Genetic analysis of the wing-form determination of the small brown planthopper,Laodelphax striatellus (Hemiptera: Delphacidae)

Summary The mechanism of the genetic control of wing form determination of the small brown planthopperLaodelphax striatellus was analyzed by selection and crossing experiments. The selection experiment for brachypterous and macropterous forms for 13 generations showed that the wing form is under some genetic control. The crossing experiment showed that the wing form polymorphism was not a simple Mendelian character but was under polygenic control. The data from the crossing experiment were applied to the liability method for the quantitative genetics by Falconer (1981). The analysis suggested that wing form determination was a threshold character under polygenic control, and the estimated minimum number of loci concerned was 2.07. The realized heritability estimated from the data of selection experiment was 0.36 for macroptery. On the other hand, the heritability in broad sense estimated on the basis of the variance of F₁ and F₂ in crossing experiment was 0.81.     

Factors and consequences of a non-functional alary polymorphism inPyrrhocoris apterus (Heteroptera: Pyrrhocoridae)

Functional alary polymorphisms have been studied rather extensively in several insect species. This review article deals with factors controlling wing polymorphism in a flightless species,Pyrrhocoris apterus (L.), and discusses its adaptive significance and mechanisms for their persistence under natural conditions. The macropterous morph is determined by a recessive allele whose penetrance depends on photoperiod and temperature. Natural populations of this species contain a small fraction of flightless macropters. The disadvantages of being a macropter (increase of development time, decrease of fecundity) are minimal, while the benefit may consist in the tendency to prereproductive arrest of ovarian development in teneral females. It prevents establishing a second generation which would mostly die during the next winter. The mechanism of alary morph regulation may be an ancestral trait linkingP. apterus with other polymorphic Heteroptera, while its decreased penetrance may be a derivative character. Variation in fitness due to alary morphs is small compared to the one associated with differences in body size. The latter is environmentally determined, and not linked to the genetic basis of wing polymorphism. In the “mosaic” of phenotypes of various size the significance of the genetic macroptery may be close to neutral.     

Why is there so much genetic variation for wing dimorphism?

Wing dimorphism appears in general to be determined either by a single locus, 2 allele system in which brachyptery is dominant, or by the additive action of numerous loci. In the latter case studies indicate that the heritability is typically quite large. It is generally postulated that wing dimorphism is under strong selection: why then is genetic variation not eroded? In this paper I consider three possible explanations. First, genetic variation may not be exposed to selection because environmental heterogeneity effectively makes heritability zero. Because wing dimorphisms are known to evolve it seems unlikely that this is the primary factor. Second, directional selection on a threshold trait may push the population almost to monomorphism but erodes genetic variance at a very slow rate. This mechanism cannot preserve variation but makes it possible for other factors to more easily maintain variability. Finally, I demonstrate that in a heterogeneous environment spatio-temporal variation in fitness will itself maintain a genetic polymorphism for wing dimorphism. This paper is dedicated to Professor Sinzo Masaki, whose work on the evolution of wing dimorphism and diapause has been so important to our understanding of the evolution of dimorphic variation.     

Genetic correlation of wing polymorphism between females and males in the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae)

Summary Genetic correlations of the wing form and the relative wing length between females and males were estimated in the oriental chinch bug,Cavelerius saccharivorus, by calculating the correlation between the sexes of the proportion macropterous or the mean relative wing length in full-sib families obtained from different wing forms of parents emerged in a high density population. There was a significantly positive genetic correlation between the sexes in both the proportion macropterous and the mean relative wing length. However, the appearance rate of macropters tended to be much lower in males than in females under the rearing conditions which promote the appearance of macropters. This was evident especially in the offspring of brachypterous parents. These indicated that inC. saccharivorus the wing polymorphism of males is not a simple result of the genetic correlation of wing morphology between the two sexes. It was considered that both of the female and male fitness advantages to wing reduction, as well as the genetic correlation between the sexes, would influence the evolution of wing polymorphism in this species.     

Genetic changes of life history and behavioral traits during mass-rearing in the melon Fly,Bactrocera cucurbitae (Diptera: Tephritidae)

Quantitative genetic studies for life history and behavioral traits are important in quality control for insect mass-rearing programs. Firstly, a brief history of quality control in mass-reared insects is described. Next, the differentiation of many traits of wild and mass-reared melon flies,Bactrocera cucurbitae, in Okinawa is reviewed, and the factors which have caused variation in these traits are considered. As artificial selection pressures are thought to be more important than inbreeding depression and genetic drift in the mass-reared strain of the Okinawan melon fly, two artificial selection experiments were conducted to evaluate genetic variations and genetic correlations among life history and behavioral traits. These are divergent selections for age at reproduction and for developmental period. The genetic relationship among 5 traits, i.e. longevity, age at reproduction, developmental period, circadian period, and time of mating was clarified and discussed in relation to genetic changes of traits during the mass-rearing. The results suggest that the genetic trade-off relationships between traits should be taken into account in mass-rearing programs.     

