The Fixed Action Pattern and animal communication:
			An important step, associated with the name of Konrad Lorenz though 
              probably due more to his teacher, Heinroth, was the identification 
              of fixed action patterns (FAPs). Lorenz popularized FAPs as instinctive 
              responses that would occur reliably in the presence of identifiable 
              stimuli (called sign stimuli or releasing stimuli). These FAPs could 
              then be compared across species, and the similarities and differences 
              between behaviour compared with the similarities and differences 
              in morphology on which taxonomy was based. An important and much 
              quoted study of the Anatidae (ducks and geese) by Heinroth used 
              this technique. The ethologists noted that the stimuli that released 
              FAPs were commonly features of the appearance or behaviour of other 
              members of their own species, and they were able to show how important 
              forms of animal communication could be mediated by a few simple 
              FAPs. The most sophisticated investigation of this kind was the 
              study by Karl von Frisch of the so-called “dance language” 
              underlying bee communication. Lorenz developed an interesting theory 
              of the evolution of animal communication based on his observations 
              of the nature of fixed action patterns and the circumstances in 
              which animals emit them.
 Imprinting:
			A second important finding of Lorenz concerned the early learning 
              of young nidifugous birds, a process he called imprinting. Lorenz 
              observed that the young of birds such as geese and chickens spontaneously 
              followed their mothers from almost the first day after they were 
              hatched, and he discovered that this following response could be 
              transferred to an arbitrary stimulus if the eggs were incubated 
              artificially and the stimulus was presented during a critical period 
              (now called a sensitive period) that covered the few days after 
              hatching. The concept of imprinting has been widely adopted in developmental 
              psychology.
 Tinbergen's four questions for ethologists:
			Lorenz’s collaborator, Niko Tinbergen, argued that ethology 
              always needed to pay attention to four kinds of explanation of any 
              instance of behaviour:
- function: how does the behaviour impact on the animal’s chances of survival and reproduction?
- causation: what are the stimuli that elicit the response, and how has it been modified by recent learning?
- development: how does the behaviour change with age, and what early experiences are necessary for the behaviour to be shown?
- evolutionary history: how does the behaviour compare with similar behaviour in related species, and how might it have arisen through the process of phylogeny?
The flowering of ethology:
			Through the work of Lorenz and Tinbergen, ethology developed strongly 
              in continental Europe in the years before World War II. After the 
              war, Tinbergen moved to the University of Oxford, and ethology became 
              stronger in the UK, with the additional influence of William Thorpe, 
              Robert Hinde and Patrick Bateson at the Sub-department of Animal 
              Behaviour of the University of Cambridge, located in the village 
              of Madingley. In this period, too, ethology began to develop strongly 
              in North America.
Lorenz, Tinbergen, and von Frisch were jointly awarded the Nobel Prize in 1973 for their work in developing ethology.
 Social ethology and recent developments:
			In 1970, the English ethologist John H. Crook published an important 
              paper in which he distinguished comparative ethology from social 
              ethology, and argued that much of the ethology that had existed 
              so far was really comparative ethology, looking at animals as individuals, 
              whereas in the future, ethologists would need to concentrate on 
              the behaviour of social groups of animals and the social structure 
              within them. This was prescient. E. O. Wilson's book "Sociobiology"; 
              appeared in 1975, and since that time the study of behaviour has 
              been much more concerned with social aspects. It has also been driven 
              by the stronger, but more sophisticated, Darwinism associated with 
              Wilson and Richard Dawkins. The related development of behavioral 
              ecology has also helped transform ethology. At the same time a substantial 
              rapprochement with comparative psychology has occurred, so the modern 
              scientific study of behaviour offers a more or less seamless spectrum 
              of approaches, from animal cognition, more traditional comparative 
              psychology, ethology, sociobiology and behavioural ecology.
 Notes:
			There are often mismatches between human senses and those of the 
              organisms they are observing. To compensate, ethologists often reach 
              all the way back to epistemology to give them the tools to predict 
              and avoid misinterpretation of data. 
List of ethologists:
			People who have made notable contributions to the field of ethology:
- W. C. Allee
- George Barlow
- Patrick Bateson
- John H. Crook
- Charles Darwin
- Richard Dawkins
- Vitus B. Droscher
- Dian Fossey
- Karl von Frisch
- Jane Goodall
- Temple Grandin
- Oskar Heinroth
- Robert Hinde
- Julian Huxley
- Julian Jaynes
- Konrad Lorenz
- Desmond Morris
- Ivan Pavlov
- B. F. Skinner
- William Thorpe
- Niko Tinbergen
- William Morton Wheeler
- E. O. Wilson
- Frans de Waal


 
            

