Fall 2003

Dr. Harold Gouzoules

Office: 324 Psychology - Phone: 727-7444 - E-mail: psyhg@emory.edu

Teaching Assistant: Jason Davis

Office: 112 Psychology -  E-mail: jdavis3@emory.edu

Recommended text:

• Goodenough, J., McGuire, B. & Wallace, R. (2001). PERSPECTIVES ON ANIMAL BEHAVIOR. Wiley & Sons, N.Y.

Course Synopsis

Early Directions and Approaches. Prior to publication of Darwin's Origin of Species (1859), animal behavior did not exist as a distinct discipline. Darwin's work generated a great deal of interest in the idea that human beings might be better understood if more were known about the behavior of animals. The first major issue, raised originally by Darwin and Spencer, was the evolutionary continuity of mental experience. The most significant contribution in this area came from Romanes. As a body of literature concerning animal behavior began to develop, controversies emerged (e.g., nature vs. nurture). I summarize this early work, noting that many of the areas of contention are still around (sometimes masquerading under different names), and some that were once considered dead (such as the question of animal awareness) have re-emerged.

Early Ethology. I next discuss the contributions of early ethologists such as Lorenz and Tinbergen. The ethological emphasis on the natural functions of behavior is highlighted. The concepts of innate releasing mechanisms, fixed action patterns, and sign stimuli are described, as are other major areas of early ethological study such as imprinting and drive. I end with a critique of the evolutionary approach taken by the early ethologists, focusing on "group selectionist" arguments that were common.

Neuroethology & Physiology. Modern ethology maintains an interest in the natural behavior of animals. Many of the conceptual tools that appeared so nebulous in their original forms, e.g., innate releasing mechanisms, are now better understood in physiological terms. (Feature-detector cells in the nervous system are sufficient to explain most IRMs.) I provide a detailed example of how developmental, mechanistic (physiology), functional, and evolutionary questions are compatible (in fact necessary) for complete explanations of behavior. The example is the co-evolution of the auditory systems of bats and moths, and I show how the sensory world in each species is tailored to meet demands imposed by the other species. Animals are sensitive to the relevant stimuli in their environments.

Orientation, Navigation and Rhythms. How animals orient themselves in their environments and find specific locations and destinations, sometimes over great distances, as in migration, is next discussed. While we have unraveled some of the mysteries of how animals get about, this fascinating topic is one where many questions remain unanswered.

Behavioral Genetics. A critical issue in any discussion of the evolution of behavior is whether or not behavior has a direct genetic basis. I discuss evidence from hybridization studies (crickets, love birds, dogs and bees), artificial selection (Drosophila, dogs and other domesticated animals, fish), early molecular approaches (Benzer's use of gynandromorphs and "fate mapping" to isolate the site of a mutation's effect on behavior), and current techniques (e.g., "knockouts" and gene transfer). Importantly, while the difference between two behaviors can be attributed to a single allele, this is not the same as concluding that the behavior itself is governed by a single allele.

Evolution of Learning. Learning has long been one of the most pivotal and, at the same time, challenging topics in animal behavior. A major problem has been lack of understanding of learning at the level of the nervous system. We examine different kinds of learning, differences in learning among species, and how approaches that integrate learning and genetic bases for behavior can help understand complex behavior such as song acquisition in birds.

Ecology and Behavior. Behavioral ecology attempts to understand behavior as a set of solutions to problems imposed by the environment. Problems include finding food, mates, rearing offspring, and avoiding predation. The "best" solution to a specific problem may not be possible because of constraints due to other problems that must also be dealt with. The apparent influence of food type and the distribution of food in the environment on most other aspects of an animal's existence (including sociality, mating system, predator defense) is illustrated with studies of birds and ungulates.

Natural Selection and Behavior. Sociobiological theory is discussed in these lectures. I briefly review the Darwinian theory of natural selection, illustrating with the finches of the Galapagos Islands. If animals are competing for access to limited resources such as food and water, why is it that only rarely do species go extinct as a result of exhausted resources? Lacks' 30 year study of reproductive output in a population of great tits is used to illustrate how individuals actually maximize the number of surviving offspring by not over-producing in any one year. The group selectionist arguments of the early ethologists are contrasted with current ideas about the levels at which selection takes place. If reproductive self-restraint for the good of the species is illusory, how do evolutionary biologists explain other apparently altruistic acts among animals? The problem of sterile castes in insects plagued Darwin and remained unresolved for years. Modern theoretical contributions such as inclusive fitness, kin selection, and reciprocal altruism are suggested to be partial solutions. Examples of behaviors that seem to be influenced by relatedness (such as agonistic aiding in primates) are provided. A discussion of the mechanisms of kin recognition is also included.

