| Theodore Garland, Jr.
Distinguished Professor of Biology
Ph.D., Univ. of California,
*** NEW!!! PBZ has a call for papers for a Functional Morphology and Biomechanics: Form Follows Function, with manuscripts due by 15 April 2018.
Physiological and Biochemical Zoology
has recently published Focused Issues on:
Early-life Effects on the Adult Phenotype: a Comparative Perspective
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.
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. Currently, I am a co-PI (along with Mark Chappell) on two grants (NSF, NIH) obtained by Wendy Saltzman to study paternal care in California mice (Peromyscus californicus).
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
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)
- Social Behavior of Desert Iguanas (Dipsosaurus dorsalis) (Jennifer M. Singleton)
1. Selective Breeding for High
Voluntary Wheel Running. Publications
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 implications 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 (June 2018) approaching 86 generations. 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 brain dopamine and endocannabinoid signaling in the selected lines, which may render them a useful model for studies of human attention-deficit hyperactivity disorder (ADHD) and understanding the control of motivated behaviors. 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. Recently, we have begun studying early-life effects on adult physical activity.
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); Daniel Pomp (genetics); Vincent Careau (quantitative genetics and animal model analyses). (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 As an additional outreach effort, 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.
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
We have conducted studies of both individual and interspecific
in snake and lizard
locomotion and life history traits, and have previously had N.S.F. funding
for this work At
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,
and horned lizards [Phrynosoma]).
Outgroups include Crotaphytidae,
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. Current efforts are channeled through IDEA, the UCR Institute for the Development of Educational Applications. For example, "Nature or Nurture? Heritability in the Classroom" can be found on the IDEA website.
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
Fall 2016 Online version Syllabus
Online Lecture Course Introduction
Online Lecture Example: Adaptation and the Comparative Method
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  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's EVOLVE license plate from Wisconsin. This can be interpreted in various ways, including as a command!
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
VIDEO - 2003 Toyota MR2 Spyder - Adams Motorsports Park Time Attack
Current Graduate Students
Ralph Lacerda de Albuquerque (EEOB) - started January 2014 - webpage Portugese- ecomorphology and locomotor physiology of lizards
Marcell D. Cadney (EEOB) - started fall 2015 - early-life effects on the adult phenotype
Alberto A. Castro (EEOB) - started fall 2016 - skeletal evolution
David Hillis (GGB) - started fall 2016 - genetic analyses of the High Runner lines of mice
Samantha Kubica (EEOB) - started fall 2018; currently rotating in our lab and co-supervised by Mark Springer
Monica P. Louis (EEOB) - started fall 2016 - microbiome of the High Runner mice
Margaret P. Schmill (Neuroscience) - started fall 2015 - behavior and neurobiology of the High Runner mice
Nicolas L. Schwartz (EEOB) - started fall 2017 (coadvised with Tim Higham) skeletal evolution of the High Runner mice
Jessica L. Tingle (EEOB) - started fall 2015 - webpage - locomotion, morphology, and biomechanics of snakes
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. home page the interplay between form, function, selection and evolutionary history. Currently Assistant Professor at Pacific Lutheran University
Vincent Careau - 2011-2012 - home page - currently Assistant Professor at the University of Ottawa
Some Former Graduate Students
Wendy Acosta (EEOB) - Finished Ph.D. fall 2015 - voluntary activity, diet choice, obesity, exercise.
Currently treaching at local community colleges.
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 Ph.D. Dec. 2001 and is presently Adjunct Assistant Professor at the Univ. of Arizona.
Gerald C. Claghorn (EEOB) - Finished Ph.D. Dec. 2016 - webpage - vertebrate exercise physiology and neurobiology.
Currently a statistical analyst in the health insurance industry.
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 - Finished Ph.D. 2011 old home page - vertebrate functional morphology and performance; herpetology.
After a postdoc at Harvard with Jonathon Losos, Gabe obtained a faculty position at Santa Monica City College.
