Patricia Gibert

CHILL‐COMA TOLERANCE, A MAJOR CLIMATIC ADAPTATION AMONG DROSOPHILA SPECIES

Abstract.— Most drosophilid species can be classified either as temperate or tropical. Adults of species were submitted to a cold treatment (0°C) and then brought back to ambient temperature. They generally exhibited a chill coma and the time needed to recover was measured. We found in a set of 26 temperate species that recovery was rapid (average 1.8 min, range 0.15–4.9). In contrast, a long recovery time (average 56 min, range 24–120) was observed for 48 tropical species. A few species, like Drosophila melanogaster, are cosmopolitan and can proliferate under temperate and tropical climates. In 9 of 10 such species, slight genetic differences were found: a shorter recovery in temperate than in tropical populations. Comparing physiological data to phylogeny suggests that chill‐coma tolerance has been a recurrent adaptation that is selected for in cold climates but tends to disappear under a permanently warm environment. This major climatic adaptation, evidenced in drosophilids, seems to occur in other insect groups also.

Proximate Causes of Rensch’s Rule: Does Sexual Size Dimorphism in Arthropods Result from Sex Differences in Development Time?

A prominent interspecific pattern of sexual size dimorphism (SSD) is Rensch’s rule, according to which male body size is more variable or evolutionarily divergent than female body size. Assuming equal growth rates of males and females, SSD would be entirely mediated, and Rensch’s rule proximately caused, by sexual differences in development times, or sexual bimaturism (SBM), with the larger sex developing for a proportionately longer time. Only a subset of the seven arthropod groups investigated in this study exhibits Rensch’s rule. Furthermore, we found only a weak positive relationship between SSD and SBM overall, suggesting that growth rate differences between the sexes are more important than development time differences in proximately mediating SSD in a wide but by no means comprehensive range of arthropod taxa. Except when protandry is of selective advantage (as in many butterflies, Hymenoptera, and spiders), male development time was equal to (in water striders and beetles) or even longer than (in drosophilid and sepsid flies) that of females. Because all taxa show female‐biased SSD, this implies faster growth of females in general, a pattern markedly different from that of primates and birds (analyzed here for comparison). We discuss three potential explanations for this pattern based on life‐history trade‐offs and sexual selection.

Cold stress tolerance in Drosophila: analysis of chill coma recovery in D. Melanogaster

Abstract Drosophila melanogaster adults, grown at 21°C, were distributed in groups of 50 after a light anaesthesia. Culture vials with flies were later submitted to a cold treatment at 0°C. All adults entered a chill coma; the recovery time was measured at ambient temperature. 2. Recovery time was strongly influenced by recovery temperature, with shorter values between 20-25°C. 3. Recovery time increased almost linearly with duration of cold treatment. 4. Recovery time was consistently larger for males than for females. It was highly variable among groups and increased with flies’ age. 5. Variability among flies of the same group was always very high, with CVs often over 25%. 6. Chill coma and its recovery seem to imply a modification of the nervous system, analogous in several aspects to what is observed with usual anaesthetics such as CO 2 .

Isofemale lines in Drosophila: an empirical approach to quantitative trait analysis in natural populations

Founding isofemale lines from wild collected females is a basic tool for investigating the genetic architecture of Drosophila natural populations. The method permits the analysis of quantitative traits under laboratory conditions, with a much broader scope than the mere evidence of a significant genetic heterogeneity among lines. Genetic variability is generally demonstrated by a significant coefficient of intraclass correlation, but several experimental precautions are needed and explained here. The relationship between classical (additive) heritability and intraclass correlation is not straightforward, presumably because the genetic bottlenecks due to the initiation of the lines unravel a significant, nonadditive genetic variance due to dominance and epistatic effects. It is thus suggested to consider intraclass correlation as a specific genetic parameter that enables comparisons between different traits, different populations or different environments. The use of isofemale lines is, however, not restricted to the calculation of an intraclass correlation. It can be used to estimate genetic correlations among traits or environments. The method is also convenient for the analysis of phenotypic plasticity in relation to an environmental gradient. A precise description of the response curves (the reaction norms) is possible, distinguishing trait parameters and plasticity parameters. A fairly general conclusion is that, for a given trait, mean value and plasticity are genetically independent. It is also possible to analyze traits, which, like sexual dimorphism, must be measured on different individuals, and even to demonstrate their genetic variability. In many cases, further empirical and theoretical analyses are possible and needed. It is argued that, in the future, isofemale lines will have an increasing significance among the various techniques appropriate to the analysis of quantitative evolutionary genetics in a diversity of species.

