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  • UC Riverside
  • College of Natural and Agricultural Sciences

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
 tgarland "at" 
 Office Phone:  951-827-3524
 Lab Phone:  951-827-5724
 Facsimile:  951-827-4286

        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.

        My graduate students and I participate in both the Evolutionary Biology and the Physiology tracks within the Department of Biology, as well as the Evolution and Ecology Graduate Research Unit, 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 phenotypes.  Through empirical, theoretical, and methodological studies, we are also helping to found the 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 pressures 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 am currently 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 current Ph.D. study, is beginning comparative studies of locomotor performance in relation to sexual selection in this group, with emphasis on 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.  This project, funded by the National Science Foundation and the National Institutes of Health, allows direct tests of the long-standing idea that behavior evolves "first."  We are now (Feb. 2004) in generation 37.  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, 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).

        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 include 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 (Univ. of Calgary: 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.

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.  Our latest series of programs (PHYSIG), written mainly in MatLab, implements multiple tests for phylogenetic signal and phylogenetic generalized least squares (PGLS) models (Regressionv2.m of Lavin et al., 2008).

        Current and recent collaborators on this research include Drs. Simon P. Blomberg (Canberra), Anthony R. Ives (Univ. of Wisconsin), Peter E. Midford (Univ. of Arizona), Ramon Diaz-Uriarte (Spain), and Francois-Joseph Lapointe (Univ. of Montreal).  Two current projects involve developing techniques for (1) incorporating measurement error into phylogenetic analyses and (2) testing hypotheses about convergent evolution.  For the latter, we are collaborating with Dr. Eric R. Pianka (Univ. of Texas at Austin) and focusing on a comparison of the lizards Phrynosoma and Moloch.

3.  Ecological and Evolutionary Physiology of Lizards.  We are conducting studies of both individual and interspecific variation in lizard locomotion and life history traits, also with recent funding from the N.S.F.  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.

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)

Picture Gallery (and a few links):

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 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)

Recent Postdoctorals
    Wendy L. Hodges   C.V.  Home Page  X-ray CT scans of Phrynosoma cornutum
    Fernando Gomes    C.V.  behavioral endocrinology of selected lines of mice, amphibian ecophysiology

Current Postdoctorals
    Kevin M. Middleton  home page - vertebrate locomotion and bone biology
    (NIH NRSA postdoc with Sharn M. Swartz at Brown University, cosponsored by TG)

Current Graduate Students
    Gabriel E. A. Gartner  home page - vertebrate functional morphology, performance, and selection in the wild; herpetology
    Rob Hannon  home page - Genetics, Genomics, and Bioinformatics graduate program
    Brooke K. Keeney  home page - neurobiology, endocrinology, and behavior in our selected lines of mice
    Scott A. Kelly  home page - mammalian ecological and evolutionary physiology, phenotypic plasticity
    Erik Kolb  home page - mammalian physiology and neurobiology
    Jessica Malisch (formerly Bunkers)  home page - behavioral endocrinology of mice and lizards
    Tom Meek  home page- evolutionary and exercise physiology
    Christopher E. Oufiero  home page - ecological and evolutionary physiology, life history of poecilid fishes

Some Former Graduate Students
   Mohammed Al-Kahtani finished Ph.D. July 2003
   Kevin E. Bonine finished his Ph.D. Dec. 2001 and is presently Adjunct Assistant Professor
                               at the Univ. of Arizona
   Richard S. King and his study organism, the massasauga rattlesnake (Sistrurus catenatus)
   Guo Li finished his Masters in August 2002 and is now in a statistics program at the Univ. of Mchigan
   Enrico L. Rezende  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 and is currently a postdoc with John C. Crabbe at
                               Oregon Health & Science University
   Ronald W. Sutherland continued his Ph.D. at the Univ. of Wisconsin with Tim Moermond

    Theodore Garland, III (Theo - born 24 October 1999)
27 Oct. 1999 Art_2004_May_13_1  
15 Jan. 2000 Art_2004_July_23_1  
31 Jan. 2000 movie Art_2004_Sep_29  
24 March 2000 Art_2004_Oct_8  
March 2002 Art_2004_Oct_26  
April 2003    
26 Aug. 2005 Swimming    
13 Oct. 2007 with pet Varanus acanthurus    

    Jaden Lee Garland (born 7 October 2001)
18 Jan. 2002 Art_2004_Sep_9  
21 July 2003    
April 2003    
"My First Roadkill" Gopher Snake 31 Aug. 2003    
Aug. 2004    
23 Oct. 2005    
26 Aug. 2005 Swimming    
April 2006 with local Rosy Boa    

    "Two Little Boys 4 Oct. 2002"
    "Two Little Boys 22 Nov. 2003"
    "Jaden_Coleen_Theo Garland 30 Oct. 2005"
    "Jaden, Ted, and Theo Garland at the San Diego Zoo April 2006"
    Big Kitty

Research Diagrams (feel free to use with due credit)
Evolutionary Physiology -- unique questions
    Phenotypic Hierarchy -- expansion of S. J. Arnold's (1983) morphology, performance, fitness paradigm
    Star_Phylogeny_vs_Hierarchical_Tree -- phylogenetic comparative methods
    Phylogenetic Pseudoreplication -- phylogenetic comparative methods
    Phylogenetically_Independent_Contrasts_1.jpg -- phylogenetic comparative methods

Pictures of (click on name) and Home Pages of Some Former Graduate Students and Postdocs:

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Last updated by T.G. 3 July 2008