Robert M. Hannon old UCR home page - Finished in Genetics, Genomics, and Bioinformatics graduate program Dec. 2010.
Subsequently faculty at Northern Virginia Community College. R.I.P.
Sean Harrington (joint doctoral program in Evolutionary Biology with SDSU) - was here at UCR for 2013-2014
academic year - clade diversification, focusing on pit vipers - coadvised by Tim Higham and Tod Reeder
Layla Hiramatsu (EEOB) - Finished Ph.D. fall 2017 - webpage Currently postdoc with Frank Chan in Germany.
Jarren C. Kay (EEOB) - Finished Ph.D. fall 2017 - muscle physiology. Currently postdoc with Stephen Secor at the University of Alabama.
Brooke K. Keeney. Finished Ph.D. 2011 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 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, now tenured, 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, became
a Lecturer in Kinesiology at USC, and is now on the faculty at Chaffee College.
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.
She is now an Assistant Professor at St. Mary's College of Maryland.
Thomas H. Meek old home page- evolutionary and exercise physiology. Tom was a postdoc at the Unviersity of Washington and now works at Novo Nordisk.
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.
He is currently on the faculty at the Universidad Andrés Bello (UNAB), Departamento de Ecología, Santiago, Chile.
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 Associate Professor at the University of Illinois
Jennifer M. Singleton (EEOB) - Finished Ph.D. Dec. 2018 - webpage - behavioral ecology and physiology of desert iguanas
Ronald W. Sutherland - home page - continued his Ph.D. at the Univ. of Wisconsin with Tim Moermond.
Zoe Thompson (Neuroscience) - Finished summer 2017 - neurobiology of exercise. Currently NIH postdoc at the University of Michigan.
Theodore Garland, III
|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|
|26 Aug. 2005 Swimming|
|13 Oct. 2007 with pet Varanus acanthurus|
|23 Aug. 1010|
Jaden Lee Garland
|18 Jan. 2002||Art_2004_Sep_9|
|21 July 2003|
|"My First Roadkill" Gopher Snake 31 Aug. 2003|
|23 Oct. 2005|
|26 Aug. 2005 Swimming|
|April 2006 with local Rosy Boa|
|16 Jan. 2011 sledding at Forest Falls, CA|
"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"
"Two Not-So-Little Boys 11 June 2010
Loki (younger half-brother of Makodo)
Pictures of (click on name) and Home Pages of Some Former Graduate Students and Postdocs:
- Nicola M. Anthony, University of New
- Anne M. Bronikowski, Iowa State University
- Kevin E. Bonine, University
- Patrick A. Carter, Washington State
- Ramon Diaz-Uriarte, Bioinformatics Unit at
the Spanish National Cancer Center (CNIO)
- Michael R. Dohm, formerly University of Hawaii at Hilo - as of Sept. 2005 is Assistant
Professor at Chaminade University, Oahu
- Isabelle A. Girard, formerly University of Wisconsin-Stevens Point, currently University of Wisconsin-Madison
- Jack P. Hayes, University of Nevada-Reno
- Wendy L. Hodges, formerly at University of Texas of the Permian Basin
- Thomas H. Meek, Novo Nordisk, Seattle, Washington
- Gad Perry, Texas Tech University
- Enrico L. Rezende, a formerly a Senior Lecturer
in Zoology at the University of Roehampton in London;
currently Associate Professor at U. Andres Bello in Santiago, Chile
- Justin S. Rhodes, University of Illinois at Urbana-Champaign
- John G. Swallow, formerly University of South Dakota, currently University of Colorado, Denver
- Jonathan L. Temte, University of Wisconsin-Madison
All Publications and PDF files
Publications on the Mouse Selection Experiment
Video of Mice Running on Wheel (Girard et al. 2001)
Garland Public Lecture on "Born to Run: Evolution of Hyperactivity in Mice" 29 Oct. 2009
Last updated by T.G. 12 January 2019