昆虫翅型分化的表型可塑性机制

翅多型现象在昆虫中广泛存在,是昆虫在飞行扩散和繁殖能力之间权衡的一种策略,对种群的环境适应性进化具有重要的意义。目前在植食性昆虫中研究较多,有关寄生蜂的翅型分化鲜见报道。综述了昆虫翅型分化的表型可塑性机制。遗传因素和环境因素均对昆虫翅的发育产生影响,基因型对翅型的决定具有显著作用,外界环境条件,包括温度、光周期、食物质量、自身密度、外源激素等因素对昆虫翅的发育也产生重要的调节作用,从而产生翅的非遗传多型性现象。此外,天敌的寄生或捕食作用可能会诱导某些昆虫的翅型产生隔代表型变化。对昆虫产生翅多型现象的生态学意义及其在生物进化过程中的作用进行了讨论,并探讨了寄生性昆虫翅型分化机制在生物防治上的可能应用途径。功能基因组学和表观遗传学的进一步发展可望为彻底揭示昆虫翅型分化机制提供新的机遇和技术手段。     

Winged presoldiers induced by a juvenile hormone analog in Zootermopsis nevadensis: implications for plasticity and evolution of caste differentiation in termites

To elucidate the switching mechanism of caste differentiation in termites and to examine the possible induction of soldier-reproductive intercastes experimentally, we investigated the effects of juvenile hormone on the morphologies of soldier caste by applying a juvenile hormone analog (JHA) to nymphs of the damp-wood termite Zootermopsis nevadensis (Isoptera : Termopsidae). JHA treatment for about 2 weeks induced a variety of intermediate castes, showing both alate and soldier morphological features. The principal component analysis (PCA) of those morphological characters showed that those intercastes were a deviation from the developmental line into alates to soldier differentiation, which is known to be triggered by juvenile hormone. Detailed morphological examination of the compound eyes, wing joint, and mandibles showed that those intercastes expressed soldier features, although they had started to develop alate characteristics. The morphology of the resultant intercastes seemed to be determined by the nymphal stage, at which JHA treatment was applied. The induced intercastes with exaggerated soldier-specific characteristics (e.g., mandibles) repressed alate-specific characteristics (e.g., wings), namely, the alate and soldier morphological characteristics in induced intercastes show opposite responses against the application of JHA. On the other hand, ovarian development was not suppressed by the JHA application, even in the soldier-like individuals. Naturally differentiated presoldiers also possessed developed ovarioles, although ovaries of mature soldiers were degenerated. Our results suggest that the juvenile hormone plays complicated roles in the expression of caste morphologies and ovarian development in termites.     

Insights into the dynamics of hind leg development in honey bee (Apis mellifera L.) queen and worker larvae - A morphology/differential gene expression analysis

Phenotypic plasticity is a hallmark of the caste systems of social insects, expressed in their life history and morphological traits. These are best studied in bees. In their co-evolution with angiosperm plants, the females of corbiculate bees have acquired a specialized structure on their hind legs for collecting pollen. In the highly eusocial bees (Apini and Meliponini), this structure is however only present in workers and absent in queens. By means of histological sections and cell proliferation analysis we followed the developmental dynamics of the hind legs of queens and workers in the fourth and fifth larval instars. In parallel, we generated subtractive cDNA libraries for hind leg discs of queen and worker larvae by means of a Representational Difference Analysis (RDA). From the total of 135 unique sequences we selected 19 for RT-qPCR analysis, where six of these were confirmed as differing significantly in their expression between the two castes in the larval spinning stage. The development of complex structures such as the bees’ hind legs, requires diverse patterning mechanisms and signaling modules, as indicated by the set of differentially expressed genes related with cell adhesion and signaling pathways.     

