Phenotypic Plasticity Of Body Size Research Articles

Egg size-dependent embryonic development in the desert locust, Schistocerca gregaria

Phenotypic plasticity in body size is a product of modification of the developmental pathway. Although hatchlings of the desert locust, Schistocerca gregaria, show egg size-dependent plasticity in body size, it remains unclear how embryogenesis during egg development regulates final embryonic body size. To determine the developmental pathway causing body size variation at hatching, we examined egg and embryonic development at the early, middle, and late egg developmental stages in S. gregaria by comparing small and large eggs. Crowd-reared females produced larger eggs than isolated-reared females. The daily egg developmental rate was similar between small and large eggs: eggs dramatically absorbed external water after days 3 to 7 and nearly doubled the initial egg weight at the late stage of day 12. Morphological measurements of eggs and embryos at different days after oviposition revealed that large eggs were longer than small eggs throughout developmental stages. However, embryo length was similar between small and large eggs at the early stage (anatrepsis). Embryos begin to absorb yolk into their bodies after blastokinesis. The size of large-egg embryos increased significantly from the middle stage (katatrepsis) due to absorption of more yolk than small eggs. Egg length and embryo length were conspicuously larger in large eggs than in small eggs on day 12 of late katatrepsis. These results suggest that egg size did not influence the egg developmental rate and initial embryo size. Large eggs had more yolk and space, resulting in larger final embryos than small eggs. The amount of yolk and size of eggshells during katatrepsis could play a key role in determining hatchling body size in S. gregaria.

Remarkable insensitivity of acorn ant morphology to temperature decouples the evolution of physiological tolerance from body size under urban heat islands.

Environmental temperature can alter body size and thermal tolerance, yet the effects of temperature rise on the size-tolerance relationship remain unclear. Terrestrial ectotherms with larger body sizes typically exhibit greater tolerance of high (and low) temperatures. However, while warming tends to increase tolerance of high temperatures through phenotypic plasticity and evolutionary change, warming tends to decrease body size through these mechanisms and thus might indirectly contribute to worse tolerance of high temperatures. These contrasting effects of warming on body size, thermal tolerance, and their relationship are increasingly important in light of global climate change. Here, we used replicated urban heat islands to explore the size-tolerance relationship in response to warming. We performed a common garden experiment with a small acorn-dwelling ant species collected from urban and rural populations across three different cities and reared under five laboratory rearing temperatures from 21 to 29 °C. We found that acorn ant body size was remarkably insensitive to laboratory rearing temperature (ant workers exhibited no phenotypic plasticity in body size across rearing temperature) and among populations experiencing cooler rural versus warmer urban environmental temperatures (no evolved differences in body size between urban and rural populations). Further, this insensitivity of body size to temperature was highly consistent across each of the three cities we examined. Because body size was robust to temperature variation, previously described plastic and evolved shifts in heat (and cold) tolerance in acorn ant responses to urbanization were shown to be independent of shifts in body size. Indeed, genetic (colony-level) correlations between heat and cold tolerance traits and body size revealed no significant association between size and tolerance. Our results show how typical trait correlations, such as between size and thermal tolerance, might be decoupled as populations respond to contemporary environmental change.

Variability in vertebral numbers does not contribute to sexual size dimorphism, interspecific variability, or phenotypic plasticity in body size in geckos (Squamata: Gekkota: Paroedura).

Body size is a fundamental trait correlated with nearly every aspect of animal life. It is influenced by numerous genetic and non-genetic factors. Despite its central importance, proximate mechanisms of intra- and interspecific variability in body size are still not well understood even in such a largely studied group as reptiles. For our study, we concentrated on the gecko species Paroedura picta. We investigated whether differences in sexual size dimorphism and in final and asymptotic snout-vent length (induced by a range of incubation and rearing temperatures) are correlated with differences in the number of presacral vertebrae. Moreover, we tested whether changes in this number were associated with evolutionary changes in sexual size dimorphism and body size in the genus Paroedura. We found that the variation in the number of presacral vertebrae is very limited both intra- and interspecifically, ranging between 26 and 28 vertebrae with most individuals possessing the modal number of 27. We conclude that changes in the number of vertebrae do not contribute to developmental plasticity or evolutionary changes in body size nor, in contrast to some other squamate lineages, to sexual size dimorphism.

