Department of Biology:

Ted Garland, Theodore Garland, Experimental Evolution, Physiological Ecology, Evolutionary Physiology, Phenotypic Plasticity, Adaptive Plasticity, Comparative Physiology, Exercise Physiology, Animal Behavior, Quantitative Genetics, Biostatistics, Herpetology, Locomotion, Conservation Biology

Theodore Garland, Jr.

Professor of Biology

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

Contact Information

E-mail: tgarland "at" ucr.edu
Office Phone: 951-827-3524
Lab Phone: 951-827-5724
Facsimile: 951-827-4286

Experimental Evolution
Phylogenetic Comparative Methods
Physiological Ecology
Evolutionary Physiology
Comparative Physiology
Exercise Physiology
Functional Morphology
Phenotypic Plasticity
Sexual Selection
Animal Behavior
Quantitative Genetics
Biostatistics
Herpetology
Locomotion
Conservation Biology

Caveats About Undergraduate Research

How to Structure and Name Data Files

Lab Wiki (members only)

Prospective graduate students should send me a letter of inquiry (email is fine) indicating their areas of interest and describing their previous research or other relevant experience. Please also indicate your grade-point average and G.R.E. scores (if available). Although I am well aware that these are not necessarily good indicators of the likelihood of success in graduate school, they are important in campus-wide competitions for fellowships and so forth. In addition, you may want to read these essays by Stephen C. Stearns (1987, Bull. Ecol. Soc. Amer. 68:145-150), Raymond B. Huey (1987, Bull. Ecol. Soc. Amer. 68:150-153), Brian W. Witz (1994, Bull. Ecol. Soc. Amer. 75:176-177), and Dan Binkley (1988. Some advice for graduate advisors. Bull. Ecol. Soc. Amer. 69:10-13 [I don't have a PDF of that one, but here is another from his web) as well as these by Massimo Pigliucci on how to choose a PhD project and a checklist for graduate students. Finally, here is an extensive website on resources and advice for graduate students, maintained by Scott Keogh.

        Most of my graduate students are in the Physiology Track of the Evolution, Ecology, and Organismal Biology Graduate Program. I also participate in the Genetics, Genomics & Bioinformatics Graduate Program, the Graduate Program in Biomedical Sciences, and the Network for Experimental Research on Evolution (NERE: a University of California Multicampus Research Project). Work in our laboratory is aimed primarily at understanding the evolution of complex traits. Through empirical, theoretical, and methodological studies, we are also helping to found the relatively new field of evolutionary physiology (e.g., see Annual Review of Physiology, 1994, 56:579-621 [PDF file]; Annual Review of Ecology and Systematics, 2000, 31:315-341). Comparative physiology and physiological ecology have developed as fields with great interest in how and why phenotypic diversity has evolved. Many studies in these fields, however, have been conducted with less than state-of-the-art approaches and analytical techniques in terms of evolutionary biology (e.g., two-species comparative studies, lack of common-garden controls). As well, evolutionary biologists have rarely worked with physiological traits, largely because of logistical problems (e.g., metabolic rates cannot be measured on museum specimens).

        My research program, therefore, focuses on the evolution of physiological systems, as well as their phenotypic plasticity. As physiology cannot properly be understood in isolation from behavior, biochemistry, and morphology, my general approach is integrative and collaborative, and 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 locomotion and activity metabolism (exercise physiology) because many natural behaviors (e.g., escaping from predators, foraging) depend crucially on capacities for locomotion. In addition, physical activity itself can have a variety of effects on behavior and physiology (e.g., training).

        We emphasize two complementary approaches, quantitative genetic and comparative. The former allows both predictions of short-term responses to hypothetical selection and tests in real time via laboratory selection experiments (one type of experimental evolution). Comparative studies, on the other hand, allow quantification of what actually has happened in a given group of organisms over evolutionary time.

        I have worked primarily on lizards, snakes, and small mammals (plus the occasional cow). However, I recognize the value of "model systems" of all types, and am always amenable to work with other organisms. For example, I was a co-PI on an NSF grant with David Reznick and Mark Springer to study the evolution of placentas and other life history traits in poecilid fishes. Chris Oufiero, a recent Ph.D. student, performed comparative studies of locomotor performance in relation to sexual selection in swordtails.

