Participating Faculty
Patrick Carter
Name:Patrick Carter
Department:School of Biological Sciences, WSU
Credentials:1993 - Ph.D., University of Colorado, Boulder- Environmental, Population and Organismic Biology
Office:Heald 217
Mailing Address:School of Biological Sciences
PO Box 644236
Pullman, WA 99164-4236

Research Interests

Evolution of the integrated phenotype.

Research Summary

My research interests are in the evolution of the integrated phenotype. The integrated phenotype can be defined in several ways, including examining multiple traits at one time and/or examining one trait at multiple values of an independent index, such as body mass as a function of age. These latter traits are called “function-valued’ traits, and a significant fraction of my current research is focused on them. Major areas of investigation in my lab include the effect of genetic variation and covariation on the rate of evolutionary change, the effect of evolution on standing levels of genetic variance and covariance, and evolution through non-adaptive processes such as genetic drift and correlated responses to selection. Some projects in my lab directly address basic questions related to these issues, while other projects apply knowledge from these areas to questions and problems in trout conservation biology.

Two basic projects in my lab involve using artificially selected organisms to examine the evolution of function-valued traits: locomotor ontogenies and growth curves in mice, and growth curves in flour beetles. In both these projects, reproduction in the experimental populations is manipulated to impose the effects of selection on specific phenotypes in the population. This manipulation allows the examination of responses in the trait that is directly selected and in traits that are correlated with the trait of selection; in addition, the effects of selection on genetic variances and covariances can be tested. In both these projects, reproductive success is manipulated so that evolutionary processes underlying reproductive success can be elucidated and examined.

Two applied projects in my lab use clonal lines of rainbow trout to examine the effects of hatchery domestication on traits related to survival and reproduction. In one study, swim performance ontogenies and growth curves are being compared between highly domesticated clones and clones that are derived from wild populations. Our data so far indicate that highly domesticated trout grow more rapidly but are poor swimmers relative to less domesticated trout, which suggests that highly domesticated trout are likely to have poor survival and reproductive success in natural environments. In the other study, the genotypes and enzyme activities of anti-oxidant enzymes, and oxidative damage to DNA and cell membranes, are being compared between highly domesticated clones and clones that are derived from wild populations. Preliminary indications are that less domesticated lines of trout have higher levels of anti-oxidant enzyme activities and potentially lower levels of oxidative damage. We think this likely leads to a phenotype that suffers less stress, survives better, and has higher reproductive success than highly domesticated trout; we plan to explicitly test this in the near future. In both of these projects, trout reproduction is being manipulated to produce specific sets of clonal lines, and in both projects the ultimate effects of domestication selection on traits related to survival and reproductive success are being examined.

Research Publications

(Note: WSU undergraduate student, ††WSU graduate student, $WSU faculty)

Bellinger††, K, G.H. Thorgaard$, and P.A. Carter. 2014. Effects of domestication on growth and swim performance in rainbow trout. Aquaculture 420-421:154-159. 

Gervini, D. and P.A. Carter. 2014. Warped functional analysis of variance. Biometrics (

Irwin††, K.K., and P.A. Carter. 2014.  Artificial selection on larval growth curves in Tribolium castaneum: Correlated responses and constraints. Journal of Evolutionary Biology (

Attanayake††, R.N., P.A. Carter, D. Jiang, L. Del Río-Mendoza, and W. Chen$. 2013. Sclerotinia sclerotiorum populations infecting canola from China and the United States are genetically and phenotypically distinct.  Phytopathology 103:750-761.

Careau, V.,  M.E. Wolak, P.A. Carter, and T. Garland, Jr. 2013. Limits to behavioral evolution: the quantitative genetics of a complex trait under directional selection. Evolution 67:3102-3119. 

Heink††, A.E., A. Parrish††, G.H. Thorgaard$ and P.A. Carter. 2013. Oxidative stress among SOD-1 genotypes in rainbow trout (Oncorhynchus mykiss). Aquatic Toxicology 144-145: 75-82.  

Irwin††, K.K., and P.A. Carter.  2013. Constraints on the evolution of function-valued traits: A study of growth in Tribolium castaneum. Journal of Evolutionary Biology 26:2633-2643.

Ware, J. V., et al. (2013). "Endocrine rhythms in the brown bear (Ursus arctos): Evidence supporting selection for decreased pineal gland size." Physiol Rep 1(3): e00048.

Serratore, V., M.P. Zalucki and P.A. Carter. 2013. Thermoregulation in molting and feeding Danaus plexippus caterpillars. Australian Journal of Entomology 52:8-13. 

Ware††, J., O. Nelson, C. Robbins$, P.A. Carter, B. Sarver, and H. Jansen$. 2013. Seasonal and daily variation of melatonin and cortisol in the brown bear (Ursus arctos). Physiological Reports 1(3)e00048:1-17. 

Stinchcombe, J.R., J. Beder, P.A. Carter, Gilchrist, G., Gervini, D., Gomulkiewicz, R. $, Hallgrimsson, B., Heckman, N., Houle, D., Kingsolver, J.G., Marquez, E., Marron, J.S., Meyer, K., Mio, W., Schmitt, J., Yao, F., and Kirkpatrick, M. 2012. Genetics and evolution of function-valued traits: understanding environmentally responsive phenotypes. Trends in Ecology and Evolution 27: 637-647.  

Center for Reproductive Biology cited as author affiliation

Washington State University