Phenotypic plasticity in sex allocation and body size leads to trade-offs between male function and growth in a simultaneously hermaphroditic fish

Phenotypic plasticity in sex allocation enables organisms to maximize reproductive success in variable environments, and thus may generate different sex allocation patterns among populations that experience different mating opportunities. In this experiment, I test whether sex allocation is phenotypically plastic in Serranus tortugarum, a simultaneously hermaphroditic fish, by using reciprocal transplants among four reef study sites with populations at high and low densities and significant differences in sex allocation. Fish transplanted across different densities were predicted to alter sex allocation and body size through trade-offs in investments to somatic growth and male and/or female reproduction. As a control for effects of transplanting, I also transplanted fish across study sites with the same densities and marked and returned fish to their original study sites. As predicted, sex allocation and body size shifted significantly for fish transplanted across different densities but not for those transplanted across the same densities. Separate analyses revealed that the treatment effect on sex allocation was driven strongly by a reduction in male investment by fish transplanted from high to low density, and this reduction in male investment was accompanied by an increase in body size. Fish transplanted from low to high density did not appear to change either male or female investments, but they were smaller than transplants from low to low density. A trade-off between male and female function was not evident, but phenotypic plasticity in body size suggested a trade-off between growth and male function when sex allocation is adjusted. Large-scale empirical tests of sex allocation in the field are relatively rare, and the results of this experiment give novel insights into how animals respond to a change in mating opportunities under natural conditions. The effects of logistical problems associated with fieldwork, such as mortality of experimental animals, are considered in the discussion.

Nonlinear effects of temperature on body form and developmental canalization in the threespine stickleback

Theoretical models predict that nonlinear environmental effects on the phenotype also affect developmental canalization, which in turn can influence the tempo and course of organismal evolution. Here, we used an oceanic population of threespine stickleback (Gasterosteus aculeatus) to investigate temperature-induced phenotypic plasticity of body size and shape using a paternal half-sibling, split-clutch experimental design and rearing offspring under three different temperature regimes (13, 17 and 21 °C). Body size and shape of 466 stickleback individuals were assessed by a set of 53 landmarks and analysed using geometric morphometric methods. At approximately 100 days, individuals differed significantly in both size and shape across the temperature groups. However, the temperature-induced differences between 13 and 17 °C (mainly comprising relative head and eye size) deviated considerably from those between 17 and 21 °C (involving the relative size of the ectocoracoid, the operculum and the ventral process of the pelvic girdle). Body size was largest at 17 °C. For both size and shape, phenotypic variance was significantly smaller at 17 °C than at 13 and 21 °C, indicating that development is most stable at the intermediate temperature matching the conditions encountered in the wild. Higher additive genetic variance at 13 and 21 °C indicates that the plastic response to temperature had a heritable basis. Understanding nonlinear effects of temperature on development and the underlying genetics are important for modelling evolution and for predicting outcomes of global warming, which can lead not only to shifts in average morphology but also to destabilization of development.

Genetic differences and phenotypic plasticity in body size between high‐ and low‐altitude populations of the ground beetle Carabus tosanus

The body size of a univoltine carabid beetle Carabus tosanus on Shikoku Island, Japan, was clearly smaller in higher-altitude populations (subspecies), which possibly represents incipient speciation. To explore the determinants of altitudinal differences in body size in this species, we studied the degree of phenotypic plasticity by conducting rearing experiments at two constant temperatures and examined genetic differences through interpopulation crosses. At 15°C, C.tosanus had a longer developmental period and a shorter adult body than at 20°C. Nevertheless, variation in body size due to temperature effects (phenotypic plasticity) was small compared to the interpopulation differences, which suggests substantial genetic differences between populations (subspecies) at different altitudes. In F(1) offspring from crosses between a low-altitude (subspecies tosanus) and a high-altitude population (subspecies ishizuchianus), adult body length was affected by the genotypes of both parents, with an interaction effect of parental genotype and offspring sex. Further analyses revealed that adult body length was affected by sex-linked factors in addition to autosomal factors. These genetic differences in body size may have resulted from adaptations to different altitudes and may be important for the process of incipient speciation because body size differences could contribute to premating reproductive isolation.

DOES PHENOTYPIC PLASTICITY FOR ADULT SIZE VERSUS FOOD LEVEL IN DROSOPHILA MELANOGASTER EVOLVE IN RESPONSE TO ADAPTATION TO DIFFERENT REARING DENSITIES?

