Form Of Phenotypic Plasticity Research Articles

A novel type of neighbour perception elicits reproductive plasticity in an annual plant with a mixed mating system.

Plants display various forms of phenotypic plasticity in anticipation of changing conditions, many of which are influenced by information obtained from neighbouring plants. Here, we tested the hypothesis that cleistogamic Lamium amplexicaule plants can adaptively modify production of chasmogamous (CH) and cleistogamous (CL) flowers based on the perception of conspecific neighbours. The production and proportion of CH and CL flowers was examined in individual L. amplexicaule grown at varying densities or treated with root leachates from plants grown at different densities. When growing at high density or treated with root leachates from high-density pots, L. amplexicaule increased production of more expensive, potentially outcrossing CH flowers. In contrast, single plants or plants treated with root leachates from empty pots or single-source plants predominantly developed cheaper, self-pollinated CL flowers. The results demonstrate a novel root-based neighbour-perception modality that enables plants to adaptively adjust production of CH and CL flowers in response to the presence of potential reproductive partners. Further research is needed to explore the broader ecological implications of this novel interplant cueing on reproductive bet-hedging and plasticity in natural settings, as well as to identify the involved cues and their mode of operation.

Plasticity and the adaptive evolution of switchlike reaction norms under environmental change.

Phenotypic plasticity is often posited as an avenue for adaptation to environmental change, whereby environmental influences on phenotypes could shift trait expression toward new optimal values. Conversely, plastic trait expression may inhibit adaptation to environmental change by reducing selective pressure on ill-adapted traits. While plastic responses are often assumed to be linear, nonlinear phenotype-environment relationships are common, especially in thermally sensitive traits. Here we examine nonlinear plasticity in a trait with great ecological and evolutionary significance: sexual phenotype in species with environmental sex determination (ESD). In species with ESD, development switches between male and female at an environmental threshold (the inflection point). The inflection point is a key trait for adaptive responses to changing environments and should evolve toward the new optimum in order to maintain evolutionarily stable sex ratios. We used an individual-based theoretical model to investigate how two forms of plasticity in the ESD reaction norm-the nonlinear slope of the reaction norm and a linear shift in the inflection point-influence the evolution of the inflection point under climate warming. We found that steeper reaction norm slopes (high nonlinear plasticity) promoted evolution toward new optimal phenotypes (higher inflection points). In contrast, increased linear plasticity in the inflection point (shift) hindered adaptive evolution. Additionally, populations in moderate warming scenarios showed greater adaptive evolution of the inflection point compared with populations in extreme warming scenarios, suggesting that the proximity of existing phenotypes to new optimal phenotypes influences evolutionary outcomes. Unexpectedly, we found greater population persistence under high climate variability, due to the increased production of rare-sex individuals in unusually cold years. Our results demonstrate that different forms of phenotypic plasticity have crucially different effects on adaptive evolution. Plasticity that prevented sex ratio bias hindered the evolution of the inflection point, while plasticity that exacerbated sex ratio bias promoted adaptation to environmental change.

Warmer and more acidic conditions enhance performance of an endemic low shore gastropod.

Changing ocean temperatures are predicted to challenge marine organisms, especially when combined with other factors, such as ocean acidification. Acclimation, as a form of phenotypic plasticity, can however, moderate the consequences of changing environments for biota. Our understanding of how altered temperature and acidification together influence species acclimation responses is, however, limited compared to responses to single stressors. This study investigated how temperature and acidification affected the thermal tolerance and righting speed of the Girdled Dogwhelk, Trochia cingulata (Linnaeus, 1771). Whelks were acclimated for two weeks to combinations of three temperatures (11°C: cold, 13°C: moderate and 15°C: warm) and two pH regimes (8.0: moderate and 7.5: acidic). We measured the temperature sensitivity of righting response by generating thermal performance curves from individual data collected at seven test temperatures and determined critical thermal minima (CTmin) and maxima (CTmax). We found that T. cingulata has a broad basal thermal tolerance range (∼38°C) and after acclimation to the warm temperature regime, both the optimal temperature for maximum righting speed and CTmax increased. Contrary to predictions, acidification did not narrow this population's thermal tolerance but increased CTmax. These plastic responses are likely driven by the predictable exposure to temperature extremes measured in the field which originate from the local tidal cycle and the periodic acidification associated with ocean upwelling in the region. This acclimation ability suggests that T. cingulata has at least some capacity to buffer the thermal changes and increased acidification predicted to occur with climate change.