Genetic control and evolution of acoustic signals in planthoppers (Homoptera: Delphacidae)

Acoustic signals are part of the specific mate recognition system of planthoppers. The genetic control of acoustic signal characters was studied in the planthopperRibautodelphax imitans. Artificial selection for interpulse interval in the female call revealed a large additive genetic component for this polygenic character. Other female call characters showed a correlated response. Some male call characters also appeared to be genetically correlated with the female character selected for, despite the rather different structure of male and female calls. Parent-offspring regression provided significant heritability estimates for those male call characters that also responded to artificial selection in the female call, one of which appeared to be influenced by sex-linked genes. It is argued that the differentiation of this mate recognition system in planthopper populations and species could be the result of founder effects, enabled by the genetic plasticity of the call characters and the existence of a wing length dimorphism in these animals.     

Genetic variation of density responses in relation to wing polymorphism in the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae)

Summary Responses to nymphal density in the determination of wing form were compared between the offspring from brachypter x brachypter crosses and those from macropter x macropter crosses of the oriental chinch bug,Cavelerius saccharivorus. In the offspring from crosses between macropters, there was a strong tendency for macropters to increase to a rather high level with increasing nymphal density in both sexes. In contrast to this, in the offspring from crosses between brachypters, the appearance rate of macropters attained a maximum value in moderately crowded conditions and conversely decreased to a lower level in more crowded conditions in both sexes. Thus density response patterns concerning the determination of wing form were quite different between the offspring from different crosses in the wing form, indicating that there is a genetic basis underlying wing polymorphism in this species. As for the body size of emerged adults, macropters tended to be larger than brachypters in the same crowded condition. Moreover, the rate of decrease of body size with nymphal density was lower in the offspring from crosses between macropters than in the offspring from crosses between brachypters. This indicated that the former offspring are more tolerant of nymphal crowding than the latter. The difference in such a tolerance against nymphal crowding between the offspring from different crosses was considered to be related to the difference in the appearance of macropters in the crowded conditions between them.     

Ecological significance of the wing polymorphism of the oriental chinch bug,Cavelerius saccharivorusOkajima (Heteroptera: Lygaeidae)

Summary The wing polymorphism of the oriental chinch bug,Cavelerius saccharivorus, was studied in relation to its reproductive strategy. The frequency distribution of wing form in terms of the relative wing length (R.W.L.) changed with increase in the population density from a single modality biased toward the brachypterous form to bimodality with both macropterous and more extreme brachypterous forms. Since some evidences in the field showed that fliers are limited to macropters, such a bimodality of wing form indicated thatC. saccharivorus employs a mixed strategy with some portion of adults staying on in the same habitat, whereas the rest disperse. Moreover, macropters were larger in body size than brachypters emerging in high density populations, although smaller than brachypters emerging in low density populations. Larger body size as well as the delay of ovarian maturation in macropters was considered to be a beneficial trait for migratory behaviour.     

Phenological studies on the wing dimorphism of a semi-aquatic bug,Microvelia douglasi (Heteroptera: Veliidae)

Summary Seasonal changes in the percentage of macropterous individuals in natural population ofMicrovelia douglasi in permanent habitats were investigated in Shimane, western part of Japan. Macropterous forms of 5th instar nymphs occurred most frequently from late June to early July. However, the seasonal appearance of macropterous adults differed between habitats and years. There were significant effects of density, temperature, photoperiod and food availability on the percentage of macropters under laboratory conditions. The percentage of macropterous forms generally increased with density, but the mode of the response was modified by temperature, photoperiod and food availability. When insects were reared under long day photperiod (16L8D) with abundant food (0.5–1.0 g/container/day), the percentage of macroptery varied most among the different density treatments at the intermediate temperature (24°C) and the difference was reduced at higher temperature (28° and 32°C). Effects of temperature were small under the short day photoperiod (12L12D). Under 12L12D photoperiod sensitivity of density responses were higher at 32°C and lower at 24°C than those under 15L8D photoperiod. When the insects were supplied with food every other day, the density response became obscure. Ecological significance of environmental responses in the wing morph determination was discussed in relation to the heterogeneous environment of water surface.     