Recent studies with Drosophila have suggested that there is extensive genetic variability for phenotypic plasticity of body size versus food level. If true, we expect that the outcome of evolution at very different food levels should yield genotypes whose adult size show different patterns of phenotypic plasticity. We have tested this prediction with six independent populations of Drosophila melanogaster kept at extreme densities for 125 generations. We found that the phenotypic plasticity of body size versus food level is not affected by selection or the presence of competitors of a different genotype. However, we document increasing among population variation in phenotypic plasticity due to random genetic drift. Several reasons are explored to explain these results including the possibility that the use of highly inbred lines to make inferences about the evolution of genetically variable populations may be misleading.

Thermal phenotypic plasticity of body size in Drosophila melanogaster: sexual dimorphism and genetic correlations

Thirty isofemale lines collected in three different years from the same wild French population were grown at seven different temperatures (12-31 °C). Two linear measures, wing and thorax length, were taken on 10 females and 10 males of each line at each temperature, also enabling the calculation of the wing/thorax (W/T) ratio, a shape index related to wing loading. Genetic correlations were calculated using family means. The W-T correlation was independent of temperature and on average, 0.75. For each line, characteristic values of the temperature reaction norm were calculated, i.e. maximum value, temperature of maximum value and curvature. Significant negative correlations were found between curvature and maximum value or temperature of maximum value. Sexual dimorphism was analysed by considering either the correlation between sexes or the female/male ratio. Female-male correlation was on average 0.75 at the within line, within temperature level but increased up to 0.90 when all temperatures were averaged for each line. The female-male ratio was genetically variable among lines but without any temperature effect. For the female/male ratio, heritability (intraclass correlation) was about 0.20 and evolvability (genetic coefficient of variation) close to 1. Although significant, these values are much less than for the traits themselves. Phenotypic plasticity of sexual dimorphism revealed very similar reaction norms for wing and thorax length, i.e. a monotonically increasing sigmoid curve from about 1.11 up to 1.17. This shows that the males are more sensitive to a thermal increase than females. In contrast, the W/T ratio was almost identical in both sexes, with only a very slight temperature effect.

Phenotypic Plasticity of Body Size in an Insular Population of a Snake

Difference in growth pattern is one of the proximate causes of geographic variation in body size. Such a difference in growth pattern could be explained by two extremes such as genetically determined outcome and phenotypic plasticity or by a combination of these. The Japanese four-lined snake (Elaphe quadrivirgata) on Yakushima Island, Japan, has small body size compared with snakes on the main islands. To examine whether the dwarfism of Yakushima snakes is a consequence of genetic modification or a direct phenotypic response to immediate environmental conditions, E. quadrivirgata from Yakushima and a main island (Shiga Prefecture in Honshu Island), were reared from hatching in a laboratory environment. Data of field surveys were used to examine feeding habits and growth in the wild. Free-ranging snakes on Yakushima had significantly smaller body size and lower growth rate than those in Shiga. Yakushima snakes consumed almost exclusively lizards, whereas frogs were the main prey for Shiga snakes. The proportion of snakes that contained food in their stomach was lower in Yakushima than in Shiga. Available information suggests that the dwarfism of Yakushima snakes cannot be explained by lower survivorship. At hatching, Yakushima snakes of both sexes were larger than Shiga snakes. However, Yakushima and Shiga snakes reached similar body size after approximately 1 year. Captive-reared snakes from the insular dwarf population can reach comparable size to those from the main island, indicating that the growth rate, and hence body size, of E. quadrivirgata is obviously highly plastic. Based on these results, I consider that the dwarfism of Yakushima snakes is attributable to a direct phenotypic response, presumably to food limitation, rather than to microevolutionary changes.

Sexual size dimorphism decreases with temperature in a blowfly, Chrysomya megacephala (Fabricius) (Diptera: Calliphoridae)

1. There is wide intra-specific variation in sexual size dimorphism (SSD). Much of this variation is probably as a result of sexual differences in the selective pressure on body size. However, environmental variables could affect males and females differently, causing variation in SSD. 2. We examined the effects of two temperatures (20 and 30 °C) on SSD in six populations of the blowfly, Chrysomya megacephala. 3. We found that body size increased with temperature in all the populations studied, and the sexes differed in phenotypic plasticity of body size in response to rearing temperature. This created substantial temperature-induced variation in SSD (i.e. sex × temperature interaction). Males were often smaller than females, but the degree of dimorphism was smaller at the higher temperature (30 °C) and larger at the lower temperature (20 °C). This change in SSD was not because of a gender difference in the effect of temperature on development time. Further studies should address whether this variation can be produced by adaptive canalisation of one sex against variation in temperature, or whether it may be a consequence of non-adaptive developmental differences between the sexes. 4. Although most studies assume that the magnitude of SSD is fixed within a species, the present study demonstrates that rearing temperature can generate considerable intra-specific variation in the degree of SSD.

