Evolution, Ecology & Organismal Biology

Theodore Garland, Jr.

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Office: (951) 827-3524
Fax: (951) 827-4286
2366 Spieth Hall
Email: tgarland@ucr.edu

Theodore Garland, Jr., Distinguished Professor

Ph.D., University of California, Irvine, 1985


Random News and Announcements:

NEW!!!  PBZ now has a call for papers for a Focused Issue on Ecoimmunology, with manuscripts due by 1 May 2018

I became Editor in Chief of the journal Physiological and Biochemical Zoology on 1 July 2014. Read my Editorial describing changes we are implementing at the journal

Hueyfest: a Symposium in Honor of Ray Huey's contributions to Physiological Ecology: 4 Oct. 2013 in Seattle 
My Talk in Honor of Ray, with help from Steve Adolph

JoeFest: a symposium in Honor of Joe Felsenstein's career
My Talk at the symposium: "Zen and the Art of Phylogenetically Independent Contrasts"

Lab Homepage

IDEA, the Institute for the Development of Educational Applications

YouTube Videos on the High Runner Mice

Lab Wiki (members only)

Profile in Google Scholar Citations 

Inquiry-Based Middle School Lesson Plan -- "Born to Run: Artificial Selection Lab"
PDF version
 on my homepage (latest, with live links to all other files

Nature or Nurture? Heritability in the Classroom (college or advanced high school)

Simulating Random Genetic Drift (college or high school)

I am featured in FAIL LAB Episode One: Evolution, part of an educational video series funded through the Discovery Digital Network and intended to target teenagers

Textbook Features on our Research:
Box on phylogenetically independent contrasts in 2004 Evolutionary Analysis text by Freeman and Herron 
Two-page Box on the mouse selection experiment in 2006 Comparative Physiology text by Moyes and Schulte

Two-pages on the mouse selection experiment in 2014 Evolutionary Analysis text by Herron and Freeman (with contributions by Hodin, Miner, Sidor)
Page discussing a selection experiment on mouse maternal aggression by Stephen Gammie that Ted helped with in 2014 Evolutionary Analysis text by Herron and Freeman 

Curriculum Vitae (not current)

Evolutionary Physiology Links

Experimental Evolution Links

Table of Contents for:
Garland, T., Jr., and M. R. Rose, eds. 2009. Experimental evolution:
     concepts, methods, and applications of selection experiments.
     University of California Press, Berkeley, California. [here's the entire book as a PDF file]

Caveats About Undergraduate Research

How to Structure and Name Data Files


     Most of the research in my laboratory involves the evolution of complex phenotypes.  Through empirical, theoretical, and methodological studies, we are also helping to develop the field of evolutionary physiology (Links: e.g., see Annual Review of Physiology [1994] 56:579-621 [PDF file]). 
     Physiology is the study of how organisms work.  Evolution is the study of how organisms have changed (genetically) across generations.  Thus, evolutionary physiology is the study of how and why the way organisms work has changed over time.  For example, does the way an organism work constrain the way it may evolve?  Answers to such questions require a deep understanding of both proximate and ultimate mechanisms, including phenotypic plasticity and early-life effects.  Accordingly, my graduate students come through the Evolution, Ecology, and Organismal Biology Graduate Program in the Department of Biology, as well as the Neuroscience Graduate ProgramGenetics, Genomics & Bioinformatics Graduate Program, and the Graduate Program in Biomedical Sciences.  I also participate in the 
Evolutionary Biology Joint-Doctoral Program with San Diego State University.  I serve as UCR co-Associate Director of NERE, the Network for Experimental Research on Evolution.  As physiology cannot properly be understood in isolation from genetics, morphology, biochemistry, and behavior, my general approach is integrative and hence crosses traditional boundaries between disciplines. 

     Our laboratory is equipped to make a variety of sophisticated whole-animal physiological and behavioral measurements.  We have concentrated on activity metabolism because many natural behaviors (e.g., escaping from predators, foraging) depend crucially on capacities for locomotion.  I have worked primarily on lizards, snakes, and small mammals, but I recognize the value of both "model" and "non-model" systems and am always amenable to work with other organisms.  Although most of our efforts involve exercise physiology and locomotor behavior, graduate students have worked on a variety of other projects, including:  the evolution of reproductive timing in seals; behavioral/physiological ecology and conservation biology of desert tortoises; reproductive and conservation biology of lizards on a Spanish island; development of a monitoring program for endangered species of small mammals.  In addition, we have collaborated with scientists from many countries, including Argentina, Australia, Belgium, Brazil, Canada, Chile, France, Germany, Mexico, The Netherlands, Spain, and The United Kingdom. 

