Topic 8. Ethology

8.1. Ethology as one of the areas of study of the psyche of animals

Ethology (from the Greek words ethos - character, temper and logos - teaching) is a science that studies the biological foundations of animal behavior, as well as its significance in the process of ontogenesis and phylogenesis for adapting to the environment.

The subject of ethology are direct acts of external activity - complete, coordinated actions of animals, connected by some expediency. Ethologists are interested in the embodied forms of animal behavior, unlike zoopsychologists, they avoid resorting to the psyche.

Ethological research is based primarily on the observation of the behavior of animals in the natural environment (ie, in the so-called "wild nature"), as well as in the course of various experiments and experiments in the laboratory. The results of such observations make it possible to compile the so-called "ethograms". Comparison of the ethograms of animals belonging to different species allows us to get closer to understanding the evolution of their behavior. Another important problem is to identify the significance of animal behavior for the process of its adaptation to living conditions.

The first works on the study of the behavioral reactions of animals date back to the XNUMXth century, when D. White и Sh.Zh. Leroy pioneered the scientific approach to the study of animal behavior. The founder of the study of animal behavior is Charles Darwin. With his theory of natural selection, he laid the foundation for an evolutionary view of animal behavior. In addition, Darwin made numerous observations of animal behavior, proving the evolutionary unity of humans as a biological species with other animals. He first formulated the idea of ​​instinct, which was successfully used in classical ethology. Darwin's work in studying animal behavior was continued by his follower G. Romanee. His work "The Mental Faculties of Animals" (1882) was the first attempt to summarize data on comparative psychology. Romanee, however, did not always critically evaluate the facts; in particular, he attributed intelligence and feelings such as jealousy to animals. His results were refuted by the work C. Morgan "Introduction to Comparative Psychology", which later led to more careful control over the conduct of experiments and a strict evaluation of the results.

The formation of ethology as an independent science dates back to the 1930s. XNUMXth century Its occurrence is associated with the work of the Austrian scientist K. Lorenz and Dutch scientist N. Tinbergen. Together with your teacher O. Heinroth they founded the "objectivist" school. Their research was based on observations in natural conditions. Mainly higher vertebrates were studied, to a lesser extent invertebrates. The scientists of this school formulated an idea about releasers (see 2.3, p. 34), about their significance in behavioral acts. Based on these ideas, a theory of behavior was developed. Lorentz and Tinbergen paid special attention to the study of the internal mechanisms of behavioral acts, thereby establishing a connection between ethology and physiology. The studies of Lorentz and Tinbergen were prepared by the work of American scientists Whitman и Craig and German scientist O. Heinroth.

Lorentz and Tinbergen emphasized the special importance of studying the behavior of animals in natural conditions. They tried to combine functional (evolutionary) and mechanistic (causal) understanding of behavior. At the same time, Lorentz's scientific approach was distinguished by a philosophical orientation.

Along with Lorentz and Tinbergen, one of the founders of ethology as an independent science is considered a German scientist K. Frisch. His research is based on careful observations of animal behavior and is distinguished by a keen understanding of the biological functions of living organisms. The main question of Frisch's scientific research was to determine how animals obtain information about the environment. His research interests were related to the study of the behavior of honey bees and fish. Frisch's most significant contribution to the development of ethology was his work on the communication of honey bees.

In 1973, K. Lorentz, N. Tinbergen and K. Frisch were awarded the Nobel Prize in Medicine.

Modern ethologists in studying animal behavior are guided by four questions formulated by N. Tinbergen in the article "Problems and Methods of Ethology" (1963).

1. What are the reasons for an animal to perform a particular behavioral act?

2. How is the formation of a behavioral act in the process of individual development of an individual?

3. What is the significance of this behavioral act for the survival of the individual?

4. How was the evolutionary development of this behavioral act?

In general, it can be noted that ethology as a science of animal behavior involves a certain range of problems that must be solved when studying each specific behavioral act. The goal of such research should not be a simple fixation of behavioral forms, but the identification of relationships between them and events in the body and outside it. These events precede this behavioral act, accompany it or follow it.

First of all, when studying the behavior of animals, it is extremely important to carry out the so-called "causal analysis". The essence of such an analysis is to clarify the relationship between the studied behavioral reactions and the events that preceded them in time. At the same time, the temporal connections between these two successive events can be complex and diverse, only sometimes being limited to the "cause - effect" scheme.

