Dr. Patrick A. Carter
Professor
School of Biological Sciences
Washington State University
Pullman WA 99164-4236
509/335-1447
pacarter@wsu.edu

 

Research Interests | Lab Group | Funding | Publications | Teaching | Pre-Vet Advising | Evolution and Ecology at WSU


Prospective Students

I may be able to take one new Ph.D. student for the fall of 2016.  Please contact me via email if you are interested in doing graduate work in my lab.

Research Interests

My research focuses on the evolution of the integrated phenotype. Using ideas and methods from evolutionary and quantitative genetics, physiological ecology, and comparative physiology, my lab group addresses questions on the evolution of function-valued traits, the correlated responses of multiple traits to selection, and the relative effects of selection and drift on trait evolution. Our goal is to provide a fundamentally deeper understanding of the evolution of the entire phenotype and of evolutionary processes.

Currently four major projects are being conducted:

    • Development and application of function-valued trait methods to the evolution of complex traits. 
    • Evolution of growth curves in laboratory populations of flour beetles using a function-valued approach.
    • Evolution of growth, body shape and swimming performance curves in  domesticated populations of rainbow trout.
    • Functional effects of variation in the Sod-1 gene in rainbow trout.

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Lab Group

I have an active lab group composed of both graduate and undergraduate students. In addition, I encourage independent research by undergraduates once they have spent some time learning basic concepts and techniques used in the lab.

Current Lab Group Members:

    • Kristy Bellinger is a Ph.D. student examining swimming performance ontogenies in rainbow trout.
    • Chris Duke is a Ph.D. student examining swimming performance and fear and startle behavior in rainbow trout.
    • Erin Weise is a new Ph.D. student in the lab who will be working on some aspect of rainbow trout biology.
    • Ben Weideback is a Ph.D. student working on the genetics of flour beetles.
    • Ka’imi Kuenzli is an undergraduate student working on temperature effects on the body shape of rainbow trout.

Past Lab Group Members:

Post-Doctoral Students:

o   Anne Bronikowski conducted much of her post-doctoral research in the lab on longevity and traits related to senescence in lines of mice selected for high voluntary wheel running activity.  Current status: Associate Professor at Iowa State University.

Graduate Students:

o   Kristen Irwin completed her Ph.D. in May 2014.  She examined genetic variation and evolution of growth curves in populations of flour beetles.  Current status: Staff Scientist at École Polytechnique Fédérale de Lausanne, Switzerland.

o   Anna Heink completed her Masters degree in May 2012.  She examined oxidative damage and Sod-1 gene variation in domesticated lines of rainbow trout.  Current status: Research associate at Cincinnati Children's Hospital.

o   Amber Walker completed her Masters degree in May 2010.  She examined genetic variation in Sod-1 enzyme activity in domesticated lines of rainbow trout.  Current status: Research Associate, WSU.

o   Amanda Castro completed her Masters degree in June 2007. She examined effects of herbicides on salamander growth and swimming performance ontogenies.  Current status: Field biologist studying Pronghorn in Arizona.

o   Drew Reinbold completed his MS degree in June 2005.  He studied genetic variation of growth trajectories and swimming performance in domesticated and semi-wild populations of rainbow trout.  Current status: Medical technician for the US Army.

o   Ted Morgan completed his Ph.D. in May 2002. He studied genetic variation in the 4 lines of mice selected for wheel-running behavior and in their 4 lines of controls, using molecular markers and phenotypic variance. In addition he analyzed behavioral and morphological ontogenetic trajectories using function-valued trait analysis. Current status: Associate Professor at Kansas State University.

o   Stephanie Kane completed her MS degree in May 2001. Her research examined the effect of activity and early-age selection on metabolic traits in aged mice. Current status: Project Manager, Social Science Research Unit, University of Idaho.

o   Greg Ragland completed his MS degree in May 2001. His research focused on quantitative genetic analysis of growth trajectories and metamorphic traits in Ambystoma macrodactylum columbianum. Current status: Adjunt Faculty at Notre Dame University.

o   Ed Leber completed his MS degree in December 1999. His research was on stress responses of active and sedentary mice selected for voluntary wheel-running behavior. Current status: Technical support staff at Molecular Probes, Inc.

o   Sara Hall completed her MS degree in May 1999. Her work was on reference and working memory in active and sedentary mice selected for high wheel-running activity.  Sara went on to a DVM program at Tufts University. Current status: Veterinarian.

