By viewing the old we learn the new Chinese Proverb Approaches to the study of animal behavior Empirical Conceptual Ethology Theoretical These descriptions represent broad categories of contributing ideas in Animal Behavior Definitions Conceptual Import ideas generated from other subdisciplines and combine them in a new cohesive way Example Hamilton s inclusive fitness model Cost Benefit Economic Model Theoretical Generation of a usually mathematical model Example Optimal foraging Pianka Empirical Experiments either manipulative or observational designed to test a concept Example Lots Remember Correlation is not equal to causation Manipulation is usually necessary to establish causation Classical Ethology Niko Tinbergen 1907 1988 Karl von Frisch 1886 1982 Konrad Lorenz 1903 1989 Ethology pre dates behavioral ecology Relying on descriptive studies ethologists discovered that many behaviors were innate INNATE COMPONENTS OF BEHAVIOR Ethology Descriptive science based on studies of animals in the natural environment Innate Behavior that has either a fixed genetic basis or a high degree of genetic preprogramming Innate behaviors may seem purposeful but animals with innate behaviors are unaware of the significance of their actions There are 2 key categories of behavior patterns identified in Ethology after Craig 1876 1954 1 Appetitive Behavior The variable actions of an animal such as its searching behavior to find food a nest site or mate 2 Consummatory Behavior Stereotypical actions that are repeated without variation such as the act of mating or killing prey Consummatory in this sense does not just refer to eating Fixed Action Patterns The most fundamental concept in classical ethology is the fixed action pattern FAP Fixed action pattern A highly stereotyped innate behavior Fixed Action Patterns can occur in response to Intraspecific Social Stimuli from conspecifics Example Courtship behaviors Feeding young Male threat displays Interspecific Stimuli from other species Example Startle response Predator avoidance Stimuli from the environment Example Migratory behavior Diurnal Nocturnal cycles Ethological studies usually begin with the creation of an ethogram of the Fixed Action Patterns Ethograms can be modified as additional information becomes available and is incorporated in the whole picture The courtship behavior of the 3 spined stickleback is an elaborate example of Fixed Action Patterns This appears to be an innate behaviour a sequence of fixed action patterns shown by all males in breeding condition each behaviour triggered by a specific external stimulus When a ripe female swollen with eggs enters his territory the male darts towards and away from the female in a so called zigzag dance 2 The female is led by the male to a nest he has constructed on the floor of the pond or stream 2 4 She may creep through the nest and spawn 5 7 The male then follows and fertilizes the eggs 8 He may chase the female away The male stays by the nest periodically fanning the nest to drive water over it and oxygenate the eggs 9 10 1 2 3 4 5 6 7 8 9 10 In ethogram form this highly stereotyped social behavior looks like this Ethogram complete inventory or descriptive catalogue of the motor patterns of a given species Ethologists asked four questions about this type of behaviour How has it evolved and developed What causes it and what is its function One way of thinking about these questions is to consider them as points on a species journey through time from the distant past into the future The evolution question How has courtship behaviour evolved in the species Is it shown by other species of sticklebacks The development question How does courtship behaviour develop during the individual s lifetime Does the male learn the zigzag dance Does he practice dancing Is the male successful in directing the female to the nest the first time he dances The cause question What causes the male to dance Are there internal factors that increase the male s tendency to dance Does increasing daylength in Spring increase the males tendency to dance Does the swollen belly of a female elicit dancing in males The function question Why does the male dance in a particular way Why does the male prod the female once she has entered the nest Why does the male fan the eggs How do these behaviours contribute to the reproductive success of the individual and ultimately the survival of the species Fruit fly fighting behavior Fruit fly fighting behavior Ethogram The researchers tallied the sequence of moves by both winners and losers into a fancy statistical chart called a Markov chain analysis left Bigger boxes show more popular tactics Smaller boxes show less common maneuvers Likewise thicker arrows show the most likely next move such as the distinct fighting loop that dominates the aggressive behavior slow approach to a wing threat to a fast approach There are no boxes or arrows for moves and sequences that occurred less often than chance In the past Ethologists generally agreed that there are distinctive characteristics of FAPs 1 FAPs are stereotyped in that they comprise sequences of motor acts which occurred in rigid predictable and highly structured sequences 2 FAPs are complex patterns a characteristic that distinguishes them from simple reflexes 3 FAPs are shown by all members of a species or at least by all members of a given sex in the appropriate age range and physiological condition 4 FAPs are elicited by simple yet highly specific stimuli More about this in just a moment 5 FAPs are regarded as self exhausting That is the mere occurrence of the FAP reduces the ease with which it can be re elicited It is not the consequence of the behavior but the mere occurrence of the behavior that makes it harder for the behavior to be elicited a second time Similar to a neural or ejaculatory refractory period 6 FAPs are regarded as triggered meaning that once elicited FAPs continue independent of external stimulation Once initiated a sequence of motor acts continues even if the environment changes so that the behavior is no longer appropriate An example of this is egg rolling behavior in geese Goose egg retrieval is an example of a Fixed Action Pattern Geese will roll an egg back into their nest by stretching out their necks Even if the egg accidentally rolls away from the goose she or he will continue the behavior in the absence of the egg Stereotyped SpeciesCommon Elicited by simple stimuli 7 The occurrence of an FAP is