Abstract Animals use a diverse array of motion to feed, escape predators, and reproduce. Linking morphology, performance, and fitness is a foundational paradigm in organismal biology and evolution. Yet, the influence of mechanical relationships on evolutionary diversity remains unresolved. Here, I focus on the many-to-one mapping of form to function, a widespread, emergent property of many mechanical systems in nature, and discuss how mechanical redundancy influences the tempo and mode of phenotypic evolution. By supplying many possible morphological pathways for functional adaptation, many-to-one mapping can release morphology from selection on performance. Consequently, many-to-one mapping decouples morphological and functional diversification. In fish, for example, parallel morphological evolution is weaker for traits that contribute to mechanically redundant motions, like suction feeding performance, than for systems with one-to-one form–function relationships, like lower jaw lever ratios. As mechanical complexity increases, historical factors play a stronger role in shaping evolutionary trajectories. Many-to-one mapping, however, does not always result in equal freedom of morphological evolution. The kinematics of complex systems can often be reduced to variation in a few traits of high mechanical effect. In various different four-bar linkage systems, for example, mechanical output (kinematic transmission) is highly sensitive to size variation in one or two links, and insensitive to variation in the others. In four-bar linkage systems, faster rates of evolution are biased to traits of high mechanical effect. Mechanical sensitivity also results in stronger parallel evolution—evolutionary transitions in mechanical output are coupled with transition in linkages of high mechanical effect. In other words, the evolutionary dynamics of complex systems can actually approximate that of simpler, one-to-one systems when mechanical sensitivity is strong. When examined in a macroevolutionary framework, the same mechanical system may experience distinct selective pressures in different groups of organisms. For example, performance tradeoffs are stronger for organisms that use the same mechanical structure for more functions. In general, stronger performance tradeoffs result in less phenotypic diversity in the system and, sometimes, a slower rate of evolution. These macroevolutionary trends can contribute to unevenness in functional and lineage diversity across the tree of life. Finally, I discuss how the evolution of mechanical systems informs our understanding of the relative roles of determinism and contingency in evolution.
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Different Traits at Different Rates: The Effects of Dynamic Strain Rate on Structural Traits in Biology
Phenotypic diversity is influenced by physical laws that govern how an organism's morphology relates to functional performance. To study comparative organismal biology, we need to quantify this diversity using biological traits (definable aspects of the morphology, behavior, and/or life history of an organism). Traits are often assumed to be immutable properties that need to be measured only a single time in each adult. However, organisms often experience changes in their biotic and abiotic environments that can alter trait function. In particular, structural traits represent the physical capabilities of an organism and may be heavily influenced by the rate at which they are exposed to physical demands (“loads”). For instance, materials tend to become more brittle when loaded at faster rates which could negatively affect structures trying to resist those loads (e.g., brittle materials are more likely to fracture). In the following perspective piece, we address the dynamic properties of structural traits and present case studies that demonstrate how dynamic strain rates affect the function of these traits in diverse groups of organisms. First, we review how strain rate affects deformation and fracture in biomaterials and demonstrate how these effects alter puncture mechanics in systems such as snake strikes. Second, we discuss how different rates of bone loading affect the locomotor biomechanics of vertebrates and their ecology. Through these examinations of diverse taxa and ecological functions, we aim to highlight how rate-dependent properties of structural traits can generate dynamic form–function relationships in response to changing environmental conditions. Findings from these studies serve as a foundation to develop more nuanced ecomechanical models that can predict how complex traits emerge and, thereby, advance progress on outlining the Rules of Life.