Local adaptation, phenotypic plasticity, and species coexistence

Understanding the mechanisms of species coexistence has always been a fundamental topic in ecology. Classical theory predicts that interspecific competition may select for traits that stabilize niche differences, although recent work shows that this is not strictly necessary. Here, we ask whether adaptive phenotypic plasticity could allow species coexistence (i.e., some stability at an equilibrium point) without ecological differentiation in habitat use. We used individual-based stochastic simulations defining a landscape composed of spatially uncorrelated or autocorrelated environmental patches, where two species with the same competitive strategies, not able to coexist without some form of phenotypic plasticity, expanded their ranges in the absence of a competition—colonization trade-off (a well-studied mechanism for species diversity). Each patch is characterized by a random environmental value that determines the optimal phenotype of its occupants. In such a scenario, only local adaptation and gene flow (migration) may interact to promote genetic variation and coexistence in the metapopulation. Results show that a competitively inferior species with adaptive phenotypic plasticity can coexist in a same patch with a competitively superior, non-plastic species, provided the migration rates and variances of the patches' environmental values are sufficiently large.

Indigenous and introduced Collembola differ in desiccation resistance but not its plasticity in response to temperature.

Biological invasions have significant ecological and economic impacts. Much attention is therefore focussed on predicting establishment and invasion success. Trait-based approaches are showing much promise, but are mostly restricted to investigations of plants. Although the application of these approaches to animals is growing rapidly, it is rare for arthropods and restricted mostly to investigations of thermal tolerance. Here we study the extent to which desiccation tolerance and its phenotypic plasticity differ between introduced (nine species) and indigenous (seven species) Collembola, specifically testing predictions of the 'ideal weed' and 'phenotypic plasticity' hypotheses of invasion biology. We do so on the F2 generation of adults in a full factorial design across two temperatures, to elicit desiccation responses, for the phenotypic plasticity trials. We also determine whether basal desiccation resistance responds to thermal laboratory natural selection. We first show experimentally that acclimation to different temperatures elicits changes to cuticular structure and function that are typically associated with water balance, justifying our experimental approach. Our main findings reveal that basal desiccation resistance differs, on average, between the indigenous and introduced species, but that this difference is weaker at higher temperatures, and is driven by particular taxa, as revealed by phylogenetic generalised least squares approaches. By contrast, the extent or form of phenotypic plasticity does not differ between the two groups, with a 'hotter is better' response being most common. Beneficial acclimation is characteristic of only a single species. Laboratory natural selection had little influence on desiccation resistance over 8-12 generations, suggesting that environmental filtering rather than adaptation to new environments may be an important factor influencing Collembola invasions.

Properties of phenotypic plasticity in discrete threshold traits.

Forms of phenotypic plasticity in key traits, and forms of selection on and genetic variation in such plasticity, fundamentally underpin phenotypic, population dynamic, and evolutionary responses to environmental variation and directional change. Accordingly, numerous theoretical and empirical studies have examined properties and consequences of plasticity, primarily considering traits that are continuously distributed on observed phenotypic scales with linear reaction norms. However, many environmentally sensitive traits are expressed as discrete alternative phenotypes and are appropriately characterized as quantitative genetic threshold traits. Here, we highlight that forms of phenotypic plasticity, genetic variation, and inheritance in plasticity, and outcomes of selection on plasticity, could differ substantially between threshold traits and continuously distributed traits (as are typically considered). We thereby highlight theoretical developments that are required to rationalize and predict phenotypic and microevolutionary dynamics involving plastic threshold traits, and outline how intrinsic properties of such traits could provide relatively straightforward explanations for apparently idiosyncratic observed patterns of phenotypic variation. We summarize how key quantitative genetic parameters underlying threshold traits can be estimated, and thereby set the scene for embedding dynamic discrete traits into theoretical and empirical understanding of the role of plasticity in driving phenotypic, population, and evolutionary responses to environmental variation and change.

