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Theodore Garland, Jr.
Theodore Garland, Jr.
Distinguished Professor of Evolution, Ecology, and Organismal Biology (EEOB)
Office: 2366 Spieth Hall
Phone: (951) 827-3524
E-mail: tgarland@ucr.edu
Degree: Ph.D., University of California, Irvine, 1985
Profile in Google Scholar Citations
AD Scientific Index 2023 (top 2% in world!?)
AI-generated Podcast of Schwartz et al. 2025
Evolutionary Physiology Labs & Links
Lab Wiki (members only)
UCR Center for Cannabinoid Research
GenBioPAC: Genetic and Biological factors that regulate Physical Activity Consortium
From 331. Galván, I., T. S. Schwartz, and T. Garland, Jr. 2021. Evolutionary physiology at +30: has the promise been fulfilled? BioEssays 2021:2100167.
Halloween 2023. We all dressed up as Ted, but I forgot to wear shorts and thongs (a.k.a. flipflops or toe-spreaders).
Natalie Whitehead presenting her poster at the American Physiological Summit in Long Beach, California, on 6 April 2024.
Garland lab and partners at the Progressive Party 9 Feb 2024.
LAB NEWS:
May 2025: Haley Cabrera and William Lampman pass their qualifying exams!
May 2025: Ted receives the The 2026 August Krogh Distinguished Lectureship of the Comparative & Evolutionary Section (CEPS) of the American Physiological Society (APS), sponsored by Novo Nordisk Foundation. This is awarded to a distinguished scientist who has made major and meritorious contributions to the scientific areas represented by CEPS and is their most prestigious award.
Jessica Tingle accepts faculty position at Brown University!
Marcell Cadney accepts faculty position at California State University, Long Beach!
NEW!!! 363. Schwartz, N. E., M. P. Schmill, M. D. Cadney, A. A. Castro, D. A. Hillis, M. P. McNamara, J. O. Rashid, W. Lampman, D. F. DeLaCruz, B. D. Tran, N. L. Trutalli, and T. Garland, Jr. 2025. Maternal exercise opportunity before, during, and after pregnancy alters maternal care behavior, offspring development, and offspring survival, but has few effects on offspring adult physical activity or body composition. Physiology & Behavior 291:114752. [Drive PDF file] https://www.sciencedirect.com/science/article/pii/S0031938424003007?via%3Dihub
NEW!!! 360. Schwartz, N. and T. Garland, Jr. 2024. A meta-analysis of whole-body and heart mass effect sizes from a long-term artificial selection experiment for high voluntary exercise. Journal of Experimental Biology 227:jeb249213. 10.1242/jeb.249213 [Drive PDF file]
NEW!!! 343. Clark, C. J., J. R. Hutchinson, and T. Garland, Jr. 2023. The Inverse-Krogh Principle: all organisms are worthy of study. Physiological and Biochemical Zoology 96:1–16. [Drive PDF file] https://www.journals.uchicago.edu/doi/10.1086/721620
NEWish!!! 336. McNamara, M. P., M. D. Cadney, A. A. Castro, D. A. Hillis, K. M. Kallini, J. C. Macbeth, M. P. Schmill, N. E. Schwartz, A. Hsiao, and T. Garland, Jr. 2022. Oral antibiotics reduce voluntary exercise behavior in athletic mice. Behavioural Processes 199:104650. [Drive PDF file] [https://doi.org/10.1016/j.beproc.2022.104650]
NEWish!!! 333. Garland, T., Jr., C. J. Downs, and A. R. Ives. 2022. Invited Perspective: Trade-offs (and constraints) in organismal biology. Physiological and Biochemical Zoology 95:82-112. [Drive PDF file] [https://www.journals.uchicago.edu/doi/full/10.1086/717897]
NEWish!!! 331. Galván, I., T. S. Schwartz, and T. Garland, Jr. 2021. Evolutionary physiology at +30: has the promise been fulfilled? BioEssays 2021:2100167. [Drive PDF file] [https://onlinelibrary.wiley.com/doi/10.1002/bies.202100167]
NEWish!!! Garland, Jr., T. 2022. Born to Run: Experimental Evolution of Exercise Behavior and Physiology in a Vertebrate Model Organism. In preparation. Riverside, California. [Draft of Chapter 1]
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. xvii + 730 pages.
Entire Book is Available Here as a fairly large PDF file
Research Overview:
Most of the research in my laboratory involves the evolution of complex traits. 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 E.E.O. Biology, as well as the Neuroscience Graduate Program, Genetics, 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 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. Moreover, physical activity is crucial for human health and well-being. 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.
News & Announcements:
New Website on the High Runner Mice!