Unusual duplication of the insulin-like receptor in the crustacean Daphnia pulex

BackgroundThe insulin signaling pathway (ISP) has a key role in major physiological events like carbohydrate metabolism and growth regulation. The ISP has been well described in vertebrates and in a few invertebrate model organisms but remains largely unexplored in non-model invertebrates. This study is the first detailed genomic study of this pathway in a crustacean species, Daphnia pulex.ResultsThe Daphnia pulex draft genome sequence assembly was scanned for major components of the ISP with a special attention to the insulin-like receptor. Twenty three putative genes are reported. The pathway appears to be generally well conserved as genes found in other invertebrates are present. Major findings include a lower number of insulin-like peptides in Daphnia as compared to other invertebrates and the presence of multiple insulin-like receptors (InR), with four genes as opposed to a single one in other invertebrates. Genes encoding for the Dappu_InR are likely the result of three duplication events and bear some unusual features. Dappu_InR-4 has undergone extensive evolutionary divergence and lacks the conserved site of the catalytic domain of the receptor tyrosine kinase. Dappu_InR-1 has a large insert and lacks the transmembranal domain in the β-subunit. This domain is also absent in Dappu_InR-3. Dappu_InR-2 is characterized by the absence of the cystein-rich region. Real-time q-PCR confirmed the expression of all four receptors. EST analyses of cDNA libraries revealed that the four receptors were differently expressed under various conditions.ConclusionsDuplications of the insulin receptor genes might represent an important evolutionary innovation in Daphnia as they are known to exhibit extensive phenotypic plasticity in body size and in the size of defensive structures in response to predation.

Environmental effects on sexual size dimorphism of a seed-feeding beetle

Sexual size dimorphism is widespread in animals but varies considerably among species and among populations within species. Much of this variation is assumed to be due to variance in selection on males versus females. However, environmental variables could affect the development of females and males differently, generating variation in dimorphism. Here we use a factorial experimental design to simultaneously examine the effects of rearing host and temperature on sexual dimorphism of the seed beetle, Callosobruchus maculatus. We found that the sexes differed in phenotypic plasticity of body size in response to rearing temperature but not rearing host, creating substantial temperature-induced variation in sexual dimorphism; females were larger than males at all temperatures, but the degree of this dimorphism was smallest at the lowest temperature. This change in dimorphism was due to a gender difference in the effect of temperature on growth rate and not due to sexual differences in plasticity of development time. Furthermore, the sex ratio (proportion males) decreased with decreasing temperature and became female-biased at the lowest temperature. This suggests that the temperature-induced change in dimorphism is potentially due to a change in non-random larval mortality of males versus females. This most important implication of this study is that rearing temperature can generate considerable intraspecific variation in the degree of sexual size dimorphism, though most studies assume that dimorphism varies little within species. Future studies should focus on whether sexual differences in phenotypic plasticity of body size are a consequence of adaptive canalization of one sex against environmental variation in temperature or whether they simply reflect a consequence of non-adaptive developmental differences between males and females.

Correlated responses to artificial body size selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies

Most life history traits are positively influenced by body size, whereas disadvantages of large body size are poorly documented. To investigate presumed intrinsic costs of large size in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae), we established two replicates each of three body size laboratory selection lines (small, control and large; selection on males only), and subjected flies of the resulting extended body size range to various abiotic stresses. Response to selection was symmetrical in the small and large lines (realized h(2) = 0.16-0.18). After 24 generations of selection body size had changed by roughly 10%. Female size showed a correlated response to selection on male size, whereas sexual size dimorphism did not change. Development time also showed a correlated response as, similar to food limited flies, small line flies emerged earlier at smaller body size. At the lowest larval food limit possible, flies of all lines emerged at the same small body size after roughly the same development time; so overall phenotypic plasticity in body size and development time strongly increased following selection. Juvenile mortality increased markedly when food was extremely limited, large line flies showing highest mortality. Winter frost disproportionately killed large (line) flies because of their longer development times. Mortality at high temperatures was high but size-selective effects were inconsistent. In all environments the larger males suffered more. Initial growth rate was higher for males and at unlimited food. Small line individuals of both sexes grew slowest at unlimited larval food but fastest at limited larval food, suggesting a physiological cost of fast growth. Overall, extension of the natural body size range by artificial selection revealed some otherwise cryptic intrinsic juvenile viability costs of large size, mediated by longer development or faster growth, but only in stressful environments.