PDAP: Phenotypic Diversity Analysis Programs  (software to perform phylogenetically based statistical analyses)

PDTREE module in Mesquite  (JAVA-based software to perform phylogenetically based statistical analyses) (Documentation as Microsoft Word file)

PHYSIG  (MatLab programs to perform phylogenetically based statistical analyses)

PHYLOGR  (R language code to perform phylogenetically based statistical analyses)

Photos by Ted Garland:  The Thorny Devil (Moloch horridus) in Australia [fullsize image], Mouse on running wheel as used in the selection experiment [fullsize image], Lizard (Lacerta agilis) in France on treadmill for endurance test [fullsize image].


Click here for a Complete Publication List with PDF Files

Selected Recent Publications:

  • Brashares, J., T. Garland, Jr., and P. Arcese. 2000. Phylogenetic analysis of coadaptation in behavior, diet, and body size in the African antelope. Behavioral Ecology 11:452-463. Abstract   [PDF file]
  • Clobert, J., A. Oppliger, G. Sorci, B. Ernande, J. G. Swallow, and T. Garland, Jr. 2000. Trade-offs in phenotypic traits: endurance at birth, growth, survival, predation, and susceptibility to parasitism in a lizard, Lacerta viviparaFunctional Ecology 14:675-684. Abstract  [PDF file]
  • Irschick, D. J., and T. Garland, Jr. 2001. Integrating function and ecology in studies of adaptation: investigations of locomotor capacity as a model system. Annual Review of Ecology and Systematics 32:367-396.  Abstract  [PDF file]
  • Girard, I., and T. Garland, Jr. 2002. Plasma corticosterone response to acute and chronic voluntary exercise in female house mice. Journal of Applied Physiology 92:1553-1561. [PDF file]
  • Perry, G., and T. Garland, Jr. 2002. Lizard home ranges revisited: effects of sex, body size, diet, habitat, and phylogeny. Ecology 83:1870-1885.  [PDF file]
  • Hutcheon, J. M., J. A. W. Kirsch, and T. Garland, Jr. 2002. A comparative analysis of brain size in relation to foraging ecology and phylogeny in the Chiroptera. Brain, Behavior and Evolution 60:165-180.  Abstract  [PDF file]
  • Abbott, D. H., E. B. Keverne, F. B. Bercovitch, C. A. Shively, S. P. Mendoza, W. Saltzman, C. T. Snowdon, T. E. Ziegler, M. Banjevic, T. Garland, Jr., and R. M. Sapolsky. 2003. Are subordinates always stressed? A comparative analysis of rank differences in cortisol levels among primates. Hormones and Behavior 43:67-82. [PDF file]
  • Blomberg, S. P., T. Garland, Jr., and A. R. Ives. 2003. Testing for phylogenetic signal in comparative data: behavioral traits are more labile. Evolution 57:717-745.  [PDF file]
  • Rhodes, J. S., H. van Praag, S. Jeffrey, I. Girard, G. S. Mitchell, T. Garland, Jr., and F. H. Gage. 2003. Exercise increases hippocampal neurogenesis to high levels but does not improve spatial learning in mice bred for increased voluntary wheel running. Behavioral Neuroscience 117:1006-1016. [PDF file]
  • Johnston, I. A., D. A. Fernandez, J. Calvo, V. L. A. Vieira, A. W. North, M. Abercomby, and T. Garland, Jr. 2003. Reduction in muscle fibre number during the adaptive radiation of notothenioid fishes: a phylogenetic perspective. Journal of Experimental Biology 206:2595-2609. [PDF file]
  • Perry, G., K. LeVering, I. Girard, and T. Garland, Jr. 2004. Locomotor performance and social dominance in male Anolis cristatellusAnimal Behaviour 67:37-47. [PDF file]
  • Rezende, E. L., F. Bozinovic, and T. Garland, Jr. 2004. Climatic adaptation and the evolution of basal and maximum rates of metabolism in rodents. Evolution 58:1361-1474.  [PDF file]
  • Al-kahtani, M. A., C. Zuleta, E. Caviedes-Vidal, and T. Garland, Jr. 2004. Kidney mass and relative medullary thickness of rodents in relation to habitat, body size, and phylogeny. Physiological and Biochemical Zoology 77:346-365. (plus online Appendix B).  [PDF file]
