Ethological approaches for studying animal learning

After many years of studying behaviour of animals placed in the boxes and forced to press bars and pull ropes like mad Rat Hats, some naturalists had come to understanding that in the natural world the organism has to cope with the challenges of the environment and of its own body rather than with artificial tasks fabricated by experimentalists. In 30-s, this led to the second birth of ethology, as a branch of science devoted to studying animal behaviour as it occur in natural contexts, often in natural surroundings.

I would like to clarify the above words about the second birth of ethology. The study of animal behaviour as a self-contained discipline has a tradition that stretches before the time of Darwin. The term “ethology” came out together with another one, “comparative psychology”. Both terms came out of the polarisation in French biology created by the debates between Cuvier and Geoffroy-Saint-Hilaire (see Jaynes, 1969). Flourens founded comparative psychology in 1864, and Saint-Hilaire founded ethology in 1859. Whereas comparative psychology as a term was eagerly taken up, there appearing 5 texts with it as the title in the late 1870's, ethology was less successful.

Saint-Hilaire defined ethology in the same sense as Haeckel implemented ecology, i.e. as a discipline concerning relations of organisms with each others and their environment. In his “Logic”, J.S.Mill (1843) called for a science of character, to be called “Ethology”. This went back to a tradition of 17-th Century when an actor, often a mime, who portrayed human characters, called ethologist. In 18-th Century Ethology was defined as a science of ethic. In Mill’s sense, Ethology' is the term for the study of how a person with certain characteristics acquires others under certain circumstances - the study of personality change. Since 1870, A. Giard suggested to distinguish the term “ethology” concerning habits and relations of animals, and “comparative psychology” concerning feelings, intelligence, moving and orientation. At last in 1895 Belgian paleontologist L. Dollo suggested “ethology” as a specific branch of science for studying behaviour of organisms. In early 1900-s, the pioneering work of American entomologist W. Wheeler with ant social organisation and behaviour became instrumental in establishing ethology as a significant and serious branch of modern biology. Wheeler advocated ethology as investigation of habits, instincts, intelligence and, in general, mode of life in animals.

In 1910 German biologist Oskar Heinroth, the founder of " Vergleichende Verhaltensforschung" (comparative ethology) and the scientific mentor of young Konrad Lorenz, the future founder of classical ethology, published his famous paper on the ethology of ducks. Heinroth defined ethology as investigation of innate habits, manners and rituals in animals, combining them as communicative systems. This interpretation surprisingly calls up with Mill’s treatment of the term (Durant, 1986).

In the 1930s, P. Pelseneer insisted on that ethology should be quantitative, comparative, and phylogenetic. At this time, investigations of Konrad Lorenz and Niko Tinbergen established ethology as a distinct branch of biology, which has had an effect on such wide-ranging disciplines as genetics, anthropology, and political science in addition to psychology. Ethologists believe that an animal must be studied on its own terms rather than primarily in relation to human beings, with a focus on its normal behaviour and environment. They study animal behaviour from the dual perspective of both " proximate explanations" (which concern the individual lifetime of an animal) and " ultimate explanations" (which concern an animal's phylogenetic past). Proximate explanations answer questions about how a specific behaviour occurs; ultimate explanations answer questions about why behaviour occurs.

The field of ethology is considered the study of evolution and functional significance of behaviour. This definition originates with C. O. Whitman in the 1800's. Whitman studied the display patterns of pigeons and coined the word instinct as a specific term to describe such behaviours (initially this word was used in the same sense by Condillac in 1754). Many of these instincts are triggered by various environmental stimuli. 100 years later, German biologist, Jakob von Uexkü ll, termed such triggers of instinctive stereotyped behaviours sign stimuli. Von Uexkü ll (1909) also used the German term " Umwelt" to describe the unique world that each species, even each individual, inhabits.Not only are the worlds of sight and smell vastly different from creature to creature, but as von Uexkü ll realised, our senses of time and space differ as well (Gould, 1982).

The notion of “private world” developed by von Uexkü ll's inspired Karl Von Frisch who blazed a trail for serious exploration of the various and distinct sensory worlds of animals. Von Frisch is most famous for his work on honey bees, most prominently for figuring out how the foragers communicate the location of food sources using the so called Dance Language which brought him Noble prize in 1973, 50 years after his first publication in this field. It is relatively little known that he started his investigations from the pioneering work on how bees and fishes perceive the world and developed this theme during his lifetime. His results in this field had often been contradictory to current concepts of that time. Thus, Von Frisch showed that fish could hear, and were able to see colours, despite the claims to the contrary by the famous psychologist von Hess. The results obtained on bees were also rather discussible. As von Frisch said in his Nobel lecture, many biologists thought that bees and other insects were totally colour-blind animals. But he was unable to believe it. For the bright colours of flowers can bee understood only as an adaptation to colour – sensitive visitors. This was the beginning of experiments on the colour sense of the bee.

The ethological approach had a strong Darwinian tradition underlying its development. Much of the work of early ethologists was further developed by Dutch biologist Niko Tinbergen and Austrian biologist Konrad Lorenz. They shared a Nobel Prize in 1973 with Von Frisch for their " discoveries in the field of the organisation and occurrence of individual and social behavioural patterns" in the animal world (Associate Press, 1988). This was the first such prize to be awarded to behavioural scientists and was shared by the founders of the field of ethology. This event definitely placed ethology on the solid-ground of well – accepted sciences.