Chill-Coma Temperature in Drosophila: Effects of Developmental Temperature, Latitude, and Phylogeny

We modify and apply a nonlethal technique for rapidly quantifying the cold tolerance of large numbers of Drosophila and other small insects. Flies are transferred to individual vials, cooled in groups in progressive 0.5°C steps, and checked for loss of righting response (chill‐coma temperature [Tcc]). Flies recover quickly when transferred to ambient temperature, and thus this technique potentially can be used in selection experiments. We applied this technique in several experiments. First, we examined the sensitivity of Tcc to developmental temperature. Drosophila melanogaster (Congo, France), Drosophila subobscura (Spain, Denmark), and Drosophila ananassae (India) were reared from egg to adult at 15°, 18°, 25°, or 29°C, transferred to 15°C for several days, and then progressively chilled: Tcc was positively related to developmental temperature, inversely related to latitude of the population, but independent of sex. The sensitivity of Tcc to developmental temperature (acclimation flexibility) was marked: Tcc shifted on average 1° for each 4°C shift in developmental temperature. Among 15 species of the obscura group of Drosophila, Tcc varied from −0.1° to 4.5°C; Tcc was inversely related to latitude in both nonphylogenetic and phylogenetically based ANCOVA (standardized independent contrasts) and was unrelated to body size.

DESICCATION AND STARVATION TOLERANCE OF ADULT DROSOPHILA: OPPOSITE LATITUDINAL CLINES IN NATURAL POPULATIONS OF THREE DIFFERENT SPECIES

Desiccation and starvation tolerance were measured along latitudinal transects in three Drosophilid species (Drosophila ananassae, D. melanogaster, and Zaprionus indianus) of the Indian subcontinent. In each case, significant latitudinal clines were observed; desiccation tolerance increased with latitude while starvation tolerance decreased. Such field observations suggest that desiccation and starvation tolerance are fitness related traits that are independently selected in nature and genetically independent. It was, however, difficult to relate these genetic changes with precise climatic variables, except winter temperature. The overall negative correlation between the two traits, which was evidenced in natural populations, contrasts with a positive correlation generally observed in various laboratory selection experiments and that also seems to exist between different species. These observations point to the difficulty of interpreting correlations among fitness‐related traits when different evolutionary levels are compared, and also different sets of data, that is, field versus laboratory studies.

Phenotypic plasticity and developmental temperature in Drosophila: Analysis and significance of reaction norms of morphometrical traits

Abstract In ectotherm species like Drosophila, morphometrical quantitative traits exhibit a large amount of phenotypic plasticity as a consequence of developmental temperature. Generally the response curves called reaction norms, are non-linear and exhibit either single maxima or minima, or a sigmoid shape. Up to now, such complex shapes were mainly considered as non-adaptive, reflecting interactions between internal developmental constraints and the environment. We show that the general, all purpose method of polynomial adjustment provides a convenient way for the description of the norms. From the polynomial parameters, characteristic values with an obvious biological significance (e.g. the temperature of a maximum size), can be calculated and used for comparing populations or species. Experimental data on body pigmentation and body size show that the shapes of the reaction norms are genetically variable and suggest adaptive responses to the environment. The possible occurrence of specific regulatory genes acting on norm shape will be a matter for further investigations.