Aggression and Competition. The question of why animals show aggressive behavior is discussed from an evolutionary perspective. Different classes of aggression (e.g., inter-sexual, intra-sexual, parental, peer, infanticide) are illustrated and the costs and benefits associated with each class are discussed. The use of game theory as a means to understand aggressive strategies and the intensity of aggression is considered. The thrust of this mathematical approach is that it does not make sense to ask what is the "best" way for an individual to fight without considering how others in the population are behaving. The costs and benefits of territoriality are discussed using sunbirds as an example.

Sexual Selection and Mating Systems. Why sex (and not asexual reproduction)? Why the disparity between males and females in terms of the costs of reproduction? What are the consequences of the differences between males and females in reproductive potential? These questions led Darwin to the idea of sexual selection, which we consider in detail. Different mating systems are described (monogamy, polygyny, etc.) as are environmental factors that are correlated with them.

Parental Care. Forms of parental care across the animal kingdom are described. Environments that have promoted the evolution of parental care are characterized and contrasted with those in which parental care is rare or nonexistent. The degree of male participation varies from species to species and, in some cases, depends upon particular environmental conditions. Trivers' theory of "parent-offspring conflict" is considered.

Animal Communication. I begin with the problem of defining communication, then move on to a consideration of the different modalities involved and why particular modalities are more prominent in some species than in others. The relationship of environment to modality is discussed. The traditional ethological view that animal signals have their origins in behaviors that served very different functions is traced back to Darwin's principle of antithesis. Ritualization is discussed. I then talk about how animal communication has been viewed in comparison to human language. Finally, I present recent data that suggest rudimentary parallels between some of the ways animals (in particular, primates) employ vocalizations, and some features of language.

Cognitive Ethology. Until very recently, scientists have been reluctant to claim that intelligence of some sort, and perhaps even self-awareness, might guide the behavior of at least some animals. The last 15 years has seen a major change in our understanding of (and attitudes about) the animal mind. We will look at attempts to train animals in artificial languages (chimps, dolphins, sea lions) and studies that explore how animals classify and view their social and physical worlds. Armed with this new evidence on animal cognition we will consider some of the key issues in animal ethics and welfare.

Human Sociobiology. Here, we consider applications of evolutionary theory to human behavior. I begin by reviewing the different ways evolutionary theory has been used in the study of animal behavior: historical, evaluative (functional), correlational and predictive uses are illustrated with examples from the previous lectures. In human sociobiology, the predictive approach is most common. Topics such as child abuse and rape are considered. I examine the premises behind the predictions (e.g., fitness enhancement), discuss the adequacy of the operational definitions employed, and evaluate the data various investigators have presented in support of their hypotheses.

Study recommendations.

There are many names - both animal and human - discussed in the text and in the lectures. You will be expected to be familiar with the names of people who have made enduring major contributions to the study of behavior. These are scientists who have pointed the way for the field, or whose work has had a significant impact in the direction research took. Darwin, Romanes, Lorenz, and Tinbergen are good examples. I will mention many scientists' names when discussing particular studies over the course of the semester, but you will not have to associate them with their work. In exams, I will always refer to names and specific studies together (for example, "Seyfarth and Cheney's study of vervet monkey alarm calls") and you will need to recall the nature of the work. I will not expect you to know scientific names of animals since we'll be considering so many over the semester; common names will do (e.g., not "Macaca mulatta" but, instead, "rhesus monkey"). When referring to a specific study, however, you should be able to recall the species of animal by its common name (e.g., "rhesus monkey," not just "monkey"). Material for exams will come from the lectures. Suggested readings should be used to help explain or provide additional examples of points I make in class. 

The three equally-weighted exams will be composed of computer-graded questions (multiple choice, true/false, matching, etc.).  The emphasis is on specific information about animal behavior and approaches to understanding it.  Note: the final exam for the course is scheduled for Wednesday, December 17, from 12:30 - 3:00 p.m.  Make your travel plans accordingly.  Please note that the college writing requirement cannot be satisfied in this course.  Also, anyone taking the course as S/U must take the final exam in order to achieve a passing grade. There will be no extra-credit opportunities: grades will be based on performance on the exams. Grades for all exams will be posted on LearnLink.

Finally, don't hesitate to stop by my office if you have problems with any of the material, or simply wish to talk - especially about animal behavior!

Office hours: Tuesdays & Thursdays 11:00 - 12:00, or other times by appointment. (When emailing me, please use my Dooley address: psyhg@emory.edu).  Jason Davis' office hours T.B.A. in class.

Last updated: August 18, 2003