Principles of Animal Behavior


Course Format

Helpful Hints


The focus of this course will be to understand the evolution of behavior. The course will begin with a discussion of how phylogenetic information can be used to reconstruct the evolution of behavior. Since evolution requires genetic change, we will then examine evidence that genes influence behavior and learn how to measure their effects. Genes, however, are inadequate to describe all behavioral change because of the inherent flexibility of behavior. For this reason we will discuss the conditions under which nongenetic modes of transmission, such as learning and imitation, should evolve. We will then examine the process of communication in animals and ask how and why animals make and receive signals using different sensory modalities. In the next part of the course predictions about which behavior to expect in a particular ecological or social setting will be made by using a variety of optimization techniques such as optimization and game theory. We will learn how to predict when competition among animals for space, food, and mates should lead to behavioral and morphological adaptations. We will also discover the situations which favor cooperative or altruistic behavior. Throughout the course we will consider how the study of animal behavior may help us to understand our own behavior as well as to better conserve threatened or endangered species.

Course format

LECTURES: two per week on Mondays and Wednesdays from 11-11:50, PLS 1140.

DISCUSSIONS: each Friday at 9, 10 or 11 in 1168 or 1113 PLS. One or two recent scientific papers will be assigned for each discussion and can downloaded from the Reading List page. Beginning with the second week, each seminar will be run by two students who will have prepared questions, that must be turned in to the instructor in charge of the section at the beginning of the period, to encourage discussion and clarify the assigned reading. Everyone is expected to read every paper every week! Discussion performance will count 15% of the course grade. Performance will be evaluated by attendance, weekly participation, and preparation as a discussion leader. See Hints for discussion preparation below.

TEXTBOOK: Perspectives on Animal Behavior, second edition (2001), by Goodenough, McGuire and Wallace is the text for the course. In the schedule of classes I have listed the chapters or page numbers that relate to the material covered in each lecture. The book is intended to supplement, not necessarily duplicate, lecture material. Supplemental textbooks are available and should be consulted if you find you are having difficulty with some of the quantitative material in the course.

LECTURE OUTLINES: To supplement your lecture notes, I will post my powerpoint presentations on the website. Students in the past have found these outlines helpful when studying for exams.

PROBLEM SETS: Most of the central concepts regarding the evolution of behavior are based on theoretical ideas that have empirical support. In lecture I will attempt to show you, at least heuristically, how some of these theoretical conclusions have been reached and provide examples. I will require that you solve some algebraic problems using the techniques I present in class. To aid you in attaining competency in these tasks, I will occasionally ask you to solve problem sets. Answers will be posted on the course website.

GRADES: Your final course grade will be based on the sum of scores for all assignments/exams. The distribution of points across assignments/exams is as follows:

Midterm I 100 points
Midterm II 100 points
Grant Proposal 100 points
Discussion 75 points
Final 125 points

I will assign letter grades, including pluses and minuses, on a curve based on how your total score ranks relative to others in the class. After each exam I will indicate how I would assign letter grades to help you track your progress.

EXAMS: There will be two midterms, October 7 and November 6, and a final, December 18 at 8:00 AM. Questions will be a combination of multiple choice, short answer, and problem solving and will integrate seminar and lecture material in a thought-provoking manner. Make-up exams will only be permitted with a valid health excuse from a doctor. Sample midterm exam questions will be posted on the course website.

GRANT PROPOSAL: The remaining 20% of your grade will be based on a report, not longer than 5 single-spaced type-written pages (excluding references), which will be written in the form of a grant proposal. Any topic which is related to animal behavior can be chosen, even if it was not discussed in class. Five copies of these reports must be handed in absolutely no later than December 2.

Proposals will be evaluated by the instructors and other students in the class and then discussed during a mock panel meeting which will take place during the last two discussion periods. Your report will be distributed to three students who will each read and prepare a written evaluation of your proposal. Then, one of the reviewers will lead the panel discussion and write a written summary of that discussion. After all proposals have been reviewed, the TA and I will rank them and assign grades using the criteria that are indicated below. I intend for this to be an enjoyable learning experience that should give you a good idea of how governmental money is allocated to scientific research as well as enable you to appreciate the efforts and ideas of your classmates.

