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Of Wing Dimorphism in the Sand Cricket, Gryllus Firm Us Heredity 65 (1990) 169—177 The Genetical Society of Great Britain Received 9 February 1990 Antagonistic pleiotropy and the evolution of wing dimorphism in the sand cricket, Gryllus firm us D. A. Roff Department of Biology, McGill University, 1205 Dr Penfield Avenue, Montreal, Quebec, Canada, H3A 1BI. At 30°C the micropterous females of the sand cricket, Gryllus firmus, begin reproduction at an earlier age after eclosion and have a larger cumulative fecundity than macropterous females. These reproductive costs may offset the advantages of being macropterous and hence capable of migration. The evolutionary significance of this phenotypic trade-off, which is characteristic of wing dimorphic insects in general, is contigent on the traits being genetically correlated. The genetic basis of the phenotypic tradeoff between flight capability and reproduction in the sand cricket, Grylius firmus, was examined by selecting for increased and decreased incidence of macroptery, and measuring the age schedules of fecundity of macropterous and micropterous females in the selected and control lines. The two traits, wing dimorphism and age schedule of reproduction, are shown to be genetically correlated. Although the mean fecundity within the selected populations changed the fecundities of macropterous and micropterous forms remained constant, suggesting that the age schedule of reproduction may itself be a threshold trait with respect to the continuously varying character controlling the expression of wing form. The relevance of antagonistic pleiotropy to the maintenance of genetic variation for wing form and the age schedule of reproduction is discussed. INTRODUCTION 1975, 1990a). But though there is abundant evidence of a phenotypic trade-off between repro- Migrationby flight is an important aspect of the duction and wing morph, there are no studies life history of many insect species, permitting them demonstrating that these tradeoffs have a genetic to colonize highly ephemeral habitats (Southwood, basis and are thus examples of antagonistic 1962; Johnson, 1969; Harrison, 1980; Dingle, pleiotropy. 1985). However, the benefits of migration may be The purpose of the present study was to test offset in part both by the energetic cost of flight the hypothesis that in the sand cricket, Gryllus and by the cost of producing the machinery of firmus, there are negative genetic correlations flight, viz the wing muscles, wings etc (Roff, 1977, between wing morph and reproductive traits. It 1986a; Roff and Fairbairn, 1990; Inglesfield and has previously been shown that in this species the Begon, 1983; Denno et a!., 1989). The latter cost macropterous (flight capable) morph has a delayed has been demonstrated by experiments with wing age at first reproduction and a decreased total dimorphic insects, (i.e., species in which some fecundity (Roff, 1984, 1989). Furthermore, the individuals within a population or family have heritability of wing morph is large (approximately functional wings and are capable of flight, while 0.65, Roll, 1986b), and hence selection for others have reduced wings and are incapable of increased or decreased incidence of macroptery flight). Within such species there is a consistent should produce rapid changes, a prediction pattern of delayed reproduction and reduced verified by artificial selection (Roll, 1990b). The fecundity of the fully winged morph (Roff, 1986a; existence of genetic correlations between wing Denno et a!., 1989; Roll and Fairbairn, 1990). If morph and reproductive characters was investi- genetically based, these trade-offs could sig- gated by measuring correlated changes in the age nificantly influence the evolution of the incidence schedule of reproduction during the course of this of wing dimorphism within a population (Roll, selection experiment. 170 D. ROFF MATERIALS AND METHODS tion design was followed from the outset and no initial group of single pair matings constructed. Experimental protocol To estimate the age schedule of fecundities individual pairs were set up as described in Roll Detailsof the species and rearing methods are (1984). Adults were fed Purina© rabbit chow and given in Roff (1986b), and only the salient points eggs removed weekly for a period of four weeks. are presented here. The stock of G. firmus used in In pairs from the lines selected for macroptery the present study was derived from approximately both males and females were macropterous, while 40 individuals (approximate sex ratio 1: 1) from a in the pairs from the micropterous lines both adults single location in northern Florida in 1981. They were micropterous. For the control lines, the male are maintained in diapause averting conditions and female within a pairing were of the same (25-30°C, no set photoperiod but the laboratory morph, one half of the pairings comprising both lights ensure a relatively long light period), with macropterous adults and one half comprising both a breeding stock of between 100—300 individuals. micropterous