Life cycles,plasticity and palaeoecology in temnospondyl amphibians

In the largest early tetrapod clade, the temnospondyls, ontogenies were diverse and quite distinct from the life cycles of extant amphibians. Three well‐studied clades exemplify the diversity of these long‐extinct ontogenies, here analysed with respect to their bearing on developmental plasticity, reaction norms and evolution. Sclerocephalus readily adjusted by means of developmental evolution to different lake environments. In addition, plasticity (reaction norm) played a significant role, apparent both morphologically and by altered developmental traits. Size increase and extension of the ontogenetic trajectory gave larger predators, a phenomenon also found in the dissorophoid Micromelerpeton. Whereas Sclerocephalus was throughout preying on the same fishes, Micromelerpeton was able to fit into different trophic levels. In the branchiosaurid Apateon, a biphasic life cycle was established, with metamorphosis producing a terrestrial morph in some species; truncation of the ontogenetic trajectory gave a sexually mature larva as an alternative morph (neoteny). Plasticity was high in the larval morphs, permitting neotenes to live as filter feeders or small carnivores. Fine‐tuning of development permitted Apateon populations to adjust to specific lake properties and readily change from a filter‐feeding to carnivorous mode of life. In the nonmetamorphosing Triassic Gerrothorax, morphology was extremely conserved, but histology reveals much plasticity at the microscopical level, correlating with fluctuating salinity and water energy. In responding to environmental fluctuations by enhanced plasticity, the studied temnospondyls managed to populate lakes inhabitable to other tetrapods and fishes.     

Temperature,size and developmental plasticity in birds

As temperatures increase, there is growing evidence that species across much of the tree of life are getting smaller. These climate change-driven size reductions are often interpreted as a temporal analogue of the observation that individuals within a species tend to be smaller in the warmer parts of the species'' range. For ectotherms, there has been a broad effort to understand the role of developmental plasticity in temperature–size relationships, but in endotherms, this mechanism has received relatively little attention in favour of selection-based explanations. We review the evidence for a role of developmental plasticity in warming-driven size reductions in birds and highlight insulin-like growth factors as a potential mechanism underlying plastic responses to temperature in endotherms. We find that, as with ectotherms, changes in temperature during development can result in shifts in body size in birds, with size reductions associated with warmer temperatures being the most frequent association. This suggests developmental plasticity may be an important, but largely overlooked, mechanism underlying warming-driven size reductions in endotherms. Plasticity and natural selection have very different constraining forces, thus understanding the mechanism linking temperature and body size in endotherms has broad implications for predicting future impacts of climate change on biodiversity.     

Diet and a developmental time constraint alter life‐history trade‐offs in a caddis fly (Trichoptera: Limnephilidae)

Environmental factors influence variation in life histories by affecting growth, development, and reproduction. We conducted an experiment in outdoor mesocosms to examine how diet and a time constraint on juvenile development (pond‐drying) influence life‐history trade‐offs (growth, development, adult body mass) in the caddis fly Limnephilus externus (Trichoptera: Limnephilidae). We predicted that: (1) diet supplementation would accelerate larval growth and development, and enhance survival to adulthood; (2) pond‐drying would accelerate development and increase larval mortality; and (3) the relationship between adult mass and age at maturity would be negative. Diet supplementation did lead to larger adult mass under nondrying conditions, but did not significantly alter growth or development rates. Contrary to predictions, pond‐drying reduced growth rates and delayed development. The slope (positive or negative) of the female mass–age at maturity relationship depended on interactions with diet or pond‐drying, but the male mass–age relationship was negative and independent of treatment. Our results suggest that pond‐drying can have negative effects on the future fitness of individuals by increasing the risk of desiccation‐induced, pre‐reproductive mortality and decreasing adult body size at maturity. These negative effects on life history cannot be overcome with additional nutritional resources in this species. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 495–504.     

Developmental plasticity and acclimation both contribute to adaptive responses to alternating seasons of plenty and of stress in <Emphasis Type="Italic">Bicyclus</Emphasis> butterflies

Plasticity is a crucial component of the life cycle of invertebrates that live as active adults throughout wet and dry seasons in the tropics. Such plasticity is seen in the numerous species of Bicyclus butterflies in Africa which exhibit seasonal polyphenism with sequential generations of adults with one or other of two alternative phenotypes. These differ not only in wing pattern but in many other traits. This divergence across a broad complex of traits is associated with survival and reproduction either in a wet season that is favourable in terms of resources, or mainly in a dry season that is more stressful. This phenomenon has led us to examine the bases of the developmental plasticity in a model species, B. anynana, and also the evolution of key adult life history traits, including starvation resistance and longevity. We now understand something about the processes that generate variation in the phenotype, and also about the ecological context of responses to environmental stress. The responses clearly involve a mix of developmental plasticity as cued by different environments in pre-adult development, and the acclimation of life history traits in adults to their prevailing environment.     