Undergraduate Students:

o   Kelsi Lakey completed her BS degree in December 2014.  She examined the effect of temperature on body shape in rainbow trout.  Current status: Graduate student at USC.

o   Kyle Johnson completed his BS degree in December 2012.  He worked on quantitative genetics of male mating traits in populations of flour beetles. Current status: DVM student at WSU.

o   Lindsay Pruett completed her BS with Honors in May 2013.  She worked on quantitative genetics of male mating traits in populations of flour beetles. Current status: Doctor of Audiology student at UW.

o   Vanessa Serratore completed her BS with Honors in May 2008.  She examined thermoregulation in feeding and molting Monarch butterfly caterpillars and went on to earn her DVM at WSU.  Current status: Surgical residency at Ohio State Unversity.

o   Abdiaziz Barre completed his BS degree in May 2008.  He examined growth trajectories of amphibian populaitons on the Palouse.  Current status: Medical student at Poznan University of Medical Sciences.

o   Chris Miller completed his BS with Honors in June 2007.  He examined muscle physiology in domesticated and semi-wild lines of rainbow trout.  Current status: Medical doctor.

o   Jamie Harpole completed her BS degree in May 2004.  She studied swimming performance in laboratory strains of rainbow trout.  Jamie went on to the DVM program at WSU. Current status: Veterinarian.

o   Brandi Irwin completed her BS degree in May 2001. Her project was on allozyme diversity in lines of mice selected for high activity.  Brandi went on to medical school at Des Moines University. Current status: Medical doctor.

o   Sharon Thomson completed her BS degree in August 1999. Her research examined activity of the superoxide dismutase enzyme in active and sedentary mice from the selected and control lines.

o   Monica Baze completed her BS with Honors in May 2002. Her research examined nesting behavior in one year old active and sedentary mice from selected and control lines. Current status: graduate student with Dr. Jack Hayes at the University of Nevada, Reno.

o   Lionel Jenkins completed his BS degree in December 2003.  He studied spontaneous tumor occurrence in active and sedentary mice from lines of mice selected for high voluntary activity.  Current status: Laboratory technician at WSU Vet School.

o   Matt Schmit completed his BS degree in May 1999. His research was on nesting behavior in one year old active and sedentary mice from selected and control lines. Current status: zookeeper, Houston Zoo.

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Current and Recent Funding

Current:

Collaborative Research: UBM - Institutional: UI-WSU Program in Undergraduate Mathematics and Biology. $700,000.  National Science Foundation Division of Mathematical Sciences.  PI’s: B. Robison, R.H Dillon; co-PI’s: R.S. Gomulkiewicz, P. Joyce and 6 others; Senior Investigators: P.A. Carter and 7 others (10/1/2010-9/30/2015).  

 

Previous:

National Genomics Research Initiative. $unspecified. Science Education Alliance, Howard Hughes Medical Institute. PI: William Davis; co-PI: P.A. Carter. (6/1/2011-5/31/2014). 

Darwinian Morphometrics: Cross-Topology Registration of Shape. $88,000. National Institute for Mathematical and Biological Synthesis. PI: P.A. Carter. co-PI’s: D.Houle, R.S. Gomulkiewicz, J.S. Marron. (9/1/09-8/30/13).

Physiological Genetics of Oxidative Stress in Rainbow Trout. $29,800. PI: P.A. Carter. USDA Washington-Idaho Aquaculture Initiative. (10/1/2010-9/30/2012).

UBM: Foundation in mathematical biology through interdisciplinary research, training, and curriculum development.  $905,000.  National Science Foundation Emerging Frontiers.  PI: R.S. Gomulkiewicz; co-PIs: Omoto, Dillon, McDonald; Senior Investigators: P.A. Carter and 13 others. (9/15/05-9/14/10).

Function-valued traits in natural populations: variation, selection, and evolution. $2,129,468. National Science Foundation Emerging Frontiers.  PI: R.S. Gomulkiewicz; co-PI: P.A. Carter. (10/1/03-9/30/09).

Exercise induced protection from cancer, III. $3000, Fraternal Order of Eagles. (1 time award).

Integrated approaches to studies of infinite dimensional trait evolution in natural populations. $3,194, National Science Foundation Biocomplexity-Incubation (Supplemental Award). PI: R.S. Gomulkiewicz; co-PI: P.A. Carter.

Exercise induced protection from cancer, II. $2500, Fraternal Order of Eagles.

Dissertation Research: The effect of artificial selection on the evolution of ontogenetic trajectories. $5640, National Science Foundation Population Biology. PI: P.A. Carter; co-PI T.J. Morgan. (7/1/01-6/30/03).

Integrated approaches to studies of infinite dimensional trait evolution in natural populations. $99,999, National Science Foundation Biocomplexity-Incubation. PI: R.S. Gomulkiewicz; co-PI: P.A. Carter. (9/1/00-8/31/03).