Abstract 2897: Dynamic transdifferentiation programs in small cell lung carcinoma

Abstract Tumors must also have the ability to rapidly adjust to a mutable environment during the lifespan of the cell. This form of phenotypic plasticity is typified by biological programs regulated by a host of transcription factors in a well-coordinated “switch” that is initiated by both cell autonomous and microenvironmental cues. We sought to study these dynamics in small cell lung carcinoma (SCLC). First, we used k-means clustering to identify discrete transcriptional programs that are represented across SCLC using gene expression values from 53 and 71 SCLC cell lines and primary tumors, respectively. To investigate these programs further, we used our inventory of 67 patient-derived xenografts (PDX) to establish an ex vivo cell culture system that reconstitutes with high fidelity the gene expression values observed in the PDXs and the primary tumors from which these cells were derived. Our data demonstrates that gene expression groupings or clusters across samples mark distinct SCLC transcriptional states, representing intertumoral variation. However, in an intriguing demonstration of intratumoral heterogeneity, the very same groupings define morphologically and phenotypically distinct subpopulations within individual samples. Subpopulations were represented by suspending aggregates of very small cells (neuroendocrine or NEn), pre-suspension aggregates that are nested above mesenchymal-like cells and appear to give rise to the suspending aggregates (proneural or PN), pleomorphic cells growing as a monolayer composed of larger cells with visible cytoplasm and spindle-like membrane extensions (mesenchymal or MS), and semi-adherent cells that are physically positioned over the latter (neuroepithelial or NEp). WES demonstrated high genetic fidelity across all subpopulations, indicating that the significant morphologic and phenotypic differences were non-genetic. RNAseq and gene ontology was applied to differentially expressed genes to inform classifications. Using network analyses, we identified transcription factors (TFs) that function as central nodes in the gene expression networks of each population. Namely, we show that ASCL1 marks a rapidly proliferating neuroendocrine population (NEn), NEUROD1 marks a primitive neural population (PN) and YAP1 marks a slow-growing mesenchymal population (MS). We also developed a promoter-reporter fluorescent protein system that positions distinctly colored fluorescent proteins under the regulatory control of the promoters of the three TFs. We show that a Markov process models the transition probability of states. Taken together, our results show that human SCLC is comprised of distinct tumor subpopulations characterized by significant, non-genetic plasticity. We posit that understanding of the SCLC intratumoral ecosystem can create new opportunities to evolutionarily steer this tumor toward improved therapeutic results. Note: This abstract was not presented at the meeting. Citation Format: Priyanka Gopal, Kevin Rogacki, Craig D. Peacock, Mohamed E. Abazeed. Dynamic transdifferentiation programs in small cell lung carcinoma [abstract]. In: Proceedings of the American Association for Cancer Research Annual Meeting 2019; 2019 Mar 29-Apr 3; Atlanta, GA. Philadelphia (PA): AACR; Cancer Res 2019;79(13 Suppl):Abstract nr 2897.

Transgenerational Developmental Effects of Immune Priming in the Red Flour Beetle Tribolium castaneum.

Immune priming, the increased chance to survive a secondary encounter with a pathogen, has been described for many invertebrate species, which lack the classical adaptive immune system of vertebrates. Priming can be specific even for closely related bacterial strains, last up to the entire lifespan of an individual, and in some species, it can also be transferred to the offspring and is then called transgenerational immune priming (TGIP). In the red flour beetle Tribolium castaneum, a pest of stored grains, TGIP has even been shown to be transferred paternally after injection of adult beetles with heat-killed Bacillus thuringiensis. Here we studied whether TGIP in T. castaneum is also transferred to the second filial generation, whether it can also occur after oral and injection priming of larvae and whether it has effects on offspring development. We found that paternal priming with B. thuringiensis does not only protect the first but also the second offspring generation. Also, fitness costs of the immune priming became apparent, when the first filial generation produced fewer offspring. Furthermore, we used two different routes of exposure to prime larvae, either by injecting them with heat-killed bacteria or orally feeding them B. thuringiensis spore culture supernatant. Neither of the parental larval priming methods led to any direct benefits regarding offspring resistance. However, the injections slowed down development of the injected individuals, while oral priming with both a pathogenic and a non-pathogenic strain of B. thuringiensis delayed offspring development. The long-lasting transgenerational nature of immune priming and its impact on offspring development indicate that potentially underlying epigenetic modifications might be stable over several generations. Therefore, this form of phenotypic plasticity might impact pest control and should be considered when using products of bacterial origin against insects.

