis of Host Plant Use in a Specialist and a Generalist Moth by Sara Jane Oppenheim A dissertation submitted to the Graduate Faculty of North Carolina State University in partial fulfillment of the requirements for the Degree of Doctor of Philosophy Entomology Raleigh, North Carolina 2010 APPROVED BY: George Kennedy Coby Schal Trudy Mackay Fred Gould Chair of Advisory Committee DEDICATION To Keith and Anna-Claire. ii BIOGRAPHY Sara was born in Raleigh, North Carolina. During her college career, she studied evolutionary biology at the Eugene Lang College of The New School for Social Research in New York City, philosophy at Smith College in Northampton, MA, and biology at Wake Forest University in Winston-Salem, NC, from which she graduated with a Bachelor of Science degree. While at Wake Forest, she studied under William Connor, who first introduced her to the many wonders of insects. After finishing her Bachelor’s degree, Sara spent six months in Southern Pines, NC as a working student in a Combined Training stable, during which time she and her horse competed in events in NC. After this break from academia, Sara began her graduate career at NC State University in the laboratory of Fred Gould. After receiving her Master’s degree in 2000, Sara continued to work in Fred’s lab doing research toward her Ph. D. In 2010, she completed her Ph. D. iii TABLE OF CONTENTS LIST OF TABLES.................................................................................................................. vii LIST OF FIGURES .................................................................................................................ix Chapter 1: Introduction............................................................................................................. 1 Adaptation and Diversification........................................................................................ 1 Significance of Research.................................................................................................. 7 Chapter 2: Literature Review....................................................................................................9 Introduction...................................................................................................................... 9 The Genetics of Host Specificity ................................................................................... 11 Heliothis ................................................................................................................. 12 Helicoverpa ............................................................................................................ 16 Papilio..................................................................................................................... 17 Euphydryas............................................................................................................. 20 Other systems ......................................................................................................... 22 Conclusions ............................................................................................................ 23 Integration of Larval and Adult Traits........................................................................... 24 Neurobiology of Host Range ......................................................................................... 27 Directionality of Host Range Evolution ........................................................................ 31 Future Prospects............................................................................................................. 33 Chapter 3: The genetic basis of Physalis use in H. subflexa .................................................. 37 Introduction.................................................................................................................... 37 Materials and Methods................................................................................................... 42 Study system........................................................................................................... 42 Insect strains and rearing........................................................................................ 44 Plants ...................................................................................................................... 44 Backcross matings.................................................................................................. 45 Measurement of Larval Phenotypes....................................................................... 46 Experimental Controls............................................................................................ 47 DNA Extraction...................................................................................................... 47 AFLP Markers........................................................................................................ 48 Gene-Based Markers .............................................................................................. 50 Linkage Mapping ................................................................................................... 51 iv QTL Mapping......................................................................................................... 52 Results...........................................................................................................................
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