Although most of the work in our lab has involved exercise physiology and locomotor behavior, graduate students have worked on a diversity of other projects, including:

Evolution of Reproductive Timing in Seals
(J. L. Temte, Ph.D. in Zoology at Wisconsin)
Behavioral/Physiological Ecology and Conservation Biology of Desert Tortoises
(S. J. Bulova, Ph.D. in Zoology at Wisconsin)
Reproductive and Conservation Biology of Lizards on a Spanish Island
(J. G. Swallow, M.S. in Conservation Biology and Sustainable Development at Wisconsin)
Development of a Monitoring Program for Endangered Species of Small Mammals in Southern
Wisconsin Prairie Fragments (N. M. Anthony, M.S. in C.B.S.D. at Wisconsin)
Evolutionary and Phenotypic Plasticity of Mammalian Kidney
(M. A. Al-kahtani, Ph.D. in Zoology at Wisconsin)

Current research in our laboratory is focused in three main areas:

1. Selective Breeding for High Voluntary Wheel Running.    Publications to date.
From a base population of randombred Hsd:ICR house mice, we have undertaken an artificial selection experiment to increase levels of voluntary wheel-running behavior, which serves as a model of voluntary exercise and has important implicatins for human biology. This project, funded primarily by the National Science Foundation and sometimes by the National Institutes of Health, allows direct tests of the long-standing idea that behavior evolves "first." We are now (Feb. 2011) in generation 65. As wheel running has evolved, mainly by increased running speed (movie), we have tested for correlated responses in a series of continuous-valued, polygenic traits (e.g., body mass, litter size, open-field behavior, sprint running speed on a photocell-timed racetrack, endurance, maximal oxygen consumption, basal metabolic rate, hematocrit, hemoglobin, heart mass, gastrocnemius muscle mass, liver mass, corticosterone and thyroid hormone levels, activities of aerobic and anaerobic indicator enzymes). Although our main focus has been understanding how exercise physiology evolves in concert with voluntary activity levels, this experiment has virtually limitless potential to uncover relationships among different aspects of behavior. For example, we have also examined nesting and parental-care behavior, resident-intruder and predatory aggression, open-field behavior, and learning. We are exploring brain structure and function through pharmacological and neuroanatomical studies. Our results indicate alterations of dopamine in the selected lines, which may render them a useful model for studies of human attention-deficit hyperactivity disorder (ADHD). In addition, the selected lines exhibit some unique responses when given a diet high in fat (Western diet), and so we are examining their utility as a model for possible resistance to the metabolic syndrome and diabetes.

        This has always been a collaborative project (Pat, Ted, John in 1995) and because of the range of possible correlated responses that we anticipate may occur, we work with various laboratories, including several at other universities. Off-campus collaborators have included Drs. Patrick A. Carter (Washington State Univ.: aging), Gary M. Diffee (Univ. of Wisconsin: muscle biology), Patricia A. Freeman (Univ. of Nebraska: skeletal morphometrics), Sharon M. Swartz (Brown Univ.: bone properties), Stephen C. Gammie (Univ. of Wisconsin: neurobiology), Helga Guderley (Laval Univ., Quebec: metabolic biochemistry), David J. Paterson (Oxford Univ.: cardiovascular physiology, magnetic resonance imaging), Steven F. Perry (Univ. of Bonn: electron microscopy to quantify lung untrastructure); Douglas A. Syme and Russell T. Hepple (currently McGill Univ.: muscle properties). (We have even found a radio station that may be interested in sponsoring our work.)

        We are also exploring the relative magnitude of the effects of the genetic selection that we have imposed ("nature") as compared with phenotypic plasticity that occurs ontogenetically when mice have access to running wheels and can self-train ("nurture"). A related question is whether mice from selected lines may have reduced or enhanced "trainability," which would constitute a genotype-by-environment interaction. For these experiments, we often house mice either with or without wheel access.