January 2024 - Physiological and Biochemical Zoology was officially renamed Ecological and Evolutionary Physiology, with Ted remaining as Editor in Chief.!
February 2023 - Over at PBZ, we organized all of the Focused Collections we've done over the years. Lots of cool topics, such as trade-offs, conservation physiology, evolution of endothermy, individual variation, and ecoimmunology.
November 2022 - Ted was named as an Associate Editor of Behavior Genetics!
August 2022 - William Lampman joins the lab as a new M.S. student in EEOB; subsequently moved into the Ph.D. program.
April 2022 - Lucas Pineiro joins the lab as a new Ph.D. student in GGB
August 2021 - Haley Cabrera joins the lab as a new Ph.D. student in EEOB
July 2021 - Natalie Holt and I received a new 4-year NSF grant titled "Trade-offs, limits to adaptation, and multiple solutions: rules elucidated by selective breeding for high activity levels"
July 2014 - I became Editor in Chief of the journal Physiological and Biochemical Zoology. Read my Editorial describing changes that we implemented at the journal. I signed on for another 5-year term in 2019.
Links & Resources:
IDEA, the Institute for the Development of Educational Applications
YouTube Videos on the High Runner Mice
Lab Wiki (members only)
Lab Wiki OLD VERSION (members only)
All Publications and PDF files
Publications on the Mouse Selection Experiment
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
JoeFest: a Symposium in Honor of Joe Felsenstein's career, 22 June 2017 in Portland
Slides from My Talk: "Zen and the Art of Phylogenetically Independent Contrasts"
Hueyfest: a Symposium in Honor of Ray Huey's contributions to Physiological Ecology, 4 Oct. 2013 in Seattle
Slides from My Talk, with help from Steve Adolph : "What We Learned from Ray Huey: What We Learned from Ray Huey: Squamates, Stones, Squamates, Stones, Science, and Style"
Videos:
Video of Mice Running on Wheel (Girard et al. 2001) [.mov file that downloads automatically]
Video (1 min 23 sec): Dr. Garland discussing "How to run far: Multiple solutions and sex-specific responses to selective breeding ..." PDF file
Video (7 min 50 sec): Dr. Garland discussing the High Runner mice in a short video
Education & Outreach:
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 Lesson Plan (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:
Software:
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].
Some Publications Since the Last Millenium:
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. [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 vivipara. Functional Ecology 14:675-684. [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. [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. [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 cristatellus. Animal 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]
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]
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]
Hillis, D., L. Yadgary, G. M. Weinstock, G. Pardo−Manuel de Villena, D. Pomp, A. Fowler, S. Xu, F. Chan, and T. Garland, Jr. 2020. Genetic basis of aerobically supported voluntary exercise: results from a selection experiment with house mice. Genetics 216:781-804. [PDF file]
Lamkin, D. M., and T. Garland, Jr. 2020. Translating preclinical research for exercise oncology: take it to the VO2max. Frontiers in Oncology 10:Article 575657. [PDF file]
Malisch, J. L., T. Garland, Jr., L. Claggett, L. Stevenson, E. A. Kohl, and H. B. John-Alder. 2020. Living on the edge: Glucocorticoid physiology in desert iguanas (Dipsosaurus dorsalis) is predicted by distance from an anthropogenic disturbance, body condition, and population density. General and Comparative Endocrinology 294:113468. [PDF file]
Verberk, W. C. E. P., P. Calosi, F. Brischoux, J. I. Spicer, T. Garland, Jr., and D. T. Bilton. 2020. Universal metabolic constraints shape the evolutionary ecology of diving in animals. Proceedings of the Royal Society B 287:20200488. [PDF file]
Cadney, M. D., L. Hiramatsu, Z. Thompson, M. Zhao, J. C. Kay, J. M. Singleton, R. L. Albuquerque, M. P. Schmill, W. Saltzman, and T. Garland, Jr. 2021. Effects of early-life exposure to Western diet and voluntary exercise on adult activity levels, exercise physiology, and associated traits in selectively bred High Runner mice. Physiology & Behavior 234:113389. [PDF file]
Selected Teaching:
Biology 105 "Evolution" Online Winter 2017
Biology 105 "Evolution" Fall 2020
Biology 174 "Ecological and Evolutionary Physiology" syllabus for Winter 2008
Biology 174 "Ecological and Evolutionary Physiology" syllabus for Winter 2014
Biology 220 "Evolutionary Physiology" syllabus for Spring 2012
Garland, T. 2022. Review: Ginetta G56 GTA. https://www.hagerty.com/media/new-car-reviews/review-ginetta-g56-gta/
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