Thermal evolution of pre-adult life history traits, geometric size and shape, and developmental stability in Drosophila subobscura

Replicated lines of Drosophila subobscura originating from a large outbred stock collected at the estimated Chilean epicentre (Puerto Montt) of the original New World invasion were allowed to evolve under controlled conditions of larval crowding for 3.5 years at three temperature levels (13, 18 and 22 degrees C). Several pre-adult life history traits (development time, survival and competitive ability), adult life history related traits (wing size, wing shape and wing-aspect ratio), and wing size and shape asymmetries were measured at the three temperatures. Cold-adapted (13 degrees C) populations evolved longer development times and showed lower survival at the highest developmental temperature. No divergence for wing size was detected following adaptation to temperature extremes (13 and 22 degrees C), in agreement with earlier observations, but wing shape changes were obvious as a result of both thermal adaptation and development at different temperatures. However, the evolutionary trends observed for the wing-aspect ratio were inconsistent with an adaptive hypothesis. There was some indication that wing shape asymmetry has evolutionarily increased in warm-adapted populations, which suggests that there is additive genetic variation for fluctuating asymmetry and that it can evolve under rapid environmental changes caused by thermal stress. Overall, our results cast strong doubts on the hypothesis that body size itself is the target of selection, and suggest that pre-adult life history traits are more closely related to thermal adaptation.     

Tropical dry and wet season polyphenism in the butterfly Melanitis leda (Satyrinae): Phenotypic plasticity and climatic correlates

Changes in wing pattern, colour, shape and size associated with seasonal polyphenism in Melanitis leda were quantified using a series of 155 butterflies collected by N. Manders on Mauritius in 1905. Butterflies of the wetter period were predominantly of the wet season form with large, well differentiated eyespots, short tails, smaller wings and a characteristic background colour. The dry season form occurred only in the drier period and has much smaller eyespots, longer tails, larger wings and a variable background. Many intermediates occurred, mainly in the drier period. These are associated with an absence of extreme seasonal change in Mauritius. The first principal component (PCI) describing the morphometric and colour data is closely related to the wing form (r = 0.80). Regression analyses using temperature and rainfall data for the 8 weeks before each capture showed that about 40% of variation in PCI could be accounted for by temperature in weeks 2–3 before capture. Many of the characters measured are redundant; a subset of seven morphometric characters yields a closely similar PCI. Analysis of is subset in an additional sample of 70 M. leda from Kenya showed that the seasonal polyphenism overrides a small degree of sexual dimorphism. The results are discussed with regard to seasonal changes in adult activity, resting backgrounds and visual predation. Wing phenotype characters are part of an array of coordinated morphological and life history traits which include ovarian dormancy and fat body development in dry season adults. A partial independence occurs in the proximal control of these traits as indicated, for example, by the larger wing and tail size, and smaller eyespots of the small number of the wet season form captured in the drier period in comparison to those of the wetter months.     

Heterochronic developmental shift caused by thyroid hormone in larval sand dollars and its implications for phenotypic plasticity and the evolution of nonfeeding development

Recent work on a diverse array of echinoderm species has demonstrated, as is true in amphibians, that thyroid hormone (TH) accelerates development to metamorphosis. Interestingly, the feeding larvae of several species of sea urchins seem to obtain TH through their diet of planktonic algae (exogenous source), whereas nonfeeding larvae of the sand dollar Peronella japonica produce TH themselves (endogenous source). Here we examine the effects of TH (thyroxine) and a TH synthesis inhibitor (thiourea) on the development of Dendraster excentricus, a sand dollar with a feeding larva. We report reduced larval skeleton lengths and more rapid development of the juvenile rudiment in the exogenous TH treatments when compared to controls. Also, larvae treated with exogenous TH reached metamorphic competence faster at a significantly reduced juvenile size, representing the greatest reduction in juvenile size ever reported for an echinoid species with feeding larvae. These effects of TH on D. excentricus larval development are strikingly similar to the phenotypically plastic response of D. excentricus larvae reared under high food conditions. We hypothesize that exogenous (algae-derived) TH is the plasticity cue in echinoid larvae, and that the larvae use ingested TH levels as an indicator for larval nutrition, ultimately signaling the attainment of metamorphic competence. Furthermore, our experiments with the TH synthesis inhibitor thiourea indicate that D. excentricus larvae can produce some TH endogenously. Endogenous TH production might, therefore, be a shared feature among sand dollars, facilitating the evolution of nonfeeding larval development in that group. Mounting evidence on the effects of thyroid hormones in echinoderm development suggests life-history models need to incorporate metamorphic hormone effects and the evolution of metamorphic hormone production.     