OFFICE HOURS: My office (BioPsych 2223) hours will be Wednesday 12-1 pm or by appointment. You can also schedule an appointment with me after class, by phone (301-405-6942) or by email ( If you discover that you are having difficulty with either the lecture or reading material, come see me or the TA. We are available to help, but you must take the initiative to meet with us. I can be most easily reached by email. If you have a short question, don't hestitate to send it to me by email. I will attempt to answer you within a day. As appropriate, I will post answers to questions I receive on the web page.

TA: Annie Bush will be the teaching assistant for this course. Annie's office is BioPsych 1215D and her office hours will be Friday 12-1. You can also schedule an appointment with her after class or by email ( .

ACADEMIC DISHONESTY : You should be aware that academic dishonesty is a serious offense that will not be tolerated in this course. This includes cheating on exams, fabricating information for a paper, helping another student to cheat, or plagiarizing material without adequately citing the source. This is especially important when writing your grant proposal. If you have any doubt about what constitutes plagiarism, please ask us.

Helpful Hints

HINTS FOR DISCUSSION PREPARATION : The intent of weekly discussions is to help you learn to think critically and speak confidently about animal behavior research. You should try to understand the assigned material and uncover its strengths and weaknesses by drawing on what you have learned to date. Identify the main points and critically examine the data and logic the authors use to support their conclusions. Don't be intimidated by statistical or scientific terms you do not understand. Just because this paper was published does not mean that it is necessarily correct. Your responsibility is not only to understand the research, but to evaluate its importance and quality. By the end of the term you should have a better appreciation for the kind of research currently being conducted on animal behavior and for the process scientists use to reach general conclusions, such as those described in the text.

When you are responsible for presenting the week's paper, do not merely reiterate what the paper says. Your job is to encourage and lead discussion. The best way to do this is to BRIEFLY summarize the work, and then either ask questions of the group or suggest an interpretation that will provoke disagreement. Come prepared with several questions and pose them to the class. You can choose either to work through the paper systematically, or identify the major weakness (or strength) and build discussion around this.

Each week those members of the class not presenting should come to discussion with at least one question or criticism (positive or negative) about the assigned paper. If you do not understand some aspect of the paper, ask your classmates for clarification. You can use the following set of questions to help guide you through understanding the main points, and evaluating the paper's strengths and weaknesses.

What is the paper's primary objective? Is it clearly stated and logically developed? Does the research attempt to test a specific hypothesis or is the work descriptive?

Are the methods appropriate for achieving the stated objective? Is the work observational or experimental? Are the observational techniques free from bias and/or is the experimental design free from flaws? Are the data analyzed appropriately?

What have the authors discovered? Are there any inconsistencies in the results?

What conclusions do the authors draw from the results? Is each conclusion supported by the results presented or by cited work? Where do the authors speculate, if at all, and are these speculations logical?

What is the contribution of this research to our understanding of the stated objectives and the larger field of which the research is a part? What kind of follow-up work would increase our understanding of the main questions.

SUPPLEMENTAL TEXTBOOKS : If you would like more detailed discussion of the genetic basis of behavior you should look at Behavioral Genetics: a Primer, third edition by Robert Plomin, John DeFries, Gerald McClearn and Michael Rutter (1997). If you feel you need additional help with topics relating to behavioral ecology, such as optimal foraging or ESS's, you should look at An Introduction to Behavioral Ecology, third edition, by John Krebs and Nick Davies.

GRANT PROPOSAL SUGGESTIONS: The grant proposal is a chance for you to explore in greater depth anything you have discovered or always thought was interesting about animal behavior. I want you to be creative and try to come up with a problem that either has not been treated in sufficient depth, which has been ignored, or appears to be involved in some kind of controversy. Be problem-oriented, not organism-oriented. If you are unable to pick a suitable organism for testing your ideas, come see me or the TA. We will be glad to discuss this assignment with you.