Causal analysis of behavior is complex and always consists of several stages. The preliminary stage is to determine the place of the behavioral act in the ethological classification. Once this place has been determined, it is necessary to establish the actual connections between the conditions that preceded the behavioral act and the act itself. From such analysis, certain causal factors can be obtained. Such factors can be real environmental factors, variables that connect these factors with a specific behavioral act, or the interdependence of the behavioral acts themselves. An example is the study of display postures in birds. If these postures are combined with hitting and attacking another individual of the same species, then these behavioral acts should be classified as aggressive behavior. If a similar reaction occurs in a bird when examining its reflection in the mirror, it becomes clear that the cause of behavioral acts are certain visual stimuli that need to be identified in further research. The dependence of this behavioral reaction on a certain time of year or time of day can also be established. In this case, attention should be paid to establishing internal factors of behavior. However, at the present stage of development of science, and ethology in particular, such a descriptive study of behavioral acts is not always sufficient. The optimal analysis would be carried out at all structural levels of the body. It is necessary not only to observe behavior, but also to note the current functioning of receptors, effectors and the nervous system itself. Such opportunities are provided by the physiology of higher nervous activity, comparative psychology and other sciences that are in close contact with ethology.

Another range of problems in ethology is connected with the analysis of the causes of behavior. At the same time, attention is drawn to the ontogenetic aspect of the formation of a behavioral act, and the influence exerted on its formation by changes in the environment is noted. From these questions, the third circle of problems of ethology arises - the identification of the consequences of behavioral acts. Such consequences can manifest themselves both after a short time period and after a long period of time. Thus, immediate effects can manifest themselves through changes in the organism itself. In this case, this behavioral reaction may be repeated in the future. In addition, the effect of a behavioral act may be remote. For example, the formation of a certain behavioral response in a young animal may have a significant impact on its participation in the process of reproduction in the distant future. Thus, individuals with "incorrect" sexual imprint often cannot find a sexual partner and, consequently, "fall out" from the process of reproduction. Individual differences in behavioral responses open wide opportunities for natural selection.

8.2. Ethology at the present stage of development

In the modern sense, ethology is the science of animal behavior. All ethologists are unanimous in their views on what range of problems this science should cover. It is believed that the whole variety of ethological problems can be reduced to four main issues that were identified by N. Tinbergen. However, if there is unity among ethologists on the questions themselves, then lively discussions flare up about the specific ways of finding answers to these questions.

So, according to a number of ethologists, only those observations of the behavior of animals that were made in their natural environment, that is, in the wild, can be attributed to the subject of ethology. Other scientists recognize the right to existence of a special branch of ethology - anthropogenic ethology. This area includes observations of animals, which are carried out not in natural conditions, but in places of human activity.

The next point of view on the subject of ethology and methods of obtaining knowledge within this science is experimental zoopsychology. Its arsenal includes methods such as modeling a variety of behavioral situations that do not occur in the natural habitat of a given animal, laboratory research and experiments. In this case, the control over the obtained results and their statistical processing are very important. Adherents of the classical direction of ethology do not recognize experimental zoopsychology as part of ethology.

According to the fourth point of view, zoopsychology is a holistic science that includes ethology (observation of animals in natural conditions), experimental psychology (experiments to model various behavioral situations), and physiology (morphological and functional studies of the brain). At the same time, in no case should all these branches of zoopsychology be considered as separate, all the more so opposed to each other parts. They complement the information provided by the other industry. For example, it is very important to consider the data of ethology in conjunction with the observations obtained by physiology. This will help to trace not only the behavioral act itself, but also to identify its causes, the mechanisms that underlie it, to streamline and systematize the facts, and to make the results of observations more visual.

Ethology at the present stage of development includes many hypotheses and theories. Recently, communicative and sociobiological concepts in ethology have been intensively developed. Sociobiology as a science is often opposed to ethology itself. Supporters of such ideas believe that the range of problems of ethology includes the study of only the biological aspects of the behavioral reactions of animals, while sociobiology studies the problems of social relations of animals and behavioral ethology. In this case, ethology is exclusively theoretical, "contemplative", it is a kind of system of philosophical concepts that has an explanatory character. Sociobiology is considered as a "computational" direction associated with the analysis of behavioral reactions at the level of mechanisms, it is a more exact science than ethology. However, sociobiology cannot be opposed to ethology, because when studying a number of behavioral forms it is difficult to divide behavior into "ethological" and "sociobiological" moments.

