Free technical library

Zoopsychology. Lecture notes: briefly, the most important

Directory / Lecture notes, cheat sheets

Comments on the article

Table of contents

1. Zoopsychology as a science (History of zoopsychology. Subject, tasks, methods and significance of zoopsychology)
2. Instinct (The concept of instinct. Modern ideas about instinct. Instinct as the basis for the formation of animal behavior. Internal and external factors. The structure of instinctive behavior)
3. Behavior. Basic forms of animal behavior
4. Learning (The learning process. The role of cognitive processes in the formation of skills. Learning and communication. Imitation in animals)
5. The development of the mental activity of animals in ontogenesis (Development of mental activity in the prenatal period. Development of the mental activity of animals in the early postnatal period. Development of mental activity in the juvenile (play) period. Animal games)
6. General characteristics of the psyche of animals. The evolution of the psyche (General characteristics of the mental activity of animals. Levels of development of the sensory psyche. Perceptual psyche. The problem of intelligence in animals)
7. Human psyche (Evolution of the human psyche in phylogenesis. The origin of labor activity, social relations and articulate speech)
8. Ethology (Ethology as one of the areas of study of the psyche of animals. Ethology at the present stage of development)

Topic 1. Animal psychology as a science

1.1. History of zoopsychology

Animal psychology from ancient times to the creation of the first evolutionary doctrine. Currently, the science of animal behavior - zoopsychology - is experiencing a period of active development. Over the past ten years alone, a number of new magazines have appeared, as well as Internet sites devoted to the problems of zoopsychology; numerous articles reflecting the development of the main branches of this science are published in periodicals on biology and psychology.

The study of animal behavior has attracted the attention of scientists at all stages of the development of human society. The science of animal behavior was created and developed by scientists who sometimes held diametrically opposed views on the nature of the same phenomena. Probably, in the way of studying these phenomena, in their interpretation, all existing philosophical systems, as well as religious views, were reflected.

It is traditional to divide the history of zoopsychology into two periods:

1) before Charles Darwin created the theory of evolution in 1859;

2) the period after Darwin. The term “scientific zoopsychology” is often used for the latter period, thereby emphasizing that before the development of evolutionary teaching, this science did not have a serious basis and therefore could not be considered independent. Nevertheless, many prominent scientists of antiquity and the Middle Ages can rightfully be classified as zoopsychologists.

One of the main questions that occupied the minds of researchers of antiquity was the question of whether there is a difference between the complex activity of animals and the rational activity of man. It was over this issue that the first clashes of philosophical schools took place. Thus, the ancient Greek philosopher Epicurus (341-270 BC) and his followers, especially the Roman poet, philosopher and scholar Lucretius (V century BC, the main work "On the Nature of Things"), argued that the animal, like man, has a soul, but at the same time they definitely defended the position of the materiality of such a "soul". Lucretius himself repeatedly said that the expedient actions of animals are the result of a kind of natural selection, since only animals with useful properties for the body can survive in changing conditions.

In contrast to the views of materialists, ancient Greek philosophers Socrates (470-399 BC) and Plato (427-347 BC) considered the soul as a divine phenomenon, not associated with the body. According to Plato, the soul is noticeably older than the body, and the souls of man and animals are different, since the human soul has a purely mental power. Animals, on the other hand, have only the lowest form of the soul - motivation, attraction. Later, on the basis of this worldview, the first ideas about instincts were formed. Most modern zoopsychologists are inclined to think that the very idea of ​​instinct was born on the basis of an idealistic opposition between the soul of man and animal.

The first naturalist among the philosophers of antiquity can rightfully be called the ancient Greek scientist and philosopher Aristotle (485-423 BC, treatise "On the Soul"). His views on the problems of the soul in man and animals were markedly different from those of his predecessors. Aristotle ascribed to man an immortal "reasonable soul" - the embodiment of the divine spirit. According to Aristotle, only the soul animates the perishable matter (body), but only the body is capable of sensory impressions and attractions. Unlike man, endowed with reason, the ability to know and free will, animals have only a mortal "sensual soul". However, Aristotle made a reservation for mammals, believing that all animals with red blood and giving birth to live young have the same five senses as humans. Throughout life, the behavior of animals is aimed at self-preservation and procreation, but it is motivated by desires and drives, sensations of pleasure or pain. But among other animals, Aristotle believed, there are rational animals, because the mind is expressed in different animals to varying degrees. Reasonable animals, in addition to the general features of behavior inherent in animals as a whole, are capable of understanding the purpose of any of their actions.

The uniqueness of Aristotle's teachings also lies in the fact that when studying the behavior of animals, he relied on specific observations. In ants, which he studied for many years, the scientist noticed the dependence of their activity on external factors, in particular on lighting. He pointed to the ability to learn from each other in a number of mammals and birds, described cases of sound communication between animals, especially during the breeding season. In addition, Aristotle was the first to conduct experiments on living objects, trying to better understand all the subtleties of animal behavior. For example, he noticed that after the removal of the chicks from their parents, they learn to sing differently than the latter, and from this he deduced the conclusion that the ability to sing is not an acquired natural gift, but arises only in the learning process.

Aristotle first began to separate the innate and acquired components of behavior. He noted in many animals the ability for individual