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Selected Publications

Campbell, J.M., P.A. Carter, P.A. Wheeler, and G.H. Thorgaard. 2015. Aggressive behavior, brain size and domestication in clonal rainbow trout lines. Behavior Genetics 45:245-254.

Kingsolver, J.G, N. Heckman, J. Zhang, P.A. Carter, J.L. Knies, and J. Stinchcombe. 2015. Genetic variation, simplicity and evolutionary constraints for function-valued traits. The American Naturalist 166: E166-E181. 

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 70:526-353.

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 27:2069-2079.

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.

Bellinger, K, G.H. Thorgaard, and P.A. Carter. 2013. Effects of domestication on growth and swim performance in rainbow trout. Aquaculture (in press). 

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 casteneum. Journal of Evolutionary Biology 26:2633-2643.

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

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.

Reinbold, D., G. H. Thorgaard, and P.A. Carter. 2009. Reduced swimming performance and increased growth are associated with domestication of rainbow trout. Canadian Journal of Fisheries & Aquatic Sciences 66:1025-1032.

Kane, S.L., T. Garland, Jr., and P.A. Carter. 2008. Basal metabolic rate of aged mice is affected by random genetic drift but not by selective breeding for high early-age locomotor activity or chronic wheel access.  Physiological and Biochemical Zoology 81:288-300.

Middleton, K.M., C.E. Shubin. D.C. Moore, P.A Carter, T. Garland, Jr., S.M. Swartz. 2008. The relative importance of  genetics and phenotypic plasticity in dictating bone morphology and mechanics in aged mice: evidence from an artificial selection experiment. Zoology 111:135-147.

Bronikowski, A.M., T.J. Morgan, T. Garland Jr. and P.A. Carter. 2006. The evolution of aging and age-related physical decline in mice selectively bred for high voluntary exercise. Evolution 60: 1494-1508.

Morgan, T.J., M.A. Evans, T. Garland, Jr., J.G. Swallow, and P.A. Carter. 2005.  Molecular and quantitative genetic divergence among populations of house mice with known evolutionary histories. Heredity 94:518-525.

Hochstetler, K.J., T. Garland, Jr., J.G. Swallow, P.A. Carter, and A. Bult-Ito. 2004. Number of arginine-vasopressin neurons in the suprachiasmatic nuclei is not related to level or circadian characteristics of wheel-running activity in house mice.  Behavior Genetics 34:131-136.

Ragland, G.J. and P.A. Carter. 2004.Genetic constraints on the evolution of growth and life history traits in the salamander Ambystoma macrodactylum. Heredity 92:569-578.

Bronikowski, A.M., P.A. Carter, T.J. Morgan, T. Garland Jr., N. Ung, T.D. Pugh, R.Weindruch, and T.A. Prolla. 2003. Lifelong voluntary exercise in the mouse prevents age-related alterations in gene expression in the heart.  Physiological Genomics 12:129-138.

Hochstetler, K.J., T. Garland, Jr., J.G. Swallow, P.A. Carter, and A. Bult-Ito. 2003. Number of arginine-vasopressin neurons in the suprachiasmatic nuclei is not related to level or circadian characteristics of wheel-running activity in house mice.  Behavior Genetics (in press).

Koteja, P., P.A. Carter, J.G. Swallow and T. Garland, Jr. 2003. Food wasting in house mice: variation among individuals, families and genetic lines.  Physiology and Behavior 80:375-383.

Koteja, P., J.G. Swallow, P.A. Carter and T Garland, Jr. 2003. Different effects of intensity and duration of locomotor activity on circadian period. Journal of Biological Rhythms 18:1-11.

Morgan, T.J., T. Garland, Jr., and P.A. Carter. 2003. Ontogenetic trajectories in mice selected for high wheel- running activity. I. Mean ontogenetic trajectories.  Evolution 57:646-657.

Morgan, T.J., T. Garland, Jr., B.L. Irwin, J.G. Swallow, and P.A. Carter. 2003. The mode of evolution of molecular markers in  populations of house mice under artificial selection for locomotor behavior. J Heredity 94:236-242.

Bronikowski, A.M., T.J. Morgan, T. Garland, Jr., and P.A. Carter. 2002. Anti-oxidant gene expression in active and sedentary house mice selected for high voluntary wheel-running behavior. Genetics 161:1763-1769.

Garland, T. Jr., M.T Morgan, J.G. Swallow, J.S. Rhodes, I. Girad, J.G. Belter, and P.A. Carter. 2002. Evolution of a small-muscle polymorphism in lines of house mice selected for high activity levels. Evolution 56:1267-1275.