Choice of Topic

Be problem-oriented, not animal-oriented. Good research in animal behavior provides answers to general questions that apply to many animal species. At least three different styles of presentation can be successful. One particularly effective method is to focus on an area of controversy. Examples of such areas are honest vs deceptive advertisement, good genes vs nonadaptive models of sexual selection, evolution of eusociality through parental manipulation of sibling cooperation, etc. Frequently, controversy exists because conflicting theories have been proposed in the absence of supporting data. A good grant proposal reviews the theory sufficiently to identify the kind of data necessary to discriminate between competing hypotheses. An alternative approach is to reexamine a traditional idea from a new perspective. This often means challenging what is commonly viewed as conventional wisdom. As an example, Wynne-Edwards notion of group selection clearly challenged the traditional view of Darwinian natural selection and stimulated extensive thinking about levels of selection and the evolution of altruistic behavior. This particular example illustrates, though, that if you overstate your case, as Wynne-Edwards did, you lose credibility. The third approach is to extend previous studies in new directions or to a finer level of analysis. Tom Seeley's work on honeybee language, learning and communication, which built upon the classic studies of von Frisch, is a fine example of how progress can be made by continuing to pursue a single area over a long time period. Regardless of which approach you decide to adopt, use as your ultimate criterion how much you like the subject. If you can't get excited about it, you won't be able to convince anyone else to give you money.

Identify a central question around which to build your grant proposal. This is without doubt the most difficult part of this assignment. You should consider the book as a good starting place to look for potential grant proposal topics. You should also refer to the references at the end of my lecture outlines if you want to pursue a lecture topic in your grant proposal. If nothing you like easily comes to mind, go to the current periodicals room of McKeldin library, and browse through all the latest issues of the behavior journals such as Animal Behavior, Proceedings of the Royal Society of London, series B, Behaviour, Behavioral Ecology, Ethology, Behavioral Ecology and Sociobiology, Ethology and Sociobiology, Behavioral Genetics, American Naturalist, Evolution, Evolutionary Ecology and Journal of Animal Ecology or review journals such as Trends in Ecology and Evolution, Annual Review of Ecology and Systematics, Quarterly Review of Biology, or Oxford Surveys in Evolutionary Biology. Then, if you find an article that sounds interesting, read it, and read some of the references that are cited in it. You should be able to trace an idea back to its origin by just reading a handful of articles and quickly decide if the topic is suitable for a grant proposal. After you identify a topic, try to develop a central question, e.g. why do large white wading birds often form foraging groups? While much of the material in this course centers on "why" questions that inquire into the adaptive significance of behavioral traits, you should not feel inhibited from asking more mechanistic questions, e.g. how do some bats manage to fly hundreds of miles from nursery colonies to winter hibernation sites and successfully return to the exact same site where they were born? Notice that both of my questions specified particular animals even though the ideas, group foraging and migration, are very general topics in animal behavior. You must also decide on an appropriate animal group to investigate after you have decided on a question. This requires careful thought because the animal you choose dictates, to a large extent, the kind of observations or experiments that can be performed. If you decide on a question, for example why do some animals seek extra-pair copulations?, but cannot think of an appropriate organism, come see me or send me email. I am most familiar with terrestrial vertebrates and insects, but I may have access to more sources on other animal groups than you can find easily. Do not choose an organism at random. You should be able to justify both your study question and animal. Thus, you should be able to claim, without too much imagination, that this animal is better than any other for investigating the topic you have chosen.

List Alternative Hypotheses

After picking a question and organism you should attempt to ennumerate all possible alternative hypotheses which can answer your original question. Typically, one of these will be a null hypothesis which often states that the observed pattern is due to chance, rather than as a consequence of past selection. You should present these hypotheses without bias, i.e. do not state that you believe one over the others unless you have direct evidence for making such a conclusion. The purpose of your research should be to test between these hypotheses. For the group foraging question, for example, you might suggest that being in a group somehow decreases predation on individuals or alternatively, increases each bird's foraging success. Note that these particular examples are not mutually exclusive. Unfortunately, this is often the case in biology and consequently, you usually need to test both alternatives, not just one. Once you have suggested as many alternatives as you can you should devise experimental or observational tests that allow you to unambiguously reject as many of these hypotheses as possible. If you initially propose a correlative study, i.e. observations on unmanipulated free-ranging animals, you should also suggest experiments that will identify causal relationships. Such a combination of observational and experimental studies often leads to the most convincing results.