Wing dimorphism and the migratory syndrome: Correlated traits for migratory tendency in wing dimorphic insects

The hypothesis that the morphological, physiological, and behavioral traits comprising the migratory syndrome in insects are genetically correlated through pleiotropic effects of genes controlling the titre of a common hormonal determinant is explored. Evidence that juvenile hormone (JH) influences the component traits of the migratory syndrome is presented, and thus JH is assumed to be the underlying, common determinant. However, readers are cautioned that this does not imply that JH is solely responsible for these traits, nor is this necessary for the arguments presented. For wing dimorphic taxa, the “correlated traits hypothesis” predicts covariance within wing morphs between JH titre and the proportion winged. Four simple genetic models for wing-morph determination are considered: single-locus with short-winged (SW) dominant; single-locus with long-winged (LW) dominant; polygenic, fixed threshold, shifting distribution; and polygenic, shifting threshold, fixed distribution. In each case, wing morphology is assumed to be a threshold trait with the liability being JH titre at some critical stage of development. All models predict covariation between %LW and the mean JH titre of at least one of the wing morphs, but the form and direction of the relationship depends critically on the genetic model used. The results suggest that we should expect the traits associated with the migratory syndrome, and hence the trade-offs associated with the evolution of wing dimorphism, to be correlated with proportion winged and, in this sense, to be frequency-dependent.     

The spatial distribution pattern of the brown planthopperNilaparvata lugens St?l (Homoptera: Delphacidae) in west Java, Indonesia

Summary Spatial distribution pattern of the brown planthopper (BPH) was analyzed at 9 experimental fields in the northern part of West Java during two consecutive rice cropping seasons, i.e., wet and dry seasons. The population of each developmental stage and wing form of BPH at each location showed consistent departure from the random (Poisson) distribution, the variances of the densities in most cases exceeding their means. Namely, the distribution pattern of BPH per hill of rice plant was found to have a general tendency to be aggregated or contagious and to fit fairly well to the negative binomial model. The tendency for aggregation was further confirmed by both the β-values of -m regression being larger than unity and theC _A -values being larger than zero for each developmental stage. Although significant variations in the distribution pattern as measured by β- orC _A -value were observed between different developmental stages, between wing forms and among locations, the degree of aggregation for a given developmental stage at each experimental field remained fairly stable throughout the crop period, despite wide temporal changes in population density. Possible factors to explain these characteristics of the spatial distribution pattern of the BPH in West Java were discussed with reference to the process generating it.     

Rapid evolution of larval development time during mass-rearing in the melon fly, Bactrocera cucurbitae

A quantitative genetic analysis of rapid evolution of a life history trait has been conducted on the first 24 generations of mass-rearing in the melon fly Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). The phenotypic variance of larval development time in each generation was divided into genetic and residual components. Mean and phenotypic coefficients of variation of larval development time decreased gradually as generations proceeded as a result of artificial selection for shorter larval period in the mass-rearing procedure. There was a trend that additive genetic coefficients of variation in larval development time decreased with generations. These changes are entirely attributed to genetic responses to laboratory selection under the mass-rearing environment because the population was maintained at a very large size so as to exclude random genetic drift and inbreeding depression, which would be other factors responsible for the observed genetic changes. The residual coefficients of variation in larval development time did not change with generations. Realized heritability of larval development time was low. The heritabilities for larval development time estimated from parent–offspring regression at generations 60 and 70, when the evolutionary plateau was asymptotically reached, were not significantly larger than 0. Received: April 22, 1999 / Accepted: September 20, 1999     

Reproduction and egg diapause of the oriental chinch bug,Cavelerius saccharivorus okajima (Heteroptera: Lygaeidae), in the subtropical winter season in relation to its wing polymorphism

Summary Reproduction and egg diapause of the oriental chinch bug,Cavelerius saccharivorus, in the subtropical winter season were investigated in relation to its wing polymorphism. Macropterous females collected from the southern part of Okinawa Is. in the autumn season delayed their oviposition and were less fecund early in the adult life period, but survived much longer than brachypterous females collected from the same locality. The total fecundity was not significantly different between wing morphs. The diapause of eggs laid by brachypters tended to be terminated more easily at a high temperature than that of eggs laid by macropters. This indicated that the eggs laid by macropters were more intense in their diapause than those laid by brachypters. However, irrespective of the parental wing form, diapause showed considerable variation in its intensity within and between clutches.     