adults. Approximately 20 replicates For the selection experiments individuals were were used for each combination, the exact number raised in batches of 60 individuals per disposable depending upon numbers emerging, space and mouse cage, as described in Roff (1986b). Food labour limitations. The fecundity of the microp- was provided, ad libitum and comprised Purina© terous females in the LW lines and the fecundity rabbit chow and fresh lettuce leaves. of the macropterous females in the SW lines were Previous studies were conducted at 30°C and not measured because by generation 5, when a photoperiod of 17 hrs L: 7 hrs D: under these fecundity trials were initiated, there were too few conditions the proportion of macropterous males to obtain a sufficient sample: as explained below and females is about 76 per cent and 64 per cent, this does not interfere with the statistical analysis. respectively. To reduce this percentage to around In the first selection experiment, measurements 50% in the females, in the present experiment were made at generations 5 and 15, and in the crickets were reared at 28°C, 15 hrs L; 9 hrs D. The second experiment at generations 5, 6 and 11. The first selected lines were initially constructed as total sample size was 419, comprising 216 macrop- follows: eggs were obtained from the stock culture terous and 203 micropterous females. In addition and the nymphs raised under the experimental to weekly egg counts, daily counts to day 14 were conditions. From this group 20 pairs were extracted made in generation 11 of the second selection comprising ten pairs LW x LW (macropterous x experiment. macropterous) and tenpairs SW x SW (micropterous x micropterous). This procedure was adopted to obtained an initial estimate of the Statisticalanalysis heritability of wing dimorphism (Roll, 1990b). Themeasurement of the generic correlation To establish a macropterous line (hereafter between a threshold trait and some other character referred to as the LW line) 200 adults (100 males, presents particular problems. To understand these, 100 females) from the ten LW x LW matings were first consider the usual situation of two con- mixed together, with approximately equal rep- tinuously varying characters X and Y (e.g., body resentation from each family. Similarly, a and size and fecundity) for which a phenotypic micropterous line (SW) was started by mixing relationship exists (fig. 1). Suppose X is increased together the offspring from the SW x SW crosses. or decreased by artificial selection; if the genetic A Control line (C) was formed by mixing the correlation, rg, between X and Y is 1 then the offspring from all crosses. For each of the lines, correlated trait Y will "slide" along the regression six cages containing 60 newly hatched nymphs per to the point A, but if rg equals 0 there will be no cage were established. Males and females of a change in Y with a change in X (point B in fig. 1). desired morph were selected upon eclosion into If rg lies between 0 and 1 trait Y will respond by adults until approximately 100 of each sex were moving to a point between A and B. obtained. The relatively large number of parents For a threshold trait we can distinguish only was used to prevent inbreeding depression. The discrete phenotypes, which are, in this case, the mass selection procedure outlined above was fol- macropterous (LW) and micropterous (SW) lowed on all generations subsequent to the first. morphs. However, we assume that there is an To provide a replicate, a second series of lines was underlying variable that shows continuous vari- set up some months after the first, the protocol ation, genotypes exceeding the threshold value followed being identical except that a mass selec- producing one phenotype and those below the ANTAGONISTIC PLEIOTROPY AND WING DIMORPHISM 171 of SW. The two scenarios can be distinguished by measuring Y at several generations during selec- tion. Note that even if the value of Y for the LW and SW morphs stay the same under selection, the mean value of Y in the selected population will change by virtue of the changing frequencies of macropterous and micropterous individuals. Y The basic statistical model used in the present analysis was, Y= a+bX1+cX2+dX3 + interaction terms + error where X1 is "wing morph" (LW or SW), X2 is "treatment" (selected or control), X3 is "trial", Selected Base population Selected and a, b, c, d are constants. Effects due to "trial" may reflect systematic differences resulting from x selection or "error" differences due to "random" Figure1 Hypothetical phenotypic relationship between two variation between trials; for this reason I used a continuously varying characters X and Y (e.g.,bodysize categorical variable for "trial". fecundity). If the genetic correlation, r,equals1 selection on X will produce a correlated response to Y to A. If Tg= 0 there will be no response (B), otherwise Y will move to a point between A and B. RESULTS Theresponse to selection was very rapid, and by threshold producing the alternate (Falconer, 1981).
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