Lorenz and Tinbergen were tied by lifelong friendship and allied research. They first met at a small symposium in " Instinct" organised by van der Klaauw in Leiden, 1936. Tinbergen became a second pupil of Lorenz (after Alfred Seitz) and did post-doctoral research under his tutoring, but they both learned equally as much from the other. Their collaboration was interrupted because of WWII, but soon after they were invited to the United States to lecture on their findings in animal behaviour. It is a common view that European and American Schools for studying animal behaviour have developed separately in isolation from each other. From 30-s, European founders of Ethology had known and visited explorers of behaviour in US, such as Schneirla, Yerkes, Thorpe, as well as evolutionists Ernst Mayr and David Lack. This all helped, as Tinbergen (1973) said, “in bridging the gap (so much wider than we had realised) between ethology and neuro-physiology”.

Lorenz is noted primarily for his work on genetically programmed behaviour in animals. He formulated many of the fundamental ideas in ethology, and developed the first coherent theory of instinct and innate behaviour (see: Lorenz, 1950, 1965, 1969). Tinbergen is famous for diverse studies of the natural behaviour of many animals including insects, fish, and birds (see: Tinbergen, 1942, 1951, 1963). He was a pioneer in experimental field ethology and developed a field methodology of high precision. In particular, Tinbergen manipulated the environment to explore the rule of behaviour and formulated a general research program of studying animal behaviour (see Part VII). Tinbergen argued that the scientific study of animal behaviour must begin with careful observation and description. We cannot explain why an animal acts in some fashion until we know the animal's ordinary activities. Ideally, description results in a complete inventory of the behaviour patterns of a species. This is called an ethogram. The ethogram can then be used to formulate questions about the causes of behaviour. However, Tinbergen cautioned against moving to causal issues before a complete ethogram had been obtained.

One of the most famous pupils of Lorenz, Irenaus Eible-Eibesfeldt has elaborated experiments on young mammals (mainly squirrels, polecats and hamsters) and revealed intricate interlacing of innate and learned behaviours during early lifetime in animals. His research on the behavioural ontogeny of mammals contributed significantly to the so called nature-nurture debate. Eible-Eibesfeldt demonstrated in which ways phylogenetic adaptations determine mammalian behaviour and how learning contributes to integrating behaviour into functional units. His studies on the communicative behaviour were aimed at tracing the process of " ritualization" by which signals evolve as well as at the functional aspect of how signals control social interactions. In 1970 he published “Ethology - the Biology of Behaviour", the first textbook in which the entire scope of the field was laid out. Eibl-Eibesfeldt (1971, 1979. 1989) is one of founders of Human Ethology as a distinct field of ethology (see also Eibl-Eibesfeldt and Salter, eds., 1998).

It is worth to note that techniques developed by behaviourists, such as classical conditioning, proved to be very valuable to ethologists. Animals trained to respond to a particular stimulus in the appropriate behavioural context can demonstrate their ability to discriminate between different types of stimuli. For example, von Frisch used classical conditioning to demonstrate colour vision in bees and fishes. At the same time, classical ethology has been based mainly on experiments that were conducted in natural surroundings for species investigated. Early ethologists demonstrated that even “simple” animals are capable of learning when their ecological situation requires it.

Coherent development of ethology and experimental comparative psychology resulted in modern branches of ethology, in particular, cognitive ethology and neuroethology.

Cognitive ethology is the comparative, evolutionary, and ecological study of nonhuman animal minds including thought processes, beliefs, rationality, information-processing, and consciousness (Becoff, 1995).

The modern era of cognitive ethology and its concentration on the evolution and evolutionary continuity of animal cognition is thought to have begun with the appearance of Donald R. Griffin's (1976) book “The Question of Animal Awareness: Evolutionary Continuity of Mental Experience”. Cognitive ethologists differ from comparative or cognitive psychologists by their approaches and methods but can learn important lessons from one another (Allen and Becoff, 1997; Becoff, 2002). Cognitive ethologists try to observe animals under natural conditions solving the kinds of problems for which their intelligence has become adapted by evolution (Fig. I-2). Surely they should mind the importance of control which is often may be problematic in changeable environment. Cognitive psychologists are specialised in highly controlled experimental procedures and they can broaden their horizons and learn more about the importance of more naturalistic methods using by ethologists. While it may be easier to study animals in captivity, they must provide with the complexity of social and other stimuli to which they are exposed in the field. As a rule, cognitive ethologists avoid talk of " lower" and " higher" animals and also mind that in many instances sweeping generalisations about the cognitive skills (or lack thereof) of species are based on small data sets from a limited number of individuals.

One of the main goals of cognitive ethology is to integrate the answers to questions traditionally asked in psychology laboratories with the answers to questions about ecology and evolution. This approach was scrutinised in Sara J. Shettleworth’s 1998 book “Cognition, Evolution and Behaviour”. Her own study on comparison of memory, cognitive abilities and brain processes in food-storing birds and animals may be considered one of the most effective examples of experimental investigation in the field of cognitive ethology.

To conclude this part, I would like to note that the turn of a century is the traditional time for asking questions about the human place in Nature. One of central problems concerns evolutionary routes that have led to human intelligence. We know now that although many animals, from ants to apes, sometimes behave “like humans”, they seem to use a variety of intricately interlocked specific-purpose subroutines to solve their vital problems. It is challengeable to understand the degree of flexibility of intelligent behaviour in non-human animals, and this is the main problem considered in the rest of this book.