Sexual selection, phenotypic plasticity and female reproductive output.

In a rapidly changing environment, does sexual selection on males elevate a population's reproductive output? If so, does phenotypic plasticity enhance or diminish any such effect? We outline two routes by which sexual selection can influence the reproductive output of a population: a genetic correlation between male sexual competitiveness and female lifetime reproductive success; and direct effects of males on females' breeding success. We then discuss how phenotypic plasticity of sexually selected male traits and/or female responses (e.g. plasticity in mate choice), as the environment changes, might influence how sexual selection affects a population's reproductive output. Two key points emerge. First, condition-dependent expression of male sexual traits makes it likely that sexual selection increases female fitness if reproductively successful males disproportionately transfer genes that are under natural selection in both sexes, such as genes for foraging efficiency. Condition-dependence is a form of phenotypic plasticity if some of the variation in net resource acquisition and assimilation is attributable to the environment rather than solely genetic in origin. Second, the optimal allocation of resources into different condition-dependent traits depends on their marginal fitness gains. As male condition improves, this can therefore increase or, though rarely highlighted, actually decrease the expression of sexually selected traits. It is therefore crucial to understand how condition determines male allocation of resources to different sexually selected traits that vary in their immediate effects on female reproductive output (e.g. ornaments versus coercive behaviour). In addition, changes in the distribution of condition among males as the environment shifts could reduce phenotypic variance in certain male traits, thereby reducing the strength of sexual selection imposed by females. Studies of adaptive evolution under rapid environmental change should consider the possibility that phenotypic plasticity of sexually selected male traits, even if it elevates male fitness, could have a negative effect on female reproductive output, thereby increasing the risk of population extinction. This article is part of the theme issue 'The role of plasticity in phenotypic adaptation to rapid environmental change'.

Competition induces increased toxin production in toad larvae without allelopathic effects on heterospecific tadpoles

Abstract Inducible defences are a form of phenotypic plasticity by which organisms respond to and mitigate the threat posed by predators, parasites and competitors. While anti‐predatory defences are often in trade‐off with anti‐competitor responses, chemicals that deter predators may have negative effects on competitors as well. Allelopathy is well known in plants and plant‐like animals, but whether the toxins of mobile, behaviourally and morphologically complex animals are induced by and exert allelopathic effects on competitors is poorly known. Common toads Bufo bufo synthesize bufadienolides which make them unpalatable or toxic to many predators. However, bufadienolide content of toad tadpoles correlates positively with the density of competitors in natural populations, suggesting that they may upregulate their toxin production to inhibit their competitors, such as heterospecific tadpoles that may be vulnerable to toad toxins. We conducted a microcosm experiment with tadpoles of common toads and agile frogs Rana dalmatina, in which we manipulated the density of conspecific and heterospecific competitors. We measured the bufadienolide content of toad tadpoles to test for competitor‐induced changes in toxin production, and we assessed the growth and development of agile frog tadpoles to test for allelopathy. We found that toad tadpoles contained higher amounts of bufadienolides at higher densities; however, heterospecific competitors did not have a stronger effect than conspecifics. Furthermore, the presence or density of toad tadpoles had no effect on the body mass and development rate of agile frog tadpoles. Our results demonstrate competitor‐induced plasticity in toxin production, but we found no support for an allelopathic function of bufadienolides. Instead, we suggest that inducible changes in bufadienolide production may serve to mitigate risks posed by competitors, including aggression, cannibalism or disease. Therefore, bufadienolides are intriguing candidates for multi‐purpose defences that may provide protection not only against predators but also against competitors. A plain language summary is available for this article.

Nutritional Regulation of Phenotypic Plasticity in a Solitary Bee (Hymenoptera: Megachilidae).

