Topic 2. 20th Century Animal BehaviorAnimal behavior is an exceptionally complex and multi-dimensional subject. It can be studied at any level in the biological hierarchy from sub-atomic particles to the Biosphere.In the 20th Century, 3 academic disciplines eventually merged to form the basis of how we study animal behavior today. In chronological order these influences include: Comparative Psychology, Ethology, and Behavioral Ecology.Proximate and UltimateOne of the major organizing principles in animal behavior (and biology in general) is the conceptthat questions, explanations, causes, etc. can be analyzed on at least 2 levels, and in most cases inat least 4 levels.Proximate causes are mechanistic and often are phrased using the interrogative word “How?”Ultimate causes are fitness and evolutionary in origin and can often be phrased with the interrogative word “Why?”An important point is that proximate and ultimate levels of analysis do not preclude each other orare mutually exclusive, but actually complement each other. They are also interactive as evolutionary forces (ultimate) operate on the physiology and behavior (proximate) of animals.Tinbergen’s Four QuestionsNiko Tinbergen (1907-1988) divided the proximate/ultimate dichotomy into 4 levels of analysis:1. Ontogeny (development)2. Physiology (the internal functioning of organ systems, including nerves, hormones, muscles)3. Fitness (survival and potential reproduction)4. Phylogeny (evolutionary history)Another interesting aspect of Tinbergen’s Four Questions is that they can also be placed on a timeline:1. Species past (Phylogeny)2. Individual’s past (Ontogeny)3. Current Behavior (Fitness component – survival)4. Individual’s future (Fitness component - reproduction)5. Species’ futureEthogramsA catalog or inventory of an animal species’ behavior is usually a first step for researchers interested in examining aspects of behavior. These catalogs are called Ethograms. An ethogram consists of a name for a behavior, its objective description, how often the behavior is expressed (time budget), and how often the behaviors transition from one to another.The information in an ethogram can enable an animal behavior researcher to construct a visual representative of the behaviors and transitions.Comparative PsychologyThe first major 20th Century influence on the academic discipline of animal behavior came from North American Comparative Psychologists.Their major characteristics are that they focused on bringing a small number of different species into the laboratory (out of their natural environments) and focused on how experience influences animal behavior (learning). Their major flaw was rejecting the concept of instinct, or innate behaviors, or any possible genetic basis for behaviors.EthologyThis aspect or approach to animal behavior began in Europe, principally The Netherlands, Germany, and Austria.In some ways diametrically opposed to the Comparative Psychologists, these researchers focusedon innate or instinctive behavior in natural settings. They were interested in behavior from an evolutionary perspective and relied very much on strictly observational methods or controlled studies that highlighted natural or instinctive behaviors.The ethologists introduced some still valid concepts in explaining how natural behaviors evolve and are triggered although much of the terminology has been superseded in modern animal behavior.Some of the basic ethological concepts include:Fixed Action Patterns (FAPs)Sign Stimuli or ReleasersInnate Releasing MechanismsAction Specific Energy (“motivation”)Behavioral EcologyMuch of animal behavior today is heavily influenced by an examination of how animals interact with their environment. We will spend about the last half of the semester exploring this topic.21st Century Animal BehaviorTechnological advances in the fields of Neuroscience and Molecular Biology drive much of today’s analyses of animal behavior and we will be covering some basic aspects of these influences later in the