Abstract
Biological systems are inherently hierarchical. Consequently, any field which aims to understand an aspect of biology holistically requires investigations at each level of the hierarchy of life, and venom research is no exception. This article aims to illustrate the structure of the field in light of a ‘levels of life’ perspective. In doing so, I highlight how traditional fields and approaches fit into this structure as focussing on describing levels or investigating links between levels, and emphasise where implicit assumptions are made due to lack of direct information. Taking a ‘levels of life’ perspective to venom research enables us to understand the complementarity of different research programmes and identify avenues for future research. Moreover, it provides a broader view that, in itself, shows how new questions can be addressed. For instance, understanding how adaptations develop and function from molecular to organismal scales, and what the consequences are of those adaptations at scales from molecular to macroevolutionary, is a general question relevant to a great deal of biology. As a trait which is molecular in nature and has clearer and more direct links between genotype and phenotype than many other traits, venom provides a relatively simple system to address such questions. Furthermore, because venom is also diverse at each level of life, the complexity within the hierarchical structure provides variation that enables powerful analytical approaches to answering questions. As a result, venom provides an excellent model system for understanding big questions in evolutionary biology.
Highlights
Venom at the interface of molecular biology and ecologyVenoms have long captured the interests of evolutionary biologists; in the early 1900s Alcock and Rogers (1902) were already investigating how snake venoms could have originated under Darwinian natural se lection
Each of the types of ecological interactions that venom is involved in are closely linked to evolutionary fitness, generating a tight linkage between venom phenotypes and fitness. They are universally characterised as antagonistic coevolutionary relationships (Arbuckle, 2017), which are known or predicted to have widespread and profound impacts on evolutionary dynamics (Queller and Strassmann, 2018)
Fundamentally a molecular trait, venom is intricately tied to the ecology and evolution of the organisms in question, providing a powerful example for studies of evolutionary processes and adaptation across biological hierarchies
Summary
Venoms have long captured the interests of evolutionary biologists; in the early 1900s Alcock and Rogers (1902) were already investigating how snake venoms could have originated under Darwinian natural se lection. Fitness consequences are often highly variable for a given phenotype based on ecological context (sometimes even reversing in sign), a single phenotype is often the outcome of a large number of underlying genes which interact in complex ways, and historical contingency can channel the underlying genetic evolution down restricted routes leading to sto chastic evolution of different fitness peaks (Elena and Lenski, 2003) While it is unlikely any natural system can be simple enough to elimi nate those limitations, venom does have some advantages. Each of the types of ecological interactions that venom is involved in are closely linked to evolutionary fitness (e.g. energy intake, survival, and competition for mating opportunities), generating a tight linkage between venom phenotypes and fitness They are universally characterised as antagonistic coevolutionary relationships (Arbuckle, 2017), which are known or predicted to have widespread and profound impacts on evolutionary dynamics (Queller and Strassmann, 2018). This realisation has recently seen a substantial presence in the literature as several recent reviews emphasising the need for and promise of behavioural, ecological, and Toxicon: X 6 (2020) 100034 evolutionary questions in venom research (Calvete, 2013; Valcu and Kempenaers, 2015; Sunagar et al, 2016; Arbuckle 2017, 2018; Jackson et al, 2019; Schendel et al, 2019)
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