A male fitness advantage to wing reduction in the oriental chinch bug,Cavelerius saccharivorus okajima (heteroptera: Lygaeidae)

Summary A male fitness advantage to wing reduction was investigated for the oriental chinch bug,Cavelerius saccharivorus, which is wing dimorphic. Field surveys for the frequency of matings between wing morphs showed that the mating probability was much higher in brachypterous males than in macropterous males in the early breeding season. Brachypterous males copulated with not only brachypterous females but also macropterous females in this season. This led to a considerable rate (30%) of insemination of macropterous females just before emigration even in the early breeding season. A rearing experiment for the pre-reproductive period of females revealed that females mated with brachypterous males copulated and oviposited earlier than females mated with macropterous males. These results indicate that there is a fitness advantage to wing reduction in the males ofC. saccharivorus in terms of the earlier sexual maturation.     

Association of late childbearing with healthy longevity among the oldest-old in China

Statistical analysis of a large and unique longitudinal data-set demonstrates that childbearing after age 35 or 40 is associated with survival and healthy survival among very old Chinese women and men. The association is stronger for women than for men. The estimates are adjusted for a variety of confounding factors: demographic characteristics, family support, social connections, health practices, and health conditions. Further analysis based on an extension of the Fixed-Attributes Dynamics method shows that late childbearing is positively associated with long-term survival and healthy survival from ages 80-85 to 90-95 and 100-105. This association exists among oldest-old women and men, but, again, the effects are substantially stronger for women than for men. We discuss four possible factors that may explain why late childbearing affects healthy longevity at advanced ages: (1) social factors; (2) biological changes caused by late pregnancy and delivery; (3) genetic and other biological characteristics; and (4) selection.     

Habitat fragmentation promotes fluctuating asymmetry but not morphological divergence in two geckos

The process of species formation is often ignored in discussions on the conservation of biodiversity. Yet the clearance of vegetation may promote divergence among populations of a species through isolation, providing conditions for rapid genetic drift and novel selection pressures. Here, stepwise discriminant function analysis and fluctuating asymmetry are used to examine variation in morphology within and among non fragmented and recently fragmented populations of two species of gecko,Oedura reticulata andGehyra variegata. High reclassification error rates using discriminant function analysis, indicate that fragmentation has had no detectable effect on morphological differentiation among populations of either species. In contrast, remnant populations of both species exhibit higher mean levels of fluctuating asymmetry than do populations in undisturbed habitat. ForOedura reticulata, levels of fluctuating asymmetry are negatively correlated with the log of adult population size. These results suggest that the changes following habitat clearance have been severe enough to cause increased developmental instability in populations of both species but not detectable morphological divergence. Given the high rate of extirpation of gecko populations in the study region and the extreme vulnerability of the remaining populations, it is unlikely that species formation will be significant in maintaining reptile diversity in that region.     

The President's Council on Bioethics: Choosing Sex of Children

The ideology of eugenics as it arose in the late nineteenth century was concerned with the perceived negative effects of differential fertility on the “quality” and composition—variously defined—of future generations. Quality was to be raised by preventing or discouraging the “less fit” from propagating themselves and by encouraging childbearing among couples seen as carriers of desirable genetic endowments. Thus, this old‐fashioned eugenics was directed, at least by intent and sometimes in practice, to select among parents and influence their procreative behavior. The rules for such selection were typically decided, democratically or otherwise, on the advice of anointed “experts.” By the mid‐twentieth century these programs had come to be thoroughly discredited, both because they were seen to lack scientific validity and, perhaps especially, as a result of Nazi racial policies. Modern technological developments have given rise to, or created the realistic prospect of, a different, “grassroots” eugenics: parents voluntarily choosing qualities they would prefer their offspring to possess. Their right to do so would seem to be a straightforward extension of the principle, endorsed by numerous international declarations, that “all couples and individuals have the basic right to decide freely and responsibly the number and spacing of their children.” Parents might exercise that right by seeking to improve their children's chances for a healthy and successful life or merely by following their own prejudices. But the application of this principle tacitly assumes that the aggregate outcome of individual reproductive decisions is in reasonable harmony with the collective interest. The technological means now available for parents to select preferred physical and intellectual characteristics of their children—to improve on the outcome of the natural genetic lottery—are rudimentary. Many of them are still in the domain of science fiction. But in one major instance the technology already exists: selecting the sex of children, especially through early determination of the sex of the fetus, which then may be followed by sex‐selective induced abortion. This option has become widely available in recent years. In societies in which there is strong preference for offspring of a particular sex—usually for boys— the result has been a marked deviation from the normal sex ratio at birth. The social implications of such uncoordinated individual choices are perceived as clearly deleterious, hence the practice of sex‐selective abortion is commonly outlawed even when abortion is otherwise permitted. Invoking a social interest that overrides the right of voluntary parental choice of course raises the question whether other untoward effects of socially unregulated fertility might also be grounds for circumscribing parental freedom of choice in childbearing. Socially undesirable patterns of childbearing could be defined as “non‐responsible.” In the United States, the President's Council on Bioethics, appointed in November 2001, is one of the many committees advising the US President on important social issues. The Council consists of 17 members, including noted scientists, physicians, ethicists, social scientists, legal experts, and theologians. It had the mandate, inter alia, to inquire into the human and moral significance of developments in biomedical and behavioral science and technology and to explore specific ethical and policy questions related to these developments. The Council was chaired by Professor Leon R. Kass, a prominent bioethicist from the University of Chicago. Its report, under the title, Beyond Therapy: Biotechnology and the Pursuit of Happiness, was submitted to the President in October 2003. This massive document, which will also be published in book form by a commercial publishing house, is available online at: http://bioethicsprint.bioethics.gov/reports/beyondtherapy . Reproduced below is a section titled Choosing sex of children from Chapter 2 of the report (titled Better Children,). It offers a penetrating analysis and commentary on issues raised by parental selection of children's sex.     