Comparative analysis of morphological traits among Drosophila melanogaster and D. simulans: genetic variability, clines and phenotypic plasticity

The two sibling cosmopolitan species, Drosophila melanogaster and D. simulans, are able to proliferate under very different climatic conditions. This has resulted in local adaptations, which are often arranged in latitudinal clines. Such clines are documented for body weight, wing and thorax length, sternopleural and abdominal bristle number, ovariole number and thoracic pigmentation. The overall magnitude of geographical differentiation is, however, much less in D. simulans than in D. melanogaster, and latitudinal clines are less pronounced.The fact that natural populations live under different climates raises the problem of interaction between temperature and phenotype. The reaction norms of morphometrical traits have been investigated as a function of growth temperature. The shapes of the response curves vary according to the investigated trait. They are generally curvilinear and can be described by calculating characteristic values after polynomial adjustments. For a given trait, the reaction norms of the two species are similar in their shape, although some significant differences may be observed.Within each species, significant differences are also observed between geographic populations: reaction norms are not parallel and the divergence is better marked when more distant populations (e.g., temperate and tropical) are compared. It thus appears that besides mean trait value, phenotypic plasticity is also a target of natural selection.A specific analysis of wing shape variation according to growth temperature was also undertaken. Reaction norms with different shapes may be observed in various parts of the wing: the major effect is found between the basis and the tip of the wing, but in a similar way in the two species. By contrast, some ratios, called wing indices by taxonomists, may exhibit completely different reaction norms in the two species.For a single developmental temperature (25°C) the phenotypic variability of morphometrical traits is generally similar in the two species, and also the genetic variability, estimated by the intraclass correlation. A difference exists, however, for the ovariole number which is less variable in D. simulans. Variance parameters may vary according to growth temperature, and a detailed analysis was made on wing dimensions. An increase of environmental variability at extreme, heat or cold temperatures, has been found in both species. Opposite trends were, however, observed for the genetic variability: a maximum heritability in D. simulans at middle temperatures, corresponding to a minimum heritability in D. melanogaster. Whether such a difference exists for other traits and in other populations deserves further investigations.In conclusion, morphometrical analyses reveal a large amount of significant differences which may be related to speciation and to the divergence of ecological niches. Within each species, numerous geographic variations are also observed which, in most cases, reflect some kinds of climatic adaptation.

LOCOMOTOR PERFORMANCE OF DROSOPHILA MELANOGASTER: INTERACTIONS AMONG DEVELOPMENTAL AND ADULT TEMPERATURES, AGE, AND GEOGRAPHY

We explored the extent to which a phenotypic trait (walking speed) of Drosophila melanogaster is influenced by population, developmental temperature, adult temperature, and age. Our goals were to estimate the importance of these factors and to test the beneficial acclimation hypothesis. We measured speed of flies from two populations (the Congo and France) that developed at different temperatures (18, 25, and 29 C) and were tested at different temperatures (18, 25, and 29°C) and ages (2, 7, 13 days). Not surprisingly, speed increased strongly with test temperature. Speed was generally greatest for flies reared at an intermediate developmental temperature, contrary to the beneficial acclimation hypothesis, which predicts that speed would be greatest when influenced by interactions involving population. For example, speed was greatest for flies from France that developed at a low temperature, but for flies from the Congo that developed at a high temperature. The impact of developmental temperature declined with age. Surprisingly, speed actually increased with age for flies raised and maintained at a low temperature, but decreased with age for flies raised and maintained at an intermediate or at a high temperature. Thus, walking performance is highly dynamic phenotypically, complicating potential attempts to predict responses to selection on performance.

Ecology and life history evolution of frugivorous Drosophila parasitoids.

Parasitoids and their hosts are linked by intimate and harmful interactions that make them well suited to analyze fundamental ecological and evolutionary processes with regard to life histories evolution of parasitic association. Drosophila aspects of what parasitoid Hymenoptera have become model organisms to study aspects that cannot be investigated with other associations. These include the genetic bases of fitness traits variations, physiology and genetics of resistance/virulence, and coevolutionary dynamics leading to local adaptation. Recent research on evolutionary ecology of Drosophila parasitoids were performed mainly on species that thrive in fermenting fruits (genera Leptopilina and Asobara). Here, we review information and add original data regarding community ecology of these parasitoids, including species distribution, pattern of abundance and diversity, host range and the nature and intensity of species interactions. Biology and the evolution of life histories in response to habitat heterogeneity and possible local adaptations leading to specialization of these wasps are reported with special emphasis on species living in southern Europe. We expose the diversity and intensity of selective constraints acting on parasitoid life history traits, which vary geographically and highlight the importance of considering both biotic and abiotic factors with their interactions to understand ecological and evolutionary dynamics of host-parasitoid associations.