Correlated evolution of phenotypic plasticity in metamorphic timing

Phenotypic plasticity has long been a focus of research, but the mechanisms of its evolution remain controversial. Many amphibian species exhibit a similar plastic response in metamorphic timing in response to multiple environmental factors; therefore, more than one environmental factor has likely influenced the evolution of plasticity. However, it is unclear whether the plastic responses to different factors have evolved independently. In this study, we examined the relationship between the plastic responses to two experimental factors (water level and food type) in larvae of the salamander Hynobius retardatus, using a cause-specific Cox proportional hazards model on the time to completion of metamorphosis. Larvae from ephemeral ponds metamorphosed earlier than those from permanent ponds when kept at a low water level or fed conspecific larvae instead of larval Chironomidae. This acceleration of metamorphosis depended only on the permanency of the larvae’s pond of origin, but not on the conspecific larval density (an indicator of the frequency of cannibalism) in the ponds. The two plastic responses were significantly correlated, indicating that they may evolve correlatively. Once plasticity evolved as an adaptation to habitat desiccation, it might have relatively easily become a response to other ecological factors, such as food type via the pre-existing developmental pathway

Correlated evolution of phenotypic plasticity in metamorphic timing

Phenotypic plasticity has long been a focus of research, but the mechanisms of its evolution remain controversial. Many amphibian species exhibit a similar plastic response in metamorphic timing in response to multiple environmental factors; therefore, more than one environmental factor has likely influenced the evolution of plasticity. However, it is unclear whether the plastic responses to different factors have evolved independently. In this study, we examined the relationship between the plastic responses to two experimental factors (water level and food type) in larvae of the salamander Hynobius retardatus, using a cause-specific Cox proportional hazards model on the time to completion of metamorphosis. Larvae from ephemeral ponds metamorphosed earlier than those from permanent ponds when kept at a low water level or fed conspecific larvae instead of larval Chironomidae. This acceleration of metamorphosis depended only on the permanency of the larvae’s pond of origin, but not on the conspecific larval density (an indicator of the frequency of cannibalism) in the ponds. The two plastic responses were significantly correlated, indicating that they may evolve correlatively. Once plasticity evolved as an adaptation to habitat desiccation, it might have relatively easily become a response to other ecological factors, such as food type via the pre-existing developmental pathway

A trade-off between prey- and predator-induced polyphenisms in larvae of the salamander Hynobius retardatus

Organisms in natural habitats participate in complex ecological interactions that include competition, predation, and foraging. Under natural aquatic environmental conditions, amphibian larvae can simultaneously receive multiple signals from conspecifics, predators, and prey, implying that predator-induced morphological defenses can occur in prey and that prey-induced offensive morphological traits may develop in predators. Although multiple adaptive plasticity, such as inducible defenses and inducible offensive traits, can be expected to have not only ecological but also evolutionary implications, few empirical studies report on species having such plasticity. The broad-headed larval morph of Hynobius retardatus, which is induced by crowding with heterospecific anuran (Rana pirica) larvae, is a representative example of prey-induced polyphenism. The morph is one of two distinct morphs that have been identified in this species; the other is the typical morph. In this paper, we report that typical larval morphs of Hynobius can respond rapidly to a predatory environment and show conspicuous predator-induced plasticity of larval tail depth, but that broad-headed morphs cannot respond similarly to a predation threat. Our findings support the hypothesis that induction or maintenance of adaptive plasticity (e.g., predator-induced polyphenism) trades off against other adaptive plastic responses (e.g., prey-induced polyphenism). For a species to retain both an ability to forage for larger prey and an ability to more effectively resist predation makes sense in light of the range of environments that many salamander larvae experience in nature. Our results suggest that the salamander larvae clearly discriminate between cues from prey and those from predators and accurately respond to each cue; that is, they adjust their phenotype to the current environment

