Although many naturalists have studied aspects of animal behavior through the centuries, the modern discipline of ethology is usually considered to have arisen with the work in the 1960s of Dutch biologist Nikolaas Tinbergen and Austrian biologist Konrad Lorenz, joint winners of the 1973 Nobel Prize in medicine. Ethology is a combination of laboratory and field science, with strong ties to certain other disciplines — e.g., neuroanatomy, ecology, evolution. Ethologists are typically interested in a behavioral process rather than in a particular animal group and often study one type of behavior (e.g. aggression) in a number of unrelated animals.
The desire to understand the animal world has made ethology a rapidly growing field, and since the turn of the 21st century, many prior understandings related to diverse fields such as animal communication, personal symbolic name use, animal emotions, animal culture and learning, and even sexual conduct, long thought to be well understood, have been revolutionized, as have new fields such as neuroethology.
The term "ethology" is derived from the Greek word "èthos" (ήθος), meaning "character". Other words derived from the Greek word "ethos" include "ethics" and "ethical". The term was first popularized in English by the American myrmecologist William Morton Wheeler in 1902. (An earlier, slightly different sense of the term was proposed by John Stuart Mill in his 1843 System of Logic. He recommended the development of a new science, "ethology," whose purpose would be the explanation of individual and national differences in character, on the basis of associationistic psychology. This use of the word was never adopted.)
Comparative psychology also studies animal behaviour, but, as opposed to ethology, construes its study as a branch of psychology rather than as one of biology. Historically, where comparative psychology sees the study of animal behaviour in the context of what is known about human psychology, ethology sees the study of animal behaviour in the context of what is known about animal anatomy, physiology, neurobiology, and phylogenetic history. This distinction is not representative of the current state of the field. Furthermore, early comparative psychologists concentrated on the study of learning and tended to look at behaviour in artificial situations, whereas early ethologists concentrated on behaviour in natural situations, tending to describe it as instinctive. The two approaches are complementary rather than competitive, but they do lead to different perspectives and sometimes to conflicts of opinion about matters of substance. In addition, for most of the twentieth century, comparative psychology developed most strongly in North America, while ethology was stronger in Europe, and this led to different emphases as well as somewhat differing philosophical underpinnings in the two disciplines. A practical difference is that early comparative psychologists concentrated on gaining extensive knowledge of the behaviour of very few species, while ethologists were more interested in gaining knowledge of behaviour in a wide range of species in order to be able to make principled comparisons across taxonomic groups. Ethologists have made much more use of a truly comparative method than comparative psychologists have. Despite the historical divergence, most ethologists (as opposed to behavioural ecologists), at least in North America, teach in psychology departments. It is a strong belief among scientists that the mechanisms on which behavioural processes are based are the same that rule the evolution of the living species: there is therefore a strong connection between these two fields.
Until the 18th century, the most common theory among scientists was still the Scala Naturae proposed by Aristotle: according to this theory, the living beings were classified on an ideal pyramid in which the simplest animals were occupying the lower floors, and then complexity would raise progressively until the top, which was occupied by the human beings. There was also an avant-garde group of biologists who were refusing the Aristotelian theory for a more anthropocentric one, according to which all living beings were created by God to serve mankind, and would behave accordingly. A well-radicated opinion in the common sense of the time in the Western world was that animal species were eternal and immutable, created with a specific purpose, as this seemed the only possible explanation for the incredible variety of the living beings and their surprising adaptation to their habitat.
The first biologist elaborating a complex evolution theory was Jean-Baptiste Lamarck (1744-1829). His theory was substantially made of two statements: the first is that animal organs and behaviour can change according to the way they are being used, and that those characteristics are capable of being transmitted from one generation to the next (well-known is the example of the giraffe whose neck becomes longer while trying to reach the upper leaves of a tree). The second affirmation is that each and every living organism, human beings included, tends to reach a greater level of perfection. At the time of his journey for the Galapagos Islands, Charles Darwin was well aware of Lamarck's theories and was deeply influenced by them.
Because ethology is understood as a branch of biology, ethologists have been particularly concerned with the evolution of behaviour and the understanding of behaviour in terms of the theory of natural selection. In one sense, the first modern ethologist was Charles Darwin, whose book, The Expression of the Emotions in Man and Animals, has influenced many ethologists. He pursued his interest in behaviour by encouraging his protégé George Romanes, who investigated animal learning and intelligence using an anthropomorphic method, anecdotal cognitivism, that did not gain scientific support.
Other early ethologists, such as Oskar Heinroth and Julian Huxley, instead concentrated on behaviours that can be called instinctive, or natural, in that they occur in all members of a species under specified circumstances. Their first step in studying the behaviour of a new species was to construct an ethogram (a description of the main types of natural behaviour with their frequencies of occurrence). This approach provided an objective, cumulative base of data about behaviour, which subsequent researchers could check and build upon.
An important step, associated with the name of Konrad Lorenz though probably due more to his teacher, Oskar 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 could be easily compared with the similarities and differences in morphology. 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.