  • Bonine, K. E., T. T. Gleeson, and T. Garland, Jr. 2005. Muscle fibre-type variation in lizards (Squamata) and phylogenetic reconstruction of hypothesized ancestral states. Journal of Experimental Biology 208:4529-4547.  [PDF file]
  • Zhang, Y., T.-S. Lee, E. M. Kolb, K. Sun, X. Lu, F. M. Sladek, G. S. Kassab, T. Garland, Jr., and J. Y.-J. Shyy. 2006. AMP-activated protein kinase is involved in endothelial nitric-oxide synthase activation in response to shear stress. Arteriosclerosis, Thrombosis, and Vascular Biology 26:1281-1287.  [PDF file]   Faculty of 1000 Selection!
  • Garland, T., Jr., and S. A. Kelly. 2006. Phenotypic plasticity and experimental evolution. Journal of Experimental Biology 209:2344-2361. [PDF file]
  • Spoor, F., T. Garland, Jr., G. Krovitz, T. M. Ryan, M. T. Silcox, and A. Walker. 2007. The primate semicircular canal system and locomotion. Proceedings of the National Academy of Science U.S.A. 104:10808-10812.  [PDF file]   UCR Press Release
  • Buchwalter, D. B., D. J. Cain, C. A. Martin, L. Xie, S. N. Luoma, and T. Garland, Jr. 2008. Aquatic insect ecophysiological traits reveal phylogenetically based differences in dissolved cadmium susceptibility. Proceedings of the National Academy of Science U.S.A. 105:8321-8326.  [PDF file
  • Huey, R. B., C. A. Deutsch, J. J. Tewksbury, L. J. Vitt, P. E. Hertz, H. J. Álvarez Pérez, and T. Garland, Jr. 2009. Why tropical forest lizards are vulnerable to climate warming. Proceedings of the Royal Society B 276:1939-1948.  [PDF file]
  • Kelly, S. A., D. L. Nehrenberg, J. L. Peirce, K. Hua, B. M. Steffy, T. Wiltshire, F. Pardo Manuel de Villena, T. Garland, Jr., and D. Pomp. 2010. Genetic architecture of voluntary exercise in an advanced intercross line of mice. Physiological Genomics 42:190-200. [PDF file]
  • Garland, T., Jr., S. A. Kelly, J. L. Malisch, E. M. Kolb, R. M. Hannon, B. K. Keeney, S. L. Van Cleave, and K. M. Middleton. 2011. How to run far: Multiple solutions and sex-specific responses to selective breeding for high voluntary activity levels. Proceedings of the Royal Society B 278:574-581.  [PDF file]
  • Garland, T., Jr., H. Schutz, M. A. Chappell, B. K. Keeney, T. H. Meek, L. E. Copes, W. Acosta, C. Drenowatz, R. C. Maciel, G. van Dijk, C. M. Kotz, and J. C. Eisenmann. 2011. The biological control of voluntary exercise, spontaneous physical activity and daily energy expenditure in relation to obesity: human and rodent perspectives. Journal of Experimental Biology 214:206-229.  [PDF file]
  • Careau, V. C., and T. Garland, Jr. 2012. Performance, personality, and energetics: correlation, causation,and mechanism. Physiological and Biochemical Zoology 85:543-571.  [PDF file]
  • Schutz, H., H. A. Jamniczky, B. Hallgrimsson, and T. Garland, Jr. 2014. Shape-shift: semicircular canal morphology responds to selective breeding for increased locomotor activity. Evolution 68:3184-3198. [PDF file]
  • Acosta, W., T. H. Meek, H. Schutz, E. M. Dlugosz, K. T. Vu, and T. Garland, Jr. 2015. Effects of early-onset voluntary exercise on adult physical activity and associated phenotypes in mice. Physiology & Behavior 149:279-286. [PDF file] [We currently have funding from the NIH to continue these studies.]

Selected Teaching:


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Tel: (951) 827-1012

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Evolution, Ecology and Organismal Biology (EEOB)
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Tel: (951) 827-5903
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Graduate Program
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Tel: (951) 827-4116
Fax: (951) 827-5517
E-mail: eeobgrad@ucr.edu