Stat Methods Med Res

Developing a personalized risk prediction model of death is fundamental for improving patient care and touches on the realm of personalized medicine. The increasing availability of genomic information and large-scale meta-analytic data sets for clinicians has motivated the extension of traditional survival prediction based on the Cox proportional hazards model. The aim of our paper is to develop a personalized risk prediction formula for death according to genetic factors and dynamic tumour progression status based on meta-analytic data. To this end, we extend the existing joint frailty-copula model to a model allowing for high-dimensional genetic factors. In addition, we propose a dynamic prediction formula to predict death given tumour progression events possibly occurring after treatment or surgery. For clinical use, we implement the computation software of the prediction formula in the joint.Cox R package. We also develop a tool to validate the performance of the prediction formula by assessing the prediction error. We illustrate the method with the meta-analysis of individual patient data on ovarian cancer patients

Top-down effects on antagonistic inducible defense and offense

Antagonistic phenotypic plasticity may strongly influence trait evolution in tightly interacting predator-prey pairs as well as the role that trait plasticity plays in community dynamics. Most work on trait plasticity has focused on single predator-prey pairs, but prey must often contend with multiple predators in natural environments. Hence, a better understanding of the evolutionary and ecological significance of phenotypic plasticity requires experiments that examine how multiple predators shape prey trait plasticity. Here, using a simple food chain consisting of a top predator (dragonfly larvae, Aeshna nigroflava), an intermediate predator (salamander larvae, Hynobius retardatus), and frog (Rana pirica) tadpoles as prey, we show that the presence of dragonfly risk cues substantially modifies the intensity of antagonistic morphological plasticity in both amphibians. In the absence of dragonflies, tadpoles produced bulgier bodies in response to salamanders, and salamanders responded to this defense by enlarging their gape size. However, in the presence of dragonfly risk cues, the expression of both antagonistic traits was significantly reduced because tadpoles and salamanders produced phenotypes that are more effective against dragonfly predators. Thus, the reduced antagonism likely emerged, in part, because the benefits of antagonistic trait expression were outweighed by the potential cost of increased vulnerability to dragonfly predation. In addition, our results suggest that when all three species were present, salamander activity levels, which influence the amount of signals required to induce antagonistic traits, were more strongly affected by dragonfly risk cues than were tadpole activity levels. This species-specific difference in activity levels was likely responsible for the reduced tadpole mortality caused by salamanders in the presence vs. absence of dragonfly risk cues. Hence, dragonflies had a positive trait-mediated indirect effect on tadpoles by modifying both the morphological and behavioral traits of salamanders

Natural variation in morphology of larval amphibians: Phenotypic plasticity in nature?

Phenotypic plasticity has been studied intensively in experimental settings but infrequently in nature, and therefore the relevance of experimental findings is poorly known. This is especially true for morphological plasticity in amphibian larvae induced by predators and competitors. This paper describes a seven-year survey of head and tail shape in eight species of anuran and newt larvae in northern Switzerland, involving 6824 individual larvae and 59 ponds. I tested relationships between geometric measures of size and shape and five habitat gradients: pond permanence, cover by forest canopy and aquatic vegetation, and the densities of predators and competitors. Responses to competitors and predators were often similar to those reported in experiments. High competitor density was associated with small size and a large head in newt larvae, a long or deep head/body in anuran larvae, and a short or shallow tail in newts and some tadpoles. High predator density was correlated with a deep tail fin and tail muscle in many species. In anurans, the change in shape between low- and high predator ponds in nature closely paralleled the plastic response to nonlethal predators in mesocosm experiments. The survey revealed many previously undescribed relationships between morphology and the other habitat features. Several species had relatively large tails in ponds that were shaded or thickly vegetated. Associations between year-to-year changes in shape and habitat within ponds implicated phenotypic plasticity rather than genetic population divergence, at least in anurans. These results inspire confidence in the relevance of experiments and highlight many new patterns that will merit further study