The cellular basis for phenotypic plasticity of body size in Western Spadefoot toad (Spea hammondi) tadpoles: patterns of cell growth and recruitment in response to food and temperature manipulations

Body size is highly variable within and among populations, both as a result of genetic variation and as a plastic response to environmental variation. From a proximate perspective, body size depends upon cell size, cell number, and extracellular matrix, but we know little about their independent contributions to size nor how these contributions vary with environmental influences. Here, I introduce the tail muscle of anuran tadpoles as a new system for studying this issue. Body size and tail size of tadpoles is sensitive to variation in food and temperature. I show first that tail muscle size is strongly correlated with overall body size, thus making it a good tissue to study size regulation. Second, the relative role of cell size and cell number, but not extracellular matrix, shows an interaction between food and temperature treatments and across ages. For example, in young tadpoles food effects on size are due exclusively to cell size at low temperatures but both cell size and number at high temperatures. This pattern partially reverses for older tadpoles. Despite the complexity of this interaction, the two populations compared show nearly identical patterns, suggesting that the plastic response is robust.

Phenotypic plasticity of body size in a temperate population ofDrosophila melanogaster: When the temperature—size rule does not apply

A natural population of Drosophila melanogaster in southern France was sampled in three different years and 10 isofemale lines were investigated from each sample. Two size-related traits, wing and thorax length, were measured and the wing/thorax ratio was also calculated. Phenotypic plasticity was analysed after development at seven different constant temperatures, ranging from 12 degrees C to 31 degrees C. The three year samples exhibited similar reaction norms, suggesting a stable genetic architecture in the natural population. The whole sample (30 lines) was used to determine precisely the shape of each reaction norm, using a derivative analysis. The practical conclusion was that polynomial adjustments could be used in all cases, but with different degrees: linear for the wing/thorax ratio, quadratic for thorax length, and cubic for wing length. Both wing and thorax length exhibited concave reaction norms, with a maximum within the viable thermal range. The temperatures of the maxima were, however, quite different, around 15 degrees C for the wing and 19.5 degrees C for the thorax. Assuming that thorax length is a better estimate of body size, it is not possible to state that increasing the temperature results in monotonically decreasing size (the temperature-size rule), although this is often seen to be the case for genetic variations in latitudinal clines. The variability of the traits was investigated at two levels-within and between lines-and expressed as a coefficient of variation. The within-line (environmental) variability revealed a regular, quadratic convex reaction norm for the three traits, with a minimum around 21 degrees C. This temperature of minimum variability may be considered as a physiological optimum, while extreme temperatures are stressful. The between-line (genetic) variability could also be adjusted to quadratic polynomials, but the curvature parameters were not significant. Our results show that the mean values of the traits and their variance are both plastic, but react in different ways along a temperature gradient. Extreme low or high temperatures decrease the size but increase the variability. These effects may be considered as a functional response to environmental stress.

Allometry for Sexual Size Dimorphism: Testing Two Hypotheses for Rensch’s Rule in the Water StriderAquarius remigis

Within any given clade, male size and female size typically covary, but male size often varies more than female size. This generates a pattern of allometry for sexual size dimorphism (SSD) known as Rensch's rule. I use allometry for SSD among populations of the water strider Aquarius remigis (Hemiptera, Gerridae) to test the hypothesis that Rensch's rule evolves in response to sexual selection on male secondary sexual traits and an alternative hypothesis that it is caused by greater phenotypic plasticity of body size in males. Comparisons of three populations reared under two temperature regimes are combined with an analysis of allometry for genital and somatic components of body size among 25 field populations. Contrary to the sexual-selection hypothesis, genital length, the target of sexual selection, shows the lowest allometric slope of all the assayed traits. Instead, the results support a novel interpretation of the differential-plasticity hypothesis: that the traits most closely associated with reproductive fitness (abdomen length in females and genital length in males) are "adaptively canalized." While this hypothesis is unlikely to explain Rensch's rule among species or higher clades, it may explain widespread patterns of intraspecific variation in SSD recently documented for many insect species.