Modern psychoanalysis defines instinct as an impulse which forces an individual to accomplish a task through pre-defined mental schemes, behaviours that are not caused by the usual learning process nor personal choice. In ethology, by instinct is meant a series of rigid and predictable actions and behavioural schemes which go under the term of fixed action patterns. Such schemes are only acted when a precise stimulating signal is present. When such signals act as communication among members of the same species, they go under the name of releasers. Notable examples of releasers are, in many bird species, the beak movements by the newborns, which stimulates the mother's regurgitating process to feed her offspring. Another well known case is the classic experiments by Tinbergen and Lorenz on the Graylag Goose. Like similar waterfowl, it will roll a displaced egg near its nest back to the others with its beak. The sight of the displaced egg triggers this mechanism. If the egg is taken away, the animal continues with the behaviour, pulling its head back as if an imaginary egg is still being maneuvered by the underside of its beak. However, it will also attempt to move other egg shaped objects, such as a golf ball, door knob, or even an egg too large to have possibly been laid by the goose itself (a supernormal stimulus). As made obvious by this last example, however, a behaviour only made of fixed action patterns would be particularly rigid and inefficient, reducing the probabilities of survival and reproduction. The learning process has therefore great importance, as the ability to change the individual's responses based on its experience. It can be said that the more the brain is complex and the life of the individual long, the more its behaviour will be "intelligent" (in the sense of guided by experience rather than rigid FAPs).
The learning process may take place in many ways, one of the most elementary is assuefaction. This process consists in ignoring a persistent or useless stimuli. An example of learning by assuefaction is the one observed in squirrels: when one of them feels in danger, the others hear its signal and go to the nearest repair. However, if the signal comes from an individual who has performed a big number of false alarms, his signal will be ignored.
Another common way of learning is by association, where a stimuli is, based on the experience, linked to another one which may not have anything to do with the first one. The first studies of associative learning were made by Russian physiologist Ivan Pavlov. An example of associative behaviour is observed when a common goldfish goes close to the water surface whenever a human is going to feed it, or the excitement of a dog whenever it sees a collar as a prelude for a walk. The associative learning process is linked to the necessity of developing discriminatory capacities, that is, the faculty of making meaningful choices. Being able to discriminate the members of your own species is of fundamental importance for the reproductive success. Such discrimination can be based on a number of factors in many species including birds, however, this important type of learning only takes place in a very limited period of time. This kind of learning is called 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 response could be imitated by an arbitrary stimulus if the eggs were incubated artificially and the stimulus was presented during a critical period (a less temporally constrained period is called a sensitive period) that continued for a few days after hatching.
Finally, imitation is often a big part of the learning process. A well-documented example of imitative learning is that of macaques in Hachijojima island, Japan. These primates used to live in the inland forest until the 60s, when a group of researchers started giving them some potatoes on the beach: soon they started venturing onto the beach, picking the potatoes from the sand, and cleaning and eating them. About one year later, an individual was observed bringing a potato to the sea, putting it into the water with one hand, and cleaning it with the other. Her behaviour was soon imitated by the individuals living in contact with her; when they gave birth, they taught this practice to their children.
The individual reproduction is with no doubt the most important phase in the proliferation of the species: for this reason, we can often observe complex mating rituals, which can reach a high level of complexity even if they are often regarded as fixed action patterns (FAPs). The Stickleback's complex mating ritual was studied by Niko Tinbergen and is regarded as a notable example of a FAP. Often in social life, males are fighting for the right of reproducing themselves as well as social supremacy. Such behaviours are common among mammals.
A common example of fight for social and sexual supremacy is the so-called pecking order among poultry. A pecking order is established every time a group of poultry co-lives for a certain amount of time. In each of these groups, a chicken is dominating among the others and can peck before anyone else without being pecked. A second chicken can peck all the others but the first, and so on. The chicken in the higher levels can be easily distinguished for their well-cured aspect, as opposed to the ones in the lower levels. During the period in which the pecking order is establishing, frequent and violent fights can happen, but once it is established it is only broken when other individuals are entering the group, in which case the pecking order has to be established from scratch.
Several animal species, including humans, tend to live in groups. Group size is a major aspect of their social environment. Social life is probably a complex and effective survival strategy. It may be regarded as a sort of symbiosis among individuals of the same species: a society is composed of a group of individuals belonging to the same species living within well-defined rules on food management, role assignments and reciprocal dependence.
The situation is, actually much more complex than it looks. When biologists interested in evolution theory first started examining social behaviour, some apparently unanswerable questions came up. How could, for instance, the birth of sterile casts, like in bees, be explained through an evolving mechanism which emphasizes the reproductive success of as many individuals as possible? Why, among animals living in small groups like squirrels, would an individual risk its own life to save the rest of the group? These behaviours may be examples of altruism. Of course, not all behaviours are altruistic, as shown in the table below. Notably, revengeful behaviour is claimed to have been observed exclusively in Homo sapiens.
|Type of behaviour||Effect on the donor||Effect on the receiver|
|Egoistic||Increases fitness||Decreases fitness|
|Cooperative||Increases fitness||Increases fitness|
|Altruistic||Decreases fitness||Increases fitness|
|Revengeful||Decreases fitness||Decreases fitness|
The existence of egoism through natural selection doesn't pose any question to the evolution theory and is, on the contrary, fully justified by it, as well as for the cooperative behaviour. It is much harder to understand the mechanism through which the altruistic behaviour initially developed.
Lorenz's collaborator, Niko Tinbergen, argued that ethology always needed to pay attention to four kinds of explanation in any instance of behaviour:
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 physiology or medicine in 1973 for their work in developing ethology.
Ethology is now a well recognised scientific discipline, and has a number of journals covering developments in the subject, such as the Ethology Journal.
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.
Also in 1970, Robert Ardrey's book The Social Contract: A Personal Inquiry into the Evolutionary Sources of Order and Disorder was published. The book and study investigated animal behaviour and then compared human behaviour as a similar phenomenon.
Indeed, E. O. Wilson's book Sociobiology: The New Synthesis 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 behavioural ecology has also helped transform ethology. Furthermore, 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 to more traditional comparative psychology, ethology, sociobiology and behavioural ecology. Sociobiology has more recently developed into evolutionary psychology.
People who have made notable contributions to the field of ethology (many are comparative psychologists):