We have also been developing middle school lesson plans that use research material from the selection experiment. One such lesson that uses digital photographs of skeletal elements has been released:
Inquiry-Based Middle School Lesson Plan -- "Born to Run: Artificial Selection Lab"
PDF version

2. Phylogenetically Based Statistical Methods (comparative methods). Phylogenies are essential for understanding the origin and maintenance of biological diversity, such as the origin of endothermy in vertebrates. We seek to promote the use of rigorous phylogenetic methods by developing and testing statistical methods for the analysis of comparative (interspecific) data. With the assistance of a separate N.S.F. grant, we make available at no cost computer programs to perform the necessary analyses (PDAP: Phenotypic Diversity Analysis Programs; PDTREE module in Mesquite; PHYLOGR). Analyses include Felsenstein's (1985) method of phylogenetically independent contrasts (with emphasis on branch-length diagnostics and transformations), Monte Carlo computer simulation to obtain phylogenetically correct ("PC") null distributions, and techniques for ancestor reconstruction.

A recent series of programs with Tony Ives (PHYSIG), written mainly in MatLab, implements multiple tests for phylogenetic signal (Blomberg et al., 2003) and phylogenetic generalized least squares (PGLS) models (Regressionv2.m of Lavin et al., 2008). Other Matlab programs allow you to incorporate "measurement error" (in the general sense) into phylogenetic analyses (Ives et al., 2007) and to perform phylogenetic logistic regression (Ives et al., 2010).

Collaborators on this research have included Drs. Simon P. Blomberg (Univ. of Queensland), Anthony R. Ives (Univ. of Wisconsin), Peter E. Midford (Univ. of Arizona), Ramon Diaz-Uriarte (Spain, home page), and Francois-Joseph Lapointe (Univ. of Montreal). One current projects involves developing techniques for testing hypotheses about convergent evolution, and has involved Dr. Eric R. Pianka (Univ. of Texas at Austin) and a focus on a comparison of the lizards Phrynosoma and Moloch.

3. Ecological and Evolutionary Physiology of Squamate Locomotion. We have conducted studies of both individual and interspecific variation in snake and lizard locomotion and life history traits, and have previously had N.S.F. funding for this work At present, we are emphasizing the family Phrynosomatidae (and select outgroups), which is diverse in North America and includes three major subclades (fence lizards [Sceloporus] and their allies, sand lizards [Callisaurus, Cophosaurus, Holbrookia, Uma], and horned lizards [Phrynosoma]). Outgroups include Crotaphytidae, Anguidae, and Teiidae.

We are testing whether capacities for speed (measured on a photocell-timed racetrack and on a high-speed treadmill) and stamina (measured on a motorized treadmill) show an ineluctable evolutionary trade-off, as predicted from physiological and biomechanical models, and we are relating locomotor capacities to variation in limb proportions and muscle fiber-type composition (in collaboration with Dr. Todd. T. Gleeson at the University of Colorado, Boulder). This work formed the basis of Kevin E. Bonine's Ph.D. dissertation. We will also be attempting to relate variation in locomotor abilities to interspecific variation in field movement, as indexed by daily movement distance, home range area, and typical foraging velocities. Future studies will examine variation in maximal rates of oxygen consumption.

Jessica Malisch (formerly Bunkers), a former graduate student, is also investigating corticosterone levels and clutch size of desert iguanas (Dipsosaurus dorsalis) in relation to distance from a road, as a possible tool for indicating the "health" of populations in human-impacted (disturbed) habitats. We collaborate with Dr. Henry B. John-Alder on this project.

We are also performing a phylogenetic analysis of life-history variation in the Phrynosomatidae. Eventually, we want to understand the nature of possible trade-offs between life-history and locomotor phenotypes.

Now that Dr. Timothy Higham has joined UCR, we are planning for some collaborative efforts in this area, and we would welcome inquiries from prospective graduate students.