more » « less- NSF-PAR ID:
- 10469028
- Publisher / Repository:
- Oxford University Press
- Date Published:
- Journal Name:
- Integrative And Comparative Biology
- Volume:
- 62
- Issue:
- 3
- ISSN:
- 1540-7063
- Format(s):
- Medium: X Size: p. 683-699
- Size(s):
- ["p. 683-699"]
- Sponsoring Org:
- National Science Foundation
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INTRODUCTION A major challenge in genomics is discerning which bases among billions alter organismal phenotypes and affect health and disease risk. Evidence of past selective pressure on a base, whether highly conserved or fast evolving, is a marker of functional importance. Bases that are unchanged in all mammals may shape phenotypes that are essential for organismal health. Bases that are evolving quickly in some species, or changed only in species that share an adaptive trait, may shape phenotypes that support survival in specific niches. Identifying bases associated with exceptional capacity for cellular recovery, such as in species that hibernate, could inform therapeutic discovery. RATIONALE The power and resolution of evolutionary analyses scale with the number and diversity of species compared. By analyzing genomes for hundreds of placental mammals, we can detect which individual bases in the genome are exceptionally conserved (constrained) and likely to be functionally important in both coding and noncoding regions. By including species that represent all orders of placental mammals and aligning genomes using a method that does not require designating humans as the reference species, we explore unusual traits in other species. RESULTS Zoonomia’s mammalian comparative genomics resources are the most comprehensive and statistically well-powered produced to date, with a protein-coding alignment of 427 mammals and a whole-genome alignment of 240 placental mammals representing all orders. We estimate that at least 10.7% of the human genome is evolutionarily conserved relative to neutrally evolving repeats and identify about 101 million significantly constrained single bases (false discovery rate < 0.05). We cataloged 4552 ultraconserved elements at least 20 bases long that are identical in more than 98% of the 240 placental mammals. Many constrained bases have no known function, illustrating the potential for discovery using evolutionary measures. Eighty percent are outside protein-coding exons, and half have no functional annotations in the Encyclopedia of DNA Elements (ENCODE) resource. Constrained bases tend to vary less within human populations, which is consistent with purifying selection. Species threatened with extinction have few substitutions at constrained sites, possibly because severely deleterious alleles have been purged from their small populations. By pairing Zoonomia’s genomic resources with phenotype annotations, we find genomic elements associated with phenotypes that differ between species, including olfaction, hibernation, brain size, and vocal learning. We associate genomic traits, such as the number of olfactory receptor genes, with physical phenotypes, such as the number of olfactory turbinals. By comparing hibernators and nonhibernators, we implicate genes involved in mitochondrial disorders, protection against heat stress, and longevity in this physiologically intriguing phenotype. Using a machine learning–based approach that predicts tissue-specific cis - regulatory activity in hundreds of species using data from just a few, we associate changes in noncoding sequence with traits for which humans are exceptional: brain size and vocal learning. CONCLUSION Large-scale comparative genomics opens new opportunities to explore how genomes evolved as mammals adapted to a wide range of ecological niches and to discover what is shared across species and what is distinctively human. High-quality data for consistently defined phenotypes are necessary to realize this potential. Through partnerships with researchers in other fields, comparative genomics can address questions in human health and basic biology while guiding efforts to protect the biodiversity that is essential to these discoveries. Comparing genomes from 240 species to explore the evolution of placental mammals. Our new phylogeny (black lines) has alternating gray and white shading, which distinguishes mammalian orders (labeled around the perimeter). Rings around the phylogeny annotate species phenotypes. Seven species with diverse traits are illustrated, with black lines marking their branch in the phylogeny. Sequence conservation across species is described at the top left. IMAGE CREDIT: K. MORRILLmore » « less
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ABSTRACT Mechanical properties including the failure behavior of physically assembled gels or physical gels are governed by their network structure. To investigate such behavior, we consider a physical gel system consisting of poly(styrene)‐poly(isoprene)‐poly(styrene)[PS‐PI‐PS] in mineral oil. In these gels, the endblock (PS) molecular weights are not significantly different, whereas, the midblock (PI) molecular weight has been varied such that we can access gels with and without midblock entanglement. Small angle X‐ray scattering data reveals that the gels are composed of collapsed PS aggregates connected by PI chains. The gelation temperature has been found to be a function of the endblock concentration. Tensile tests display stretch‐rate dependent modulus at high strain for the gels with midblock entanglement. Creep failure behavior has also been found to be influenced by the entanglement. Fracture experiments with predefined cracks show that the energy release rate scales linearly with the crack‐tip velocity for all gels considered here. In addition, increase of midblock chain length resulted in higher viscous dissipation leading to a higher energy release rate. The results provide an insight into how midblock entanglement can possibly affect the mechanical properties of physically assembled triblock copolymer gels in a midblock selective solvent. © 2019 Wiley Periodicals, Inc. J. Polym. Sci., Part B: Polym. Phys.
2019 ,57 , 1014–1026 -
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Abstract Phenotypes are the target of selection and affect the ability of organisms to persist in variable environments. Phenotypes can be influenced directly by genes and/or by phenotypic plasticity. The amphibian‐killing fungus
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