The following outline and page lengths are merely a suggestion. If your project more easily fits another format, feel free to use it. However, you should cover all of these topics in your proposal and write no more than 5, single-spaced typewritten pages. References cited, figures and tables can be included in addition to the 5 pages of text.

Purpose and Objectives: (1/2 page) This introductory section can be very brief - a single paragraph is often enough. Most people state the underlying question and then describe how they will answer it, i.e. will this be lab work, field studies, both, etc. Alternative hypotheses can be included here, but they often make more sense in the Proposed Work section after more background has been provided.

Background: (1 page) This often consists of two parts: a review of relevant theoretical and empirical studies and justification for why the animal system which you plan to use was chosen. This section should be sufficiently detailed to enable the reader to place your study in the broader context of related work and make it clear why your study is needed or will be important. If you have pilot data that relates to the proposed experiments, this is the appropriate place to include and discuss it.

Proposed Work: (2 pages) This should be a description of the experiments, observations, and analyses you propose to conduct to test your alternative hypotheses and answer your original question. Enough detail is needed to show you know what you are talking about and to convince the reader that these are practical things to do in the time period of the grant . Ideally, this should be a logical progression of experiments such that the results from one influence the next. Clearly state how each experiment or observation addresses each hypothesis. For this paper, you need not worry about time limits, but try to think about 1 to 3 year time periods. In a formal proposal you should be conscious of statistical design and include such details as sample sizes and appropriate statistical procedures you plan to use to evaluate the results of each experiment. You needn't worry about statistics for this proposal. If you propose to do several different experiments or have multiple observation periods, you should consider including a table that clearly outlines the schedule you plan to follow. The more organized you appear to be, the more likely a reviewer will believe that you can accomplish what you propose.

Potential Results: (1 page) This is an optional section in which you predict the outcome of each experiment. You should attempt to interpret the results of each experiment such that you can foresee each possible outcome. Preferably, it should be made clear that exciting results will be forthcoming no matter what result you obtain.

Impact: (1/2 page) How will your results affect the big picture? Who should care about what you discover and why? Why should someone give you money rather than contribute money to find a cure for AIDS? These are hard questions, but every scientist has to be able to justify why their research is of value. This need not require any kind of immediate benefit nor relate to improving human living conditions. The quest for knowledge is sufficient if you can provide evidence (i.e. list recent relevant references) that lots of people are also interested in the same problem.

References: You should use citations in the text, e.g. (Smith, 1996), whenever you mention the results or ideas of a previous study. I expect you to locate primary references, i.e. original research articles published in journals, rather than secondary references, i.e. summaries of studies from textbooks, review articles, or articles written for a lay audience, such as newspaper or popular magazine articles. Any articles cited in the text must be fully referenced in a literature cited section at the end of your paper. You should adopt a consistent format for these references. A good example is provided by the journal Animal Behaviour at the end of every article. You can also adopt the format I use in my lecture outlines.

Dos and Don'ts

Create subdivisions within sections to highlight topics and improve readability.
Include figures that convey information simply and dramatically
Include a flowchart to link experiments if more than three or four are planned.
Hand in five copies of your proposal.

Don'ts: Go beyond page limits.
Miss deadline for submission.
Make grammatical or typographical errors.
Write for the specialist; rather, write for the informed lay person.
Forget to summarize importance of project at the end of the proposal.


Each student will submit five copies of their proposal. Three of your classmates will read and comment on your proposal. During the last two discussion section meetings we will discuss our reviews and the class will rank the proposals and recommend the best for funding. The instructions for reviewers follows. This is very much like the process used by governmental organizations, such as the National Science Fondation, when they solicit reviews from external sources.

Please provide a frank, critical appraisal of this project proposal. Evaluate the grant on creativity in choice of subject and design of research, adequacy of experimental or observational protocol in testing hypotheses, logic and clarity of presentation, and potential impact of the research on the scientific community as well as on society. Use the scale at the bottom of the page to categorize your overall impression of the proposal. If you do not sign the proposal, your comments will be returned to the applicant anonymously.

Area Possible Points

Originality: 20
Experimental Design: 20
Feasibility: 20
Presentation: 20
Impact: 20

___ Excellent _____ Very good ____ Good ____ Fair ____ Poor