Girad, I.A., J.G. Swallow, P.A. Carter, P. Koteja, J.S. Rhodes, and T. Garland, Jr. 2002. Maternal-care behavior and life-history traits in house mice (Mus domesticus) artificially selected for high voluntary wheel-running activity.  Behavioral Processes 57:37-50.

Klomberg, K.F., T. Garland, Jr., J.G. Swallow, and P.A. Carter. 2002. Aggression, plasma testosterone levels, and testis size in house mice artificially selected for high activity levels. Physiology and Behavior 77:27-38.

Nunez, N.P., P.A. Carter, and G.G. Meadows. 2002. Alcohol consumption promotes body weight loss in melanoma-bearing mice. Alcoholism: Clinical and Experimental Research 26:617-626.

Thomson, S.L., T. Garland, Jr., J.G. Swallow, and P.A. Carter. 2002. Response of Sod-2 enzyme activity to selection for high voluntary wheel running. Heredity 88:52-61.

Bronikowski, A. M., P. A. Carter, J. G. Swallow, I. A. Girard, J. S. Rhodes, and T. Garland, Jr. 2001. Open-field behavior of house mice artificially selected for high voluntary wheel running. Behavior Genetics 31:309-316.

Kingsolver, J.G., R.S. Gomulkiewicz, and P.A. Carter. 2001. Variation, selection, and evolution of function-valued traits.  Genetica 112-113:87-104.

Koteja, P., J.G. Swallow, P.A. Carter, and T. Garland, Jr. 2001.  Maximum cold-induced food consumption in mice selected for high locomotor activity: implications for evolution of energy budgets of endotherms. The Journal of Experimental Biology 204(6) 1177-1190.

Swallow, J.G., P. Koteja, P.A. Carter, and T. Garland, Jr. 2001. Food consumption and body composition in mice selected for high wheel running activity. Journal of Comparative Physiology B 171: 651-659.

Carter, P.A., J.B. Mitton, T.D. Kocher, and J.R. Coehlo. 2000. Maintenance of the Adh polymorphism in tiger salamanders II. Differences in biochemical function among allozymes.  Functional Ecology 14:70-76.

Carter, P.A., J.G. Swallow, S. Davis, and T. Garland, Jr. 2000. Nesting behavior of house mice (Mus domesticus) selected for increased wheel-running activity.  Behavior Genetics 30(2):85-94.

Koteja, P., J.G. Swallow, P.A. Carter, and T. Garland, Jr. 2000. Individual variation and repeatability of maximum cold-induced energy assimilation in house mice. Acta theriologica 45(4):455-470.

Rhodes, J.S., P. Koteja, J.G. Swallow, P.A. Carter, and T. Garland, Jr.  2000.  Body temperatures of house mice artificially selected for high voluntary wheel-running behavior: repeatability and effect of genetic selection.  Journal of Thermal Biology 25:391-400.

Carter, P.A., T. Garland, Jr., M.R. Dohm, and J.P Hayes. 1999. Genetic variation and correlations between genotype and locomotor physiology in outbred laboratory house mice (Mus domesticus). Comparative Biochemistry and Physiology 123:157-164.

Koteja, P., T. Garland, Jr., J. K. Sax, J. G. Swallow, and P. A. Carter. 1999. Behaviour of house mice artificially selected for voluntary wheel running. Animal Behaviour 58:1307-1318.

Koteja, P., J.G. Swallow, P.A. Carter, and T. Garland, Jr. 1999. Energy cost of wheel running in house mice: implications for coadaptation of locomotion and energy budgets. Physiological and Biochemical Zoology 72:238-249.

Swallow, J.G., T. Garland, Jr., P. Koteja, and P.A. Carter. 1999. Artificial selection for increased wheel-running activity in house mice results in decreased body mass. The Journal of Experimental Biology 202:2513-2520.

Zhan, W-Z, J.G. Swallow, T. Garland, Jr., D.N. Proctor, P.A. Carter and G.C. Sieck. 1999. Effects of genetic selection and voluntary activity on the medialgastrocnemius muscle in house mice. Journal of Applied Physiology 87: 2326-2333.

Coleman, M.A., T. Garland, Jr., C.A. Marler, S.S. Newton, J.G. Swallow, and P.A. Carter. 1998.  Glucocorticoid response to forced exercise in laboratory house mice (Mus domesticus). Physiology and Behavior 63:279-285.

Krugner-Higby, L.A., A. Gendron, T. Garland, Jr., J.G. Swallow, P.A. Carter, and J.J. Lee. 1998. Eosinophylic polymyositis in a mouse. Contemporary Topics in Laboratory Animal Science 37(6): 94-97.