In this regard, some authors single out the so-called "non-linguistic" hypothesis of behavior. This hypothesis is based on the idea of ​​the equivalence of the ways in which animals respond to various stimuli. In this case, the same nature of the reaction will serve as a way to establish social ties. At the same time, the concept of the formation of an equivalent class is introduced - responding to different stimuli in the same way (it is assumed that these stimuli belong to the same class. In this case, a kind of combination of sensory keys occurs, which serve for individual recognition of individuals and situations. Such equivalence of stimuli will help describe the formation of abstract representations in animals, such as sameness, symmetry, transitivity, or equivalence Abstract representations can be used by animals in a variety of social and communicative relationships, for example, in danger signals, rivalry for territory, hierarchical ties in groups, kinship or friendship interactions. at the present stage of development, suggests the possibility of the formation of abstract images in animals based on the generalization of the properties of different objects.However, reliable data on this issue are still insufficient.

In modern ethology, a comparative approach to the study of the behavioral reactions of animals is widespread. Most often, interspecies differences in forms of behavior are considered. The extensive material accumulated to date on the behavior of animals belonging to various systematic groups is being refined and processed statistically. The comparative approach makes it possible to identify such types and forms of behavior that are common to representatives of different systematic groups, to determine differences in their behavior, i.e., to identify independent behavioral variables. In addition, on the basis of a comparative analysis, hypotheses of the evolutionary formation of behavioral forms can be put forward, refined and tested.

The comparative approach also has its own characteristics, which must be taken into account when applying it. First of all, it is very difficult to isolate data on the behavior of animals at different levels of historical development. Some abilities of animals at a high level of evolutionary development may look simple in comparison with similar properties of more primitive animals. In addition, it is extremely important to pay special attention to intraspecific variability in the behavior of animals of the same species. The level of development of any form of behavior in an individual of one evolutionary level may exceed the development of the same ability in a particular individual of a higher level.

It should also be taken into account that the similarity in the behavior of animals belonging to different species may be associated with the emergence of parallel evolutionary adaptation and be based on completely different reasons. That is why, in order to conduct a deep analysis of the similarities and differences in behavioral forms, one must begin with a study of the behavioral acts of closely related species, and then move on to more distant species. In this case, generalization and comparison will serve as the main methods.

As an example of the problems of comparative ethology, we can consider the problem of establishing the hierarchical status of animals according to the degree of development of their intellectual abilities. In this case, the difficulty lies primarily in finding ways to adequately assess the capabilities of the animal's intelligence. Classifications based on an approximate assessment, without the development of special assessment methods, can be erroneous and subjective. However, a number of experimental methods have been developed for assessing the mental abilities of animals, for example, determining the level of intelligence development in solving experimentally set learning tasks. The animal is asked to solve a learning problem, while scientists determine the differences in the mental activity of animals, in the decision-making strategy. It is important to take into account the characteristics of the habitat of animals in natural conditions, and the behavioral skills that an individual possesses. At the same time, by solving additional tasks on the choice of a general rule from a set of various stimuli, it is possible to increase the accuracy of experimental conclusions by an order of magnitude. As an example of the application of this approach to assessing the intellectual abilities of animals of different species, one can cite the results of experiments on birds - crows and pigeons. As a result of the experiments, it was revealed that if pigeons memorize the solution when solving problems, then crows are able to learn the general rule of solution. Thus, according to this assessment approach, ravens are superior to pigeons in terms of intelligence.

Another problem of comparative ethology is the selection of such tasks for animals that would be adequate for many species, and, moreover, would be comparable with each other.

Modern theoretical ethology pays great attention to the problem of studying the cognitive abilities of animals. The cognitive approach allows us to interpret specific behavioral acts and contributes to the creation of new theories of behavior. Within the framework of this approach, the results of sociological, psychological, cybernetic, linguistic and philosophical studies of thinking are integrated. In general, the cognitive approach is completely developed within the framework of human psychology, but it can also be applied to the study of animal behavior, i.e. in ethology. However, in this case a number of problems arise.

