Testing the ecological consequences of evolutionary change using elements

Understanding the ecological consequences of evolutionary change is a central challenge in contemporary biology. We propose a framework based on the ~25 elements represented in biology, which can serve as a conduit for a general exploration of poorly understood evolution-to-ecology links. In this framework, known as ecological stoichiometry, the quantity of elements in the inorganic realm is a fundamental environment, while the flow of elements from the abiotic to the biotic realm is due to the action of genomes, with the unused elements excreted back into the inorganic realm affecting ecological processes at higher levels of organization. Ecological stoichiometry purposefully assumes distinct elemental composition of species, enabling powerful predictions about the ecological functions of species. However, this assumption results in a simplified view of the evolutionary mechanisms underlying diversification in the elemental composition of species. Recent research indicates substantial intraspecific variation in elemental composition and associated ecological functions such as nutrient excretion. We posit that attention to intraspecific variation in elemental composition will facilitate a synthesis of stoichiometric information in light of population genetics theory for a rigorous exploration of the ecological consequences of evolutionary change.Peer reviewedZoolog

SEED: A framework for integrating ecological stoichiometry and eco‐evolutionary dynamics

Characterising the extent and sources of intraspecific variation and their ecological consequences is a central challenge in the study of eco‐evolutionary dynamics. Ecological stoichiometry, which uses elemental variation of organisms and their environment to understand ecosystem patterns and processes, can be a powerful framework for characterising eco‐evolutionary dynamics. However, the current emphasis on the relative content of elements in the body (i.e. organismal stoichiometry) has constrained its application. Intraspecific variation in the rates at which elements are acquired, assimilated, allocated or lost is often greater than the variation in organismal stoichiometry. There is much to gain from studying these traits together as components of an ‘elemental phenotype’. Furthermore, each of these traits can have distinct ecological effects that are underappreciated in the current literature. We propose a conceptual framework that explores how microevolutionary change in the elemental phenotype occurs, how its components interact with each other and with other traits, and how its changes can affect a wide range of ecological processes. We demonstrate how the framework can be used to generate novel hypotheses and outline pathways for future research that enhance our ability to explain, analyse and predict eco‐evolutionary dynamics

Ecological stoichiometry beyond Redfield: An ionomic perspective on elemental homeostasis

Elemental homeostasis has been largely characterized using three important elements that were part of the Redfield ratio (i.e., carbon: nitrogen: phosphorus). These efforts have revealed substantial diversity in homeostasis among taxonomic groups and even within populations. Understanding the evolutionary basis, and ecological consequences of such diversity is a central challenge. Here, we propose that a more complete understanding of homeostasis necessitates the consideration of other elements beyond C, N, and P. Specifically, we posit that physiological complexity underlying maintenance of elemental homeostasis along a single elemental axis impacts processing of other elements, thus altering elemental homeostasis along other axes. Indeed, transcriptomic studies in a wide variety of organisms have found that individuals differentially express significant proportions of the genome in response to variability in supply stoichiometry in order to maintain varying levels of homeostasis. We review the literature from the emergent field of ionomics that has established the consequences of such physiological trade-offs on the content of the entire suite of elements in an individual. Further, we present experimental data on bacteria exhibiting divergent phosphorus homeostasis phenotypes demonstrating the fundamental interconnectedness among elemental quotas. These observations suggest that physiological adjustments can lead to unexpected patterns in biomass stoichiometry, such as correlated changes among suites of non-limiting microelements in response to limitation by macroelements. Including the entire suite of elements that comprise biomass will foster improved quantitative understanding of the links between chemical cycles and the physiology of organisms.Peer reviewedIntegrative Biolog

Understanding variation in salamander ionomes: A nutrient balance approach

Ecological stoichiometry uses information on a few key biological elements (C, N, and P) to explain complex ecological patterns. Although factors driving variation in these elements are well-established, expanding stoichiometric principles to explore dynamics of the many other essential elements comprising biological tissues (i.e., the ionome) is needed to determine their metabolic relationships and better understand biological control of elemental flows through ecosystems. 2. In this paper, we report observations of ionomic variation in two species of salamander (Ambystoma opacum and A. talpoideum) across ontogenic stages using specimens from biological collections of two wetlands sampled over a 30-year period. This unique data set allowed us to explore the extent of ionomic variation between species, among ontogenic stages, between sites, and through time. 3. We found species- and to a lesser extent site-specific differences in C, N, and P along with 13 other elements forming salamander ionomes but saw no evidence of temporal changes. Salamander ionomic composition was most strongly related to ontogeny with relatively higher concentrations of many elements in adult males (i.e., Ca, P, S, Mg, Zn, and Cu) compared to metamorphic juveniles, which had greater amounts of C, Fe, and Mn. 4. In addition to patterns of individual elements, covariance among elements was used to construct multi-elemental nutrient balances, which revealed differences in salamander elemental composition between species and sites and changes in elemental proportions across ontogenic development. These multi-elemental balances distinguished among species-site-ontogenic stage groups better than using only C, N, and P. 5. Overall, this study highlights the responsiveness of consumer ionomes to life-history and environmental variation while reflecting underlying relationships among elements tied to biological function. As such, ionomic studies can provide important insights into factors shaping consumer elemental composition and for predicting how these changes might affect higher-order ecological processes