Phenotypic plasticity involves adaptive responses to predictable environmental fluctuations and may promote evolutionary change. We studied the regulation of phenotypic plasticity in an important agricultural pollinator, the solitary alfalfa leafcutting bee (Megachile rotundata F.). Specifically, we investigated how larval nutrition affects M. rotundata diapause plasticity and how diapause plasticity affects adult female reproductive behavior. Field surveys and laboratory manipulations of aspects of larval diet demonstrated nutritional regulation of M. rotundata diapause plasticity. Manipulation of larval diet quality through the addition of royal jelly, the caste-determining substance of the honey bee Apis mellifera L., increased the probability of diapause in M. rotundata. We also found that larval nutrition and diapause status affected M. rotundata adult female reproductive behavior. Nutritional effects on larval diapause that also impact adult fitness have intriguing implications for the evolution of developmental plasticity in bees. In particular, as the solitary lifestyle of M. rotundata is considered to be the ancestral condition in bees, nutritionally regulated plasticity may have been an ancestral condition in all bees that facilitated the evolution of other forms of phenotypic plasticity, such as the castes of social bees.

Behaviorally Induced Camouflage: A New Mechanism of Avian Egg Protection.

When animals potentially occupy diverse microhabitats, how can camouflage be achieved? Here we combine descriptive and experimental methods to uncover a novel form of phenotypic plasticity in the camouflage of bird eggs that may be present in other avian taxa. Soil from the bare substrate adheres to the blue-footed booby's (Sula nebouxii's) pale eggs, which parents manipulate both under and on top of their webs. Analysis of digital images confirmed that dirtiness increases progressively during the first 16 days of the incubation period, making eggs more similar to the nest substrate. Observations of 3,668 single-egg clutches showed that the probability of egg loss declines progressively over the same time frame and then remains low for the rest of the 41-day incubation period. An experiment showed that when chicken eggs are soiled and exposed in artificial booby nests, they are less likely to be taken by Heermann's gulls (Larus heermanni) than clean eggs.

'Do you remember the first time?' Host plant preference in a moth is modulated by experiences during larval feeding and adult mating.

In insects, like in other animals, experience-based modulation of preference, a form of phenotypic plasticity, is common in heterogeneous environments. However, the role of multiple fitness-relevant experiences on insect preference remains largely unexplored. For the multivoltine polyphagous moth Spodoptera littoralis we investigated effects of larval and adult experiences on subsequent reproductive behaviours. We demonstrate, for the first time in male and female insects, that mating experience on a plant modulates plant preference in subsequent reproductive behaviours, whereas exposure to the plant alone or plant together with sex pheromone does not affect this preference. When including larval feeding experiences, we found that both larval rearing and adult mating experiences modulate host plant preference. These findings represent the first evidence that host plant preferences in polyphagous insects are determined by a combination of innate preferences modulated by sensory feedback triggered by multiple rewarding experiences throughout their lifetime.

Cross‐generational costs of compensatory growth in nine‐spined sticklebacks

Compensatory growth (CG) is a form of phenotypic plasticity allowing individuals’ growth trajectories to rebound after a period of resource limitation, but little is known about the reproductive and cross‐generational costs of CG. We studied the potential costs of CG by exposing female nine‐spined sticklebacks Pungitius pungitius to 1) high (favourable), 2) low (stressful), and 3) recovery (initially stressful, subsequently favourable) feeding treatments, and quantified the effects of these treatments on female reproductive traits (clutch, egg and yolk size), and on the size of their offspring. The low feeding treatment reduced female size relative to the high and recovery feeding treatments, which produced equally large females. Hence, females from the recovery treatment demonstrated CG and full growth compensation. Feeding treatments had significant effects on clutch, yolk, egg and larval size, also when the effect of female size was controlled for. However, these effects came about mostly because females from the low feeding treatment produced small clutches with large eggs (containing little yolk) and larvae, whereas females from the recovery feeding treatment produced as large clutches, eggs (with similar amounts of yolk) and larvae as females from the high feeding treatment. Yet, structural equation modelling revealed that while a direct effect of female size on offspring size was positive in the low and high feeding treatments, it was negative in the recovery feeding treatment, independently of egg and clutch size. This indicates a cross‐generational tradeoff between female and offspring sizes in the recovery feeding treatment. Also the tradeoff between clutch and larval size was more pronounced in recovery than in low or high feeding treatments. Hence, apart from demonstrating that environmental influences experienced by females during their development have the potential to influence their size, fecundity and reproductive traits, the results also provide evidence for complex cross‐generational costs of recovery growth.