An analysis of animal behavior from the standpoint of any model of the cognitive process is very difficult. Thus, it is extremely difficult to correctly prove the use of deduction or induction by animals as methods of reasoning in solving a problem. The proof of a similar method of reasoning is simpler, but the model of the cognitive process inevitably simplifies. The use of semantic and syntactic models is even more unrealistic, because they are very far from animal contact forms. The idea of ​​thinking as a manipulation of models of the external environment can be used as the basis of the cognitive approach in ethology.

The cognitive approach involves the study of the ontogenetic aspect of learning in animals. The concept of the mechanism of cognitive development is introduced. These are various mental processes that improve the ability of a developing organism to process information. Several types of such mechanisms of cognitive development have been identified. All of them are manifested in the cognitive activity of both animals and humans. According to psychologists, cognitive development is based on such neural mechanisms as associative competition, coding, analogies, and the choice of a behavior strategy. However, for animals the existence of such mechanisms has not been conclusively proven.

For ethology, the theory is of great importance, according to which a constant characteristic of any neural mechanism is the competitive interaction between the psychological and physiological processes that occur in the animal body. Such interaction allows behavior to be changeable, capable of adapting to changing environmental conditions. In addition, due to the competition of these processes in the body, there is a constant selection of the most effective mechanisms of cognitive learning in a given environment.

There are three main concepts in modern ethology, each of which has its supporters. The most popular of these is the concept of behaviorism. The theoretical basis of behaviorism is scientific positivism, while the behavior of animals within the framework of the behaviorist concept is studied using objective methods. Scientific experiments are built on the basis of scientific positivism, and explanations of behavioral acts are also built accordingly. Internal variables are introduced into explanations, with the help of which a connection is established between the reaction and the stimulus that causes it.

The second trend common in modern ethology is functionalism. Functionalism involves the study of the activity and structure of an organism from a biological as well as from a phylogenetic point of view. At the same time, it is believed that knowledge about its structure is quite enough to predict the behavior of an animal. Behavior is considered as adaptive in nature, during the life of an individual, structure and function may change.

The third concept, which is the opposite of the first two, is cognitive psychology. It studies the diverse processes of information processing, while internal processing of external information is allowed. Methods of demonstrating the structures of consciousness that cognitive psychology uses are often not accepted by ethologists, as these methods are more applicable to the study and description of human behavior.

All these trends complement each other, they do not have fundamental differences, but only affect the methodological methods of description.

The material substratum of ethology is the data of functional anatomy, physiology, endocrinology and other sections of the natural sciences. All these data are extremely important for the analysis and prediction of many forms of animal and human behavior. Ethology at the present stage of development has a neurobiological basis. The study of the nervous system is extremely important for explaining the results of observations of animals in natural or experimental conditions. There is a direct relationship between the behavior of an animal and the development of its nervous system. The higher the animal in terms of development, the more complex the ways of its interaction with the outside world and the more complex its nervous system is.

Neurobiology includes many biological disciplines: human and animal physiology and psychology, embryology, anatomy, genetics, molecular biology, cytology, biophysics and biochemistry. Neurobiology considers the issue of controlling the nervous system of all animal life processes. It includes molecular neuroscience, neurochemistry, neurogenetics and neuroembryology. All these branches of neuroscience collect information about the mechanisms and location of information storage in the nervous system, its origin and properties.

Modern ethology closely cooperates with such biological branches as the physiology of higher nervous activity, biochemistry and biophysics. These sciences supplement ethology with knowledge about the laws by which the nervous system works during the performance of behavioral acts, what patterns underlie them. Often in close collaboration with ethology and neuroscience are evolutionary morphology and anthropology. Anthropology allows us to consider the evolutionary development of the human brain, and evolutionary morphology involves the study of the evolutionary development and formation of the nervous system of animals, from protozoa to humans.

The boundaries of neurobiology are fuzzy, but it is possible to accurately determine the common material substratum of all branches of knowledge that are part of it. This substrate is the functional morphology of the nervous system. When studying any processes of the molecular, biochemical or physiological level, it is important as a structural basis to pay attention to the organization of the central and peripheral nervous system at all levels of its organization: anatomical, histological and cytological. However, one should not forget that if the structure of the nervous system in general is not considered in the study of the behavioral acts of animals, then the causes of these behavioral forms will remain unexplained. Thus, neurobiology is not only the basis of modern ethology, but also an independent subject.

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Authors: Stupina S.B., Filipechev A.O.

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