4. Development of Inquiry-Based Lesson Plans for Middle School and High School Science Students. In collaboration with Dr. Tricia Radojcic of Bella Vista Middle School (Murrieta, California), and with support from a grant from the National Science Foundation, we have developed the first of what may prove to be a series of lesson plans that involve research materials from our lines of selectively bred lines of High Runner mice. The first such plan (Born to Run: Artificial Selection Lab) is now available from the Evolution and the Nature of Science Institutes website. Subsequent plans may involve analyses of digital research photographs derived from magnetic resonance imaging brain scans (e.g., see Kolb et al. 2013) or histological analyses.

List of Publications (includes links to some abstracts and most PDF files)

Curriculum Vitae (includes links to various people and places, as well as photos)

Biology 105 "Evolution"
    Fall 2003 Syllabus
Biology 174 "Ecological and Evolutionary Physiology"
    Winter 2002 Syllabus
    Fall 2002 Syllabus
    Fall 2004 Syllabus
    Winter 2006 Syllabus
    Winter 2007 Syllabus

Biology 282 "Seminar in Genetics and Evolution" - Phylogenies and the Comparative Method
   Winter 2003 Syllabus
   Spring 2006 Syllabus

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

PDTREE module in Mesquite (JAVA-based software to perform phylogenetically based statistical analyses)

PHYSIG (MatLab programs to perform phylogenetically based statistical analyses)

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

Research Diagrams (feel free to use with due credit)
    Evolutionary Physiology -- unique "emergent" questions
    Phenotypic Hierarchy -- expansion of S. J. Arnold's (1983) morphology, performance, fitness paradigm
    Phenotypic Plasticity Genotypes to Phenotypes -- illustrating genotype-by-environment interaction & reactions norms
    Star_Phylogeny_vs_Hierarchical_Tree -- phylogenetic comparative methods
    Phylogenetic Pseudoreplication -- phylogenetic comparative methods
    Phylogenetically_Independent_Contrasts_1.jpg -- phylogenetic comparative methods (how to compute Felsenstein's [1985] phylogenetically independent contrasts
    Phenotypic_Plasticity_Genotypes_to_Phenotypes.jpg -- phenotypic plasticity, illustrating reaction norms and genotype-environment interaction

Picture Gallery (and a few links):

Ted Garland catches his first snake in Wisconsin, summer 1966
Justin Rhodes, Pat Carter, Isabelle Girard, John Swallow, Ted Garland (at the Evolution meetings in Madison, Wisconsin 1999
Pat Carter, Ted Garland, and John Swallow in 1995
What Ted Garland did before he became a scientist
Why Ted Garland had to leave Wisconsin before his kids got too old
Ted Garland at the Dec. 1995 ASZ Meetings in Washington, D.C.
John Swallow at the Dec. 1995 ASZ Meetings in Washington, D.C.
Michael Rose, Steve Britton, Ted Garland, and Al Bennett at the Experimental Biology meetings in Washington, D.C., 30 April 2007
Graduation_June_2008_Morris_Maduro_Ted_Garland_Mark_Chappell_John_Rotenberry (MM)

Current Graduate Students
    Wendy Acosta (EEOB) - started fall 2008 - voluntary activity, diet choice, obesity, exercise
    Gerald C. Claghorn (EEOB) - started fall 2010 - webpage - vertebrate exercise physiology and neurobiology
    Layla Hiramatsu (EEOB) - started fall 2012 - webpage
    Jarren C. Kay (EEOB) - started fall 2011
    Jennifer Singleton (EEOB) - started fall 2012 - webpage - behavioral ecology and physiology of desert iguanas
    Zoe Thompson (Neuroscience) - started fall 2012 - neurobiology of exercise