植物表型可塑性研究进展

表型可塑性已成为生态进化发育生物学的核心概念,很大程度上由于植物可塑性研究的主要贡献,但人们仍远未完全了解表型可塑性的原因和结果。从整体角度理出表型可塑性研究发展的基本脉络,介绍研究内容、途径和简史,聚焦于几个主要方面的研究进展及发展方向。现代可塑性研究的兴盛始于关于可塑性的进化学重要性的一篇综述,从现象的描述、对其遗传基础和可塑性本身进化的讨论,发展到探索其背后的发育机制、植物生长与适应策略、生态学影响等。未来可塑性研究应在重新理解和评价表型可塑性及其适应性的基础上,更关注自然条件下环境因子和可塑响应的复杂性。表型可塑性的生态-进化学意义仍将是未来研究的重点。     

Ergatoid queen development in the ant Myrmecina nipponica: modular and heterochronic regulation of caste differentiation

Caste polyphenism in social insects provides us with excellent opportunities to examine the plasticity and robustness underlying developmental pathways. Several ant species have evolved unusual castes showing intermediate morphologies between alate queens and wingless workers. In some low-temperature habitats, the ant Myrmecina nipponica produces such intermediate reproductives (i.e. ergatoids), which can mate and store sperm but cannot fly. To gain insight into the developmental and evolutionary aspects associated with ergatoid production, we conducted morphological and histological examinations of the post-embryonic development of compound eyes, gonads and wings during the process of caste differentiation. In compound eyes, both the queen-worker and ergatoid-worker differences were already recognized at the third larval instar. In gonads, queen-worker differentiation began at the larval stage, and ergatoid-worker differentiation began between the prepupal and pupal stages. Wing development in ergatoids was generally similar to that in workers throughout post-embryonic development. Our results showed that the developmental rate and timing of differentiation in body parts differed among castes and among body parts. These differences suggest that the rearrangement of modular body parts by heterochronic developmental regulation is responsible for the origination of novel castes, which are considered to be adaptations to specific ecological niches.     

Toward a population genetic framework of developmental evolution: the costs,limits, and consequences of phenotypic plasticity

Adaptive phenotypic plasticity allows organisms to cope with environmental variability, and yet, despite its adaptive significance, phenotypic plasticity is neither ubiquitous nor infinite. In this review, we merge developmental and population genetic perspectives to explore costs and limits on the evolution of plasticity. Specifically, we focus on the role of modularity in developmental genetic networks as a mechanism underlying phenotypic plasticity, and apply to it lessons learned from population genetic theory on the interplay between relaxed selection and mutation accumulation. We argue that the environmental specificity of gene expression and the associated reduction in pleiotropic constraints drive a fundamental tradeoff between the range of plasticity that can be accommodated and mutation accumulation in alternative developmental networks. This tradeoff has broad implications for understanding the origin and maintenance of plasticity and may contribute to a better understanding of the role of plasticity in the origin, diversification, and loss of phenotypic diversity.     

Ageing in a eusocial insect: molecular and physiological characteristics of life span plasticity in the honey bee

Commonly held views assume that ageing, or senescence, represents an inevitable, passive, and random decline in function that is strongly linked to chronological age. In recent years, genetic intervention of life span regulating pathways, for example, in Drosophila as well as case studies in non-classical animal models, have provided compelling evidence to challenge these views.Rather than comprehensively revisiting studies on the established genetic model systems of ageing, we here focus on an alternative model organism with a wild type (unselected genotype) characterized by a unique diversity in longevity - the honey bee.Honey bee (Apis mellifera) life span varies from a few weeks to more than 2 years. This plasticity is largely controlled by environmental factors. Thereby, although individuals are closely related genetically, distinct life histories can emerge as a function of social environmental change.Another remarkable feature of the honey bee is the occurrence of reverted behavioural ontogeny in the worker (female helper) caste. This behavioural peculiarity is associated with alterations in somatic maintenance functions that are indicative of reverted senescence. Thus, although intraspecific variation in organismal life span is not uncommon, the honey bee holds great promise for gaining insights into regulatory pathways that can shape the time-course of ageing by delaying, halting or even reversing processes of senescence. These aspects provide the setting of our review.We will highlight comparative findings from Drosophila melanogaster and Caenorhabditis elegans in particular, and focus on knowledge spanning from molecular- to behavioural-senescence to elucidate how the honey bee can contribute to novel insights into regulatory mechanisms that underlie plasticity and robustness or irreversibility in ageing.     

Honey supplementation and its developmental consequences: evidence for food limitation in a paper wasp, Polistes metricus

ABSTRACT.