Alpine activity patterns of Mitopus morio (Fabricius, 1779) are induced by variations in temperature and humidity at different scales in central Norway

Our research addresses questions about how micro-climate affects activity abundance of a common and widespread harvestman in an alpine ecosystem. Activity patterns of the Harvestman Mitopus morio (Fabricius, 1779) were studied along different alpine gradients in the central Norwegian Scandes. Within a nested design, we surveyed 18 alpine habitats with pitfall traps and micro-climatological equipment along oceanic-continental, two elevational, and (fine-scaled) microtopographic gradients. Sites in the oceanic region of the Scandes showed generally higher abundance of M. morio than sites in the continental region. Furthermore, along the elevational gradient, middle-alpine sites showed higher abundances than low-alpine sites. These general patterns are best explained by higher humidity in the oceanic region and in the middle-alpine belt. Focusing at a finer scale, i.e. one elevational level within each region, revealed partly opposing activity patterns within relatively short distances. While in the western middle-alpine belt these patterns were best explained by humidity-related measures but now with higher activity abundance during drier conditions, in the drier eastern middle-alpine belt heat sums rather than humidity were found to be the best explanatory variables for the observed patterns. Hence, our results imply a pronounced different reaction of the two populations towards climatic variables that partly even contradict the previously described general pattern. Regardless whether these differences in activity abundance in M. morio are a form of phenotypic plasticity or adaptation, our findings stress the importance of detailed autecological knowledge combined with fine-scaled climatic measurements when aiming at predictions about possible future ecosystem structures and spatio-temporal phenomena. M. morio proves to be an ideal biogeographic model organism for understanding spatio-temporal responses of alpine ecosystems under modified climatic conditions.

Orchids do not pay costs at emergence for prolonged dormancy

In plants, prolonged dormancy is often considered a response to resource depletion or environmental stress that comes at a fitness cost. However, apparent costs of dormancy could reflect the state in which plants entered dormancy, rather than effects of dormancy per se. We tested this hypothesis for a terrestrial orchid, Epipactis atrorubens, by analyzing differences in vital rates of dormant and emergent plants using generalized linear mixed models, applied to eight years of demographic data. Dormant E. atrorubens plants did not form one homogeneous stage class. Instead, the vital rates of dormant plants mirrored performance of plants in their life stage before dormancy. Plants emerging from dormancy were slightly (albeit only marginally statistically significantly) larger than plants transitioning from the matching aboveground stage class, especially for smaller and younger stage classes. Because small plants were most likely to go dormant, plants emerging from dormancy were also smaller than average, if one were to compare all previously dormant plants to all previously emergent plants. Therefore, misclassifying all dormant plants into a single stage class changes whether we view dormancy as intrinsically costly, in terms of future performance upon emergence. We suggest that prolonged dormancy may be a form of phenotypic plasticity in which plants distribute their performance and reproductive effort through time, rather than a simple stress response.

Pristionchus pacificus daf-16is essential for dauer formation but dispensable for mouth form dimorphism

The nematode Pristionchus pacificus shows two forms of phenotypic plasticity: dauer formation and dimorphism of mouth form morphologies. It can therefore serve as a model for studying the evolutionary mechanisms that underlie phenotypic plasticity. Formation of dauer larvae is observed in many other species and constitutes one of the most crucial survival strategies in nematodes, whereas the mouth form dimorphism is an evolutionary novelty observed only in P. pacificus and related nematodes. We have previously shown that the same environmental cues and steroid signaling control both dauer formation and mouth form dimorphism. Here, we examine by mutational analysis and whole-genome sequencing the function of P. pacificus (Ppa) daf-16, which encodes a forkhead transcription factor; in C. elegans, daf-16 is the target of insulin signaling and plays important roles in dauer formation. We found that mutations in Ppa-daf-16 cause strong dauer formation-defective phenotypes, suggesting that Ppa-daf-16 represents one of the evolutionarily conserved regulators of dauer formation. Upon strong dauer induction with lophenol, Ppa-daf-16 individuals formed arrested larvae that partially resemble wild-type dauer larvae, indicating that Ppa-daf-16 is also required for dauer morphogenesis. By contrast, regulation of mouth form dimorphism was unaffected by Ppa-daf-16 mutations and mutant animals responded normally to environmental cues. Our results suggest that mechanisms for dauer formation and mouth form regulation overlap partially, but not completely, and one of two key transcriptional regulators of the dauer regulatory network was either independently co-opted for, or subsequently lost by, the mouth form regulatory network.