Socio-economic correlates of fecundability in a sample of taiwanese women

Estimates of fecundability (monthly probability of conception) in the absence of contraception are derived from the frequency distribution of conceptive delays immediately following marriage, reported by 2,443 married women aged 20 to 39 included in the Taichung (Taiwan) Intensive Fertility Survey of 1962. Average fecundability of women is positively associated with their socio-economic status. These differentials are not accounted for by differences among socio-economic groups with respect to memory and truncation biases (associated with the marriage duration), wife’s age at marriage, or unreported premarital conceptions. A Multiple Classification Analysis suggests that among the socio-economic characteristics, husband’s education, rural background, and modern family type are the more important predictors of fecundability. The importance of genetic factors as opposed to cultural factors in producing these socio-economic differences in fecundability can not be evaluated systematically. Moreover, the relation of a couple’s privacy, their attitude toward family building, and patterns of mate selection to their socioeconomic status would have to be taken into account before the differences in fecundability could be attributed to factors such as nutrition, health, or infections which might directly influence their physiological ability to conceive.     

Population Composition,Public Policy,and the Genetics of Smoking

In this article, we explore the effect of public policy on the extent to which genes influence smoking desistance. Using a sample of adult twins (n _mz = 363, n _dz = 233) from a large population registry, we estimate Cox proportional hazards models that describe similarity in the timing of smoking desistance among adult twin pairs. We show that identical twin pairs are significantly more likely to quit smoking within a similar time frame compared with fraternal twin pairs. Importantly, we then show that genetic factors for smoking desistance increase in importance following restrictive legislation on smoking behaviors that occurred in the early and mid-1970s. These findings support the social push perspective and make important contributions to the social demography and genetic epidemiology of smoking as well as to the gene-environment interaction literatures.     

The comparison of the interpersonal action component of woman-centred care reported by healthy pregnant women in different sized practices in the Netherlands: A cross-sectional study

BackgroundThe number of interventions is lower, and the level of satisfaction is higher among women who receive midwife-led primary care from one or two midwives, compared to more midwives. This suggests that midwives in small-sized practices practice more women-centred. This has yet to be explored.ObjectiveTo examine pregnant women’s perceptions, of the interpersonal action component of woman-centred care by primary care midwives, working in different sized practices.MethodsA cross-sectional study using the Client Centred Care Questionnaire (CCCQ), administered during the third trimester of pregnancy among Dutch women receiving midwife-led primary care from midwives organised in small-sized practices (1−2 midwives), medium-sized (3−4 midwives) and large-sized practices (≥5 midwives). A Welch ANOVA with post hoc Bonferroni correction was performed to examine the differences.Results553 completed questionnaires were received from 91 small-sized practices/104 women, 98 medium-sized practices/258 women and 65 large-sized practices/191 women. The overall sum scores varied between 57–72 on a minimum/maximum scoring range of 15–75. Women reported significantly higher woman-centred care scores of midwives in small-sized practices (score 70.7) compared with midwives in medium-sized practices (score 63.6) (p < .001) and large-sized practices (score 57.9) (p < .001), showing a large effect (d .88; d 1.56). Women reported statistically significant higher woman-centred care scores of midwives in medium-sized practices compared with large-sized practices (p < .001), showing a medium effect (d .69).ConclusionThere is a significant variance in woman-centred care based on women’s perceptions of woman-midwife interactions in primary care midwifery, with highest scores reported by women receiving care from a maximum of two midwives. Although the CCCQ scores of all practices are relatively high, the significant differences in favour of small-sized practices may contribute to moving woman-centred care practice from ‘good’ to ‘excellent’ practice.