Current Postdoc

Former Postdocs
Jack P. Hayes - 1988-1990 - home page - Currently Associate Professor at the University of Nevada, Reno
Aurora Martinez de Castilla Munoz - 1990, 1993 - Currently employed at the Qatar Foundation. Ted assisted with a biodiversity survey there in October 2012.
Patrick A. Carter - 1993-1996 - home page - Currently Associate Professor at Washington Staste University
Isabelle Girard - 1999-2002 - Currently employed in the Research Animal Resources Center at the University of Wisconsin-Madison
Gad Perry - 1999-2002 - home page - Currently Associatre Professor at Texas Tech University
Simon P. Blomberg - 2001-2002 - home page - Currently Lecturer and Consultant Statistician in the School of Biological Sciences at the University of Queensland
Wendy L. Hodges - 2002-2004 - C.V. old home page X-ray CT scans of Phrynosoma cornutum
Fernando Gomes - 2003-2005 - old C.V. home page behavioral endocrinology of selected lines of mice, amphibian ecophysiology
Kevin M. Middleton - 2005-2007 - home page - vertebrate locomotion and bone biology
        (NIH NRSA postdoc with Sharon M. Swartz at Brown University, cosponsored by TG)
         currently Associate Professor at the University of Missouri
Heidi Schutz - 2008-2010 - C.V.    old home page the interplay between form, function, selection and evolutionary history. Heidi will assume a faculty position at Pacific Lutheran University in summer 2013.
Vincent Careau - 2011-2012 - home page - currently Alfred Deakin Postdoctoral fellow at Deakin University

Some Former Graduate Students
Mohammed Al-Kahtani - Finished Ph.D. July 2003 and then returned to a position at King Faisal University in Saudi Arabia.
Kevin E. Bonine - home page - Finished his Ph.D. Dec. 2001 and is presently Adjunct Assistant Professor at the Univ. of Arizona.
Ramon Diaz-Uriarte - home page - Finished his Master's in Biometry at Wisconsin in 1992 and is currently Associate Professor
   at the Department of Biochemistry, Universidad Autónoma de Madrid (UAM).
Gabriel E. A. Gartner old home page - vertebrate functional morphology and performance; herpetology.
   Currently postdoc at Harvard with Jonathon Losos.
Robert M. Hannon old UCR home page - Finished in Genetics, Genomics, and Bioinformatics graduate program Dec. 2010.
   Currently faculty at Northern Virginia Community College.
Brooke K. Keeney old home page - neurobiology, endocrinology, and behavior in our selected lines of mice.
      Brooke Keeney joins LFHCfS
Scott A. Kelly old home page   home page finished his Ph.D. summer 2008. Here at UCR, Scott worked on mammalian ecological
   and evolutionary physiology, and phenotypic plasticity. After a postdoc with our collaborator Daniel Pomp
   at the University of North Carolina, Scott became an Assistant Professor at Ohio Wesleyan University.
Richard S. King and his study organism, the massasauga rattlesnake (Sistrurus catenatus).
Erik M. Kolb old UCR home page - mammalian physiology and neurobiology. Finished Sept. 2010 and is
    now a Lecturer in Kinesiology at USC.
Guo Li finished his Masters in August 2002, enrolled in a statistics program at the Univ. of Michigan, and is now a statistician at the University of Washington.
Jessica Malisch (formerly Bunkers) old UCR home page - behavioral endocrinology of mice and lizards.
    After a postdoc with another of our collaborators, Creagh Breuner at the University of Montana, Jessica was a Visiting Assistant Professor
    in the Joint Science Department at Claremont McKenna College/Pitzer College/Scripps College.
Thomas H. Meek old home page- evolutionary and exercise physiology. Tom is now a postdoc at the Unviersity of Washington.
Christopher E. Oufiero old home page home page- ecological and evolutionary physiology, sexual selection and locomotor performance in
    swordtail fishes. Chris was a postdoc with Peter C. Wainwright at U.C. Davis and is now Assistant Professor at Towson University.
Enrico L. Rezende old UCR home page - [Enrico left for a postdoc in Spain in Sept. 2005, but I am maintaining a home page
    here until he gets a new one] activity physiology of selected lines of house mice, mammalian ecophysiology.
Justin S. Rhodes finished his Ph.D. in Dec. 2002, was then a postdoc with John C. Crabbe at
   Oregon Health & Science University, and is now an Assistant Professor at the University of Illinois
Ronald W. Sutherland - home page - continued his Ph.D. at the Univ. of Wisconsin with Tim Moermond.