A ROLE FOR LEARNING IN POPULATION DIVERGENCE OF MATE PREFERENCES

Learning and other forms of phenotypic plasticity have been suggested to enhance population divergence. Mate preferences can develop by learning, and species recognition might not be entirely genetic. We present data on female mate preferences of the banded demoiselle (Calopteryx splendens) that suggest a role for learning in population divergence and species recognition. Populations of this species are either allopatric or sympatric with a phenotypically similar congener (C. virgo). These two species differ mainly in the amount of wing melanization in males, and wing patches thus mediate sexual isolation. In sympatry, sexually experienced females discriminate against large melanin wing patches in heterospecific males. In contrast, in allopatric populations within the same geographic region, females show positive ("open-ended") preferences for such large wing patches. Virgin C. splendens females do not discriminate against heterospecific males. Moreover, physical exposure experiments of such virgin females to con- or hetero-specific males significantly influences their subsequent mate preferences. Species recognition is thus not entirely genetic and it is partly influenced by interactions with mates. Learning causes pronounced population divergence in mate preferences between these weakly genetically differentiated populations, and results in a highly divergent pattern of species recognition at a small geographic scale.

Environmental Dependence of Thermal Reaction Norms: Host Plant Quality Can Reverse the Temperature‐Size Rule

The temperature-size rule, a form of phenotypic plasticity in which decreased temperature increases final size, is one of the most widespread patterns in biology, particularly for ectotherms. Identifying the environmental conditions in which this pattern is reversed is key to understanding the generality of the rule. We use wild and domesticated populations of the tobacco hornworm Manduca sexta and the natural host plants of this species to explore the consequences of resource quality for the temperature-size rule. Manduca sexta reared on a high-quality host, tobacco (Nicotiana tabacum), followed the temperature-size rule, with larger final sizes at lower temperatures. In contrast, M. sexta reared on a low-quality host, devil's claw (Proboscidea louisianica), showed the reverse response. Wild and domesticated M. sexta exhibited qualitatively similar responses. Survival, growth and development rates, fecundity, and final size decreased with decreasing temperature in M. sexta reared on devil's claw. We propose that the reversal of the temperature-size rule results from the stressful combination of low temperatures and low dietary quality. Such reversals may impact seasonal and geographic patterns of host use in Manduca and other systems. Our results suggest that the temperature-size rule occurs for a restricted range of nonstressful environmental conditions, limiting the robustness of this widespread pattern of phenotypic plasticity.

Ecological dynamics and the basis of sympatric phenotypic diversification

AbstractTheoretical and empirical studies are showing evidence in support of evolutionary branching and sympatric speciation due to frequency‐dependent competition. However, phenotypic diversification due to underlying genetic diversification is only one possible evolutionary response to disruptive selection. Another potentially general response is phenotypic diversification in the form of phenotypic plasticity. It has been suggested that genetic variation is favored in stable environments, whereas phenotypic plasticity is favored in unstable and fluctuating environments. We investigate the “competition” between the processes of evolutionary branching and the evolution of phenotypic plasticity in a predator‐prey model that allows both processes to occur. In this model, environmental fluctuations can be caused by complicated population dynamics. We found that the evolution of phenotypic plasticity was generally more likely than evolutionary branching when the ecological dynamics exhibited pronounced predator‐prey cycles, whereas the opposite was true when the ecological dynamics was more stable. At intermediate levels of density cycling, trimorphisms with two specialist branches and a phenotypically plastic generalist branch sometimes occurred. Our theoretical results suggest that ecological dynamics and evolutionary dynamics can often be tightly linked and that an explicit consideration of population dynamics may be essential to explain the evolutionary dynamics of diversification in natural populations.