Comparative and Evolutionary Physiology

Much our research uses comparative approaches to answer fundamental questions about how organisms adapt and evolve to better cope in challenging environments. What are the integrative mechanisms of adaptive evolution and phenotypic plasticity? To what extent do interacting and integrated traits (e.g., steps in the O2 transport pathway) evolve in concert? Are changes in some traits more likely to constitute the first steps of adaptation? Do evolved changes in some traits alter the adaptive value of changes in other traits? Does the evolution of organismal performance arise via similar underlying mechanisms across different lineages? Our research examines these questions in different vertebrate groups.

Evolution and Plasticity in High-Altitude Deer Mice

High-altitude environments are among the most challenging environments experienced by terrestrial animals. Temperatures are much colder at high altitude than at sea level, increasing the O2 demands of thermogenesis, but high altitude also has thin air with low O2 availability (hypoxia) to support these high O2 demands. In fact, O2 levels atop the highest mountains in the world are scarcely sufficient to support life in many species – they are so low that unacclimatized lowland animals can be rendered comatose within minutes. However, every mountain range contains animals that have overcome this challenge and thrive at high altitudes.

The emphasis of much of our current work in this area focusses on deer mice (Peromyscus maniculatus), the species with the broadest altitudinal distribution of any North American mammal. In high-altitude populations, natural selection favours a high capacity to produce body heat (‘thermogenic capacity’), a key performance trait that is underpinned by several intersecting areas of physiology (cardiorespiratory physiology and the O2 transport pathway, mitochondrial metabolism, thermoregulation, etc.). We maintain lab colonies of mice derived from wild populations at high altitude (4300m in the Rocky Mountains) and low altitude (Great Plains) to study these processes in captivity.

Deer mouse in our lab colony, rewarded with a marshmallow for a job well done (photo courtesy of Kayla Garvey).

Mice can be held in control conditions reflective of sea level, and can also be exposed to conditions that simulate the cold and hypoxic conditions at high altitude. Comparisons between environments are used to uncover mechanisms of phenotypic plasticity, which we examine during adulthood (acclimation), early life (developmental plasticity), and even across generations (trans-generational plasticity, epigenetics). Comparisons between high-altitude and low-altitude populations are used to elucidate the mechanisms underlying evolutionary adaptation and the evolution of phenotypic plasticity.

The figure above shows a hypothetical scenario for how phenotypic plasticity and evolutionary adaptation can affect phenotypes and contribute to fitness in a new challenging environment, such as during the initial colonization of high altitude (Storz and Scott 2021). (A) In lowland populations, fitness in low-altitude environments varies as a function of phenotype (i.e., trait values) and fitness is maximized at a ‘fitness optimum’ (vertical dashed line). (B-D) In high-altitude environments, the fitness optimum may be shifted relative to that in low-altitude environments (solid vertical line). (B) Plasticity could move the population mean phenotype to the new optimum (black arrow), such that there is no opportunity for natural selection to further improve fitness, and the trait does not evolve. (C) Plasticity could move the population mean phenotype part way to the new optimum (black arrow), in which case selection on genetically based variation (red arrow) can then shift it the rest of the way. (D) Plasticity can in some cases be maladaptive and move the population mean phenotype further from the optimum (black arrow), in which case selection on genetically based variation (red arrow) can lead to a large compensatory change in phenotype that opposes the ancestral plasticity response.

Evolutionary Physiology of High-Altitude Birds

The Andes of South America contain a great diversity of species that have adapted to high-altitude environments. Many high-altitude lakes contain several species of ducks, geese, and other waterbirds that have independently colonized high altitude. Some of these taxa are well established and endemic to high altitude, whereas some others are relatively new colonists of the high-altitude environment. We compare these high-altitude taxa to their close low-altitude relatives to uncover mechanisms of high-altitude adaptation that are conserved across species, to examine how time at high altitude affects high-altitude phenotypes, and to understand how and why differences between species may alter the responses to high-altitude in different lineages.


Phylogenetic relationships of bird taxa used in the comparative analyses of the O2 transport pathway and mitochondrial physiology in high-altitude natives (Dawson et al. 2020). Multiple pairs of high- and low-altitude taxa are used to evalute whether common physiological changes have arisen across high-altitude birds (e.g., convergent evolution), or whether birds in different lineages exhibit distinct changes at high altitude.

Key Publications:

Scott GR, Dalziel AC. 2021. Physiological insight into the evolution of complex phenotypes: aerobic performance and the O2 transport pathway of vertebrates. J Exp Biol. 224, jeb210849.

Wearing OH, Scott GR. 2021. Commentary: Hierarchical reductionism approach to understanding adaptive variation in animal performance. Comp Biochem Physiol B. 256, 110636.

Milsom WK, Scott GR, McCracken KG, Frappell PB. 2021. Different strategies for convective O2 transport in high altitude birds: a graphical analysis. Comp Biochem Physiol A. 253, 110871.

Storz JF, Scott GR. 2021. Phenotypic plasticity, genetic assimilation, and genetic compensation in hypoxia adaptation of high-altitude vertebrates. Comp Biochem Physiol A. 253, 110865.

Storz JF, Scott GR. 2019. Life ascending: mechanism and process in physiological adaptation to high-altitude hypoxia. Annu Rev Ecol Evol Syst. 50, 503-526.

Storz JF, Cheviron ZA, McClelland GB, Scott GR. 2019. Evolution of physiological performance capacities and environmental adaptation: insights from high-elevation deer mice (Peromyscus maniculatus). J Mammal. 100, 910-922.

McClelland GB, Scott GR. 2019. Evolved mechanisms of aerobic performance and hypoxia resistance in high-altitude natives. Annu Rev Physiol. 81, 561-583.

Ivy CM, Scott GR. 2015. Control of breathing and the circulation in high-altitude mammals and birds. Comp Biochem Physiol A. 186, 66-74.

Storz JF, Scott GR, Cheviron ZA. 2010. Phenotypic plasticity and genetic adaptation to high-altitude hypoxia in vertebrates. J Exp Biol. 213, 4125-4136.