Does differential iron supply to algae affect Daphnia life history? : An ionome-wide study

The availability of iron (Fe) varies considerably among diet items, as well as ecosystems. Availability of Fe has also changed due to anthropogenic environmental changes in oceanic as well as inland ecosystems. We know little about its role in the nutrition of ecologically important consumers, particularly in inland ecosystems. Physiological studies in several taxa indicate marked effects of dietary Fe on oogenesis. We predicted that differential Fe supply to algae will impact algal Fe concentration with consequences on the life history of the freshwater grazer, Daphnia magna. We found that algal Fe concentration increased with Fe supply, but did not affect algal growth, indicating that the majority of experimental Fe additions were likely adsorbed to, or stored in algal cells. Regardless, data indicate that algal Fe impacted the reproductive traits (age and size at maturity) but not juvenile growth rate of Daphnia. A subsequent experiment revealed that Fe concentration in eggs was significantly higher than the rest of Daphnia. These results indicate that the concentration of Fe in or on algal cells may vary considerably among ecosystems overlying distinct geological formations differing in Fe, possibly with important implications for zooplankton life histories. Understanding the mechanisms underlying this response is unlikely to be accomplished by a strict focus on Fe because we found correlated shifts in the algal ionome, with concomitant ionome-wide adjustments in Daphnia. Information on ionome-wide responses may be useful in better understanding the responses of biota to changes in the supply of any one element.peerReviewe

Crop Depredation by Wildlife Along the Eastern Boundary of the Kalakad mundanthurai Tiger Reserve, Southern India

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Adaptation to a limiting element involves mitigation of multiple elemental imbalances

About 20 elements underlie biology and thus constrain biomass production. Recent systems-level observations indicate that altered supply of one element impacts the processing of most elements encompassing an organism (i.e. ionome). Little is known about the evolutionary tendencies of ionomes as populations adapt to distinct biogeochemical environments. We evolved the bacterium Serratia marcescens under five conditions (i.e. low carbon, nitrogen, phosphorus, iron or manganese) that limited the yield of the ancestor compared with replete medium, and measured the concentrations and use efficiency of these five, and five other elements. Both physiological responses of the ancestor, as well as evolutionary responses of descendants to experimental environments involved changes in the content and use efficiencies of the limiting element, and several others. Differences in coefficients of variation in elemental contents based on biological functions were evident, with those involved in biochemical building (C, N, P, S) varying least, followed by biochemical balance (Ca, K, Mg, Na), and biochemical catalysis (Fe, Mn). Finally, descendants evolved to mitigate elemental imbalances evident in the ancestor in response to limiting conditions. Understanding the tendencies of such ionomic responses will be useful to better forecast biological responses to geochemical changes.peerReviewe

Endopolyploidy as a potential driver of animal ecology and evolution

Endopolyploidy – the existence of higher-ploidy cells within organisms that are otherwise of a lower ploidy level (generally diploid) – was discovered decades ago, but remains poorly studied relative to other genomic phenomena, especially in animals. Our synthetic review suggests that endopolyploidy is more common in animals than often recognized and probably influences a number of fitness-related and ecologically important traits. In particular, we argue that endopolyploidy is likely to play a central role in key traits such as gene expression, body and cell size, and growth rate, and in a variety of cell types, including those responsible for tissue regeneration, nutrient storage, and inducible anti-predator defences. We also summarize evidence for intraspecific genetic variation in endopolyploid levels and make the case that the existence of this variation suggests that endopolyploid levels are likely to be heritable and thus a potential target for natural selection. We then discuss why, in light of evident benefits of endopolyploidy, animals remain primarily diploid. We conclude by highlighting key areas for future research such as comprehensive evaluation of the heritability of endopolyploidy and the adaptive scope of endopolyploid-related traits, the extent to which endopolyploid induction incurs costs, and characterization of the relationships between environmental variability and endopolyploid levels. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Versions. © 2015 Wile