Phenotypic plasticity of body size at different temperatures in a planktonic rotifer: mechanisms and adaptive significance

Summary Larger body size at low temperatures is a commonly observed phenomenon in ectothermic organisms. The mechanisms that lead to this pattern and its possible adaptive significance were studied in laboratory experiments using the parthenogenetically reproducing rotifer Synchaeta pectinata. At low temperatures of 4 °C mean body volume was 46% larger than in individuals cultured at 12 °C. Egg volume was 35% larger in low vs high temperatures. Larger body size at low temperatures was caused by two mechanisms. First, when exposed to low temperatures, mothers laid larger eggs and the hatchlings of these eggs developed into larger adults (irrespective of temperature). Second, individuals cultured at low temperatures grew to a larger body size during their juvenile phase. The former mechanism had a greater influence on adult size than the latter. The production of larger eggs at low temperatures seemed to be due to a higher reproductive investment into individual offspring as it occurred independently of differences in maternal size. Life table experiments showed that offspring from small eggs (produced at high temperatures) had a significantly higher population growth rate than offspring from large eggs, when cultured at high temperatures. This was mainly due to an increase in fertility during the first days of adult life.

Phenotypic plasticity of body size in Drosophila: effects of a daily periodicity of growth temperature in two sibling species

Abstract. Variation of wing and thorax length under thermoperiodic growth conditions was analysed in four strains of two sibling species, Drosophila melanogaster and D. simulans, from two European localities. Results were compared to those obtained with constant temperatures ranging from 12 to 31 °C.Under constant temperatures the data basically confirmed previous results: concave reaction norms for wing and thorax length; a monotonically decreasing norm for wing : thorax ratio; and an increasing norm for sex dimorphism (female : male ratio). Phenotypic variability was maximum at extreme temperatures and minimum at middle ones. Slight differences were observed according to the geographical origin: the difference between strains from Bordeaux (France) and Cordoba (Spain) was maximum at low temperatures but disappeared at about 28 °C.According to the temperatures chosen, alternating thermal regimens had either no effect or produced a significant size reduction, probably reflecting a periodic stress. The magnitude of this effect was proportional to the amplitude of the thermoperiod but not to the quality (cold or heat) of the stress. In a similar way, the wing : thorax ratio was either not modified or reduced significantly, indicating that wing length was relatively more affected than thorax length by alternating thermal regimens. Sex dimorphism also showed either no change or a significant increase, indicating that males were relatively more reactive than females to alternating conditions. Finally, regimens of broad amplitudes increased the phenotypic variability, again an indication of stressful effects. All these observations should be taken into account when analysing phenotypic variability in nature and trying to understand natural selection in wild‐living populations.

Genotype–environment interaction and the ontogeny of diet-induced phenotypic plasticity in size and shape of Melanoplus femurrubrum (Orthoptera: Acrididae)

Abstract The developmental origin of phenotypic plasticity in morphological shape can be attributed to environment-specific changes in growth of overall body size, localized growth of a morphological structure or a combination of both. I monitored morphological development in the first four nymphal instars of grasshoppers (Melanoplus femurrubrum) raised on two different plant diets to determine the ontogenetic origins of diet-induced phenotypic plasticity and to quantify genetic variation for phenotypic plasticity. I measured diet-induced phenotypic plasticity in body size (tibia length), head size (articular width and mandible depth) and head shape (residual articular width and residual mandible depth) for grasshoppers from 37 full-sib families raised on either a hard plant diet (Lolium perenne) or a soft plant diet (Trifolium repens). By the second to third nymphal instar, grasshoppers raised on a hard plant diet had significantly smaller mean tibia length and greater mean residual articular width (distance between mandibles adjusted for body size) compared with full-sibs raised on a soft plant diet. However, there was no significant phenotypic plasticity in mean unadjusted articular width and mandible depth, and in mean residual mandible depth. At the population level, development of diet-induced phenotypic plasticity in grasshopper head shape is mediated by plastic changes in allocation to tissue growth that maintain growth of head size on hard, low-nutrient diets while reducing growth of body size. Within the population, there was substantial variation in the plasticity of growth trajectories since different full-sib families developed phenotypic plasticity of residual articular width through different combinations of head and body size growth. Genetic variation for diet-induced phenotypic plasticity of residual articular width, residual mandible depth and tibia length, as estimated by genotype–environment interaction, exhibited significant fluctuation through ontogeny (repeated measures MANOVA, family × plant × instar, P < 0.01). For example, there was significant genetic variation for phenotypic plasticity of residual articular width in the third nymphal instar, but not earlier or later in ontogeny. The observed patterns of genetic variation are discussed with reference to short-term constraints and the evolution of phenotypic plasticity.

ECTOTHERMS FOLLOW THE CONVERSE TO BERGMANN'S RULE.

ECTOTHERMS FOLLOW THE CONVERSE TO BERGMANN'S RULE.