• 1 Polistes metricus Say (Hymenoptera: Vespidae, Polistinae) field colonies were supplemented with dilute honey during the pre-emergence and early post-emergence phases of colony development.
• 2 Supplementation did not increase number of nest cells constructed or rates of loss to predation or foundress disappearance compared with controls.
• 3 Colonies receiving honey supplementation produced first offspring earlier in the season than control colonies. The difference is due to a shorter time span between founding and first emergence.
• 4 Foundresses of supplemented and control colonies did not differ in wing length or per cent body fat.
• 5 Offspring of both supplemented and control colonies had shorter wing lengths than did foundresses of supplemented colonies. Offspring of control colonies had shorter wing lengths than did foundresses of control colonies but not foundresses of supplemented colonies.
• 6 The per cent body fat of offspring from control colonies was lower than that of all foundresses and of offspring from supplemented colonies.
• 7 The per cent body fat of offspring from supplemented colonies was higher than that of all foundresses.

     

Plasticity and artificial selection for developmental mode in a poecilogonous sea slug

The contribution of phenotypically plastic traits to evolution depends on the degree of environmental influence on the target of selection (the phenotype) as well as the underlying genetic structure of the trait and plastic response. Likewise, maternal effects can help or hinder evolution through affects to the response to selection. The sacoglossan sea slug Alderia willowi exhibits intraspecific variation for developmental mode (= poecilogony) that is environmentally modulated with populations producing more yolk‐feeding (lecithotrophic) larvae during the summer, and more planktonic‐feeding (planktotrophic) larvae in the winter. I found significant family‐level variation in the reaction norms between 17 maternal families of A. willowi when reared in a split‐brood design in low (16 ppt) versus high (32 ppt) salinity, conditions which mimic seasonal variation in salinity of natural populations. I documented a significant response to selection for lecithotrophic larvae in high and low salinity. The slope of the reaction norm was maintained following one generation of selection for lecithotrophy. When the maternal environment was controlled in the laboratory, I found significant maternal effects, which reduced the response to selection. These results suggest there is standing genetic variation for egg‐mass type in A. willowi, but the ability of selection to act on that variation may depend on the environment in which the phenotype is expressed in preceding generations.     

Simulating plasticity as a framework for understanding habitat selection and its role in adaptive capacity and extinction risk through an expansion of CDMetaPOP

Adaptive capacity can present challenges for modelling as it encompasses multiple ecological and evolutionary processes such as natural selection, genetic drift, gene flow and phenotypic plasticity. Spatially explicit, individual-based models provide an outlet for simulating these complex interacting eco-evolutionary processes. We expanded the existing Cost-Distance Meta-POPulation (CDMetaPOP) framework with inducible plasticity modelled as a habitat selection behaviour, using temperature or habitat quality variables, with a genetically based selection threshold conditioned on past individual experience. To demonstrate expected results in the new module, we simulated hypothetical populations and then evaluated model performance in populations of redband trout (Oncorhynchus mykiss gairdneri) across three watersheds where temperatures induce physiological stress in parts of the stream network. We ran simulations using projected warming stream temperature data under four scenarios for alleles that: (1) confer thermal tolerance, (2) bestow plastic habitat selection, (3) give both thermal tolerance and habitat selection preference and (4) do not provide either thermal tolerance or habitat selection. Inclusion of an adaptive allele decreased declines in population sizes, but this impact was greatly reduced in the relatively cool stream networks. As anticipated with the new module, high-temperature patches remained unoccupied by individuals with the allele operating plastically after exposure to warm temperatures. Using complete habitat avoidance above the stressful temperature threshold, habitat selection reduced the overall population size due to the opportunity cost of avoiding areas with increased, but not guaranteed, mortality. Inclusion of plasticity within CDMetaPOP will provide the potential for genetic or plastic traits and ‘rescue’ to affect eco-evolutionary dynamics for research questions and conservation applications.     

Sex-specific plasticity of growth and maturation size in a spider: implications for sexual size dimorphism

Sex-specific plasticity in body size has been recently proposed to cause intraspecific patterns of variation in sexual size dimorphism (SSD). We reared juvenile male and female Mediterranean tarantulas (Lycosa tarantula) under two feeding regimes and monitored their growth until maturation. Selection gradients calculated across studies show how maturation size is under net stabilizing selection in females and under directional selection in males. This pattern was used to predict that body size should be more canalized in females than in males. As expected, feeding affected male but not female maturation size. The sex-specific response of maturation size was related to a dramatic divergence between subadult male and female growth pathways. These results demonstrate the existence of sex-specific canalization and resource allocation to maturation size in this species, which causes variation in SSD depending on developmental conditions consistent with the differential-plasticity hypothesis explaining Rensch's Rule.