Swallow, J.G., P.A. Carter, and T. Garland, Jr. 1998. Artificial selection for increased wheel-running behavior in house mice. Behavior Genetics 28:227-237.

Swallow, J.G., T Garland, Jr., P.A. Carter, W.Z. Zhan, G.C. Sieck. 1998. Effects of voluntary activity and genetic selection on aerobic capacity in house mice (Mus domesticus). Journal of Applied Physiology 84(1):69-76.

Carter, P.A. 1997. Maintenance of the Adh polymorphism in tiger salamanders I. Genotypic differences in time to metamorphosis in extreme oxygen environments. Heredity 78:101-109.

Mitton, J.B., P.A. Carter and A. DiGiacomo. 1997. Resting oxygen consumption varies among lactate dehydrogenase genotypes in the sow bug, Porcellio scaber. Proc. R. Soc. Lond. B 264:1543-1546.

Watt, W.B., K. Donohue, and P.A. Carter. 1996. Adaptation at specific loci. VI. Divergence vs. parallelism of polymorphic allozymes in molecular function and fitness-component effects among Colias species. Molecular Biology and Evolution 13(5):699-709.

Garland, T., Jr, and P.A. Carter. 1994. Evolutionary physiology. Annual Review of Physiology 56:579-621.

Mitton, J.B., D.J. Zelenka, and P.A. Carter. 1994. Selection of breeding stock in pigs favours 6PGD heterozygotes. Heredity 73:177-184.

Zalucki, M.P., J.M. Hughes, J.M. Arthur, and P.A. Carter. 1993. Seasonal variation at four loci in a continuously breeding population of Danaus plexippus L. Heredity 70:205-213.

Hughes, J.M., M.P. Zalucki, and P.A. Carter. 1992. Consistent differences among individual Monarch butterflies (Danaus plexippus L.) in flight activities at low temperatures. Biological Journal of the Linnean Society 47:217-231.

Carter, P.A., M.P. Zalucki, and J.M. Hughes. 1989. Genetic variation in a continuously breeding population of Danaus plexippus L.: An examination of heterozygosity at four loci in relation to activity times.  Heredity 63:191-194.

Watt, W.B., C. Kremen, and P.A. Carter. 1989. Testing  the "mimicry" explanation for the Colias "alba" polymorphism: Patterns of co-occurrence of Colias and Pierine butterflies. Functional Ecology 3:193-199.

Carter, P.A., and W.B. Watt. 1988.  Adaptation at specific loci. V. Metabolically adjacent enzyme loci may have very distinct experiences of selective pressures.  Genetics 119:913-924.

Zalucki, M.P., J.M. Hughes, and P.A. Carter. 1987. Genetic variation in Danaus plexippus L.: habitat selection or differences in activity times?  Heredity 59: 213-221.

Watt, W.B., P.A. Carter, and K. Donohue. 1986. Females' choice of "good genotypes" as mates is promoted by an insect mating system. Science 233:1187-1190.

Watt, W.B., P.A. Carter, and S.M. Blower. 1985. Adaptation at specific loci. IV. Differential mating success among glycolytic allozyme genotypes of Colias butterflies. Genetics 109:157-175.

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Teaching

In Fall semesters I teach Human Physiology:

o    Human Physiology (Biology 251) (every year)

In Spring semesters I teach either Quantitative Genetics (Biol 521) or Environmental Physiology (Biol 561):

o    Environmental Physiology (offered in 2011)

o    Quantitative Genetics (offered in 2012)

 

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Pre-Vet Advising

I am a pre-Vet advisor for the School of Biological Sciences.  We offer a BS in Zoology with a pre-Vet option that prepares students for entry into a DVM program.  Students in this option take all the courses required for entry into Vet School.  In addition, they take classes such comparative anatomy, comparative physiology, animal development, parasitology, animal behavior and mammology which prepare them for entry into Vet School and which train them in the methods and approaches of modern biology.  More information about the DVM program at WSU can be found on this link:  Prospective Vet Student Page.  If you would like to talk to me about pre-Vet studies generally or the Zoology pre-Vet option specifically, please call or email me to set up an appointment.

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Evolution and Ecology at WSU

WSU has a very active group in Ecology and Evolutionary Biology in the School of Biological Sciences. We gather weekly for a presentation of recent work by one of our members (EcoEvo Lunch, Tuesdays at noon); once a week EEGR (Evolution, Ecology and Genetic Reading group) meets, and monthly we meet, along with colleagues from the Biological Sciences Department at the University of Idaho, for the PEES (Palouse Ecology, Evolution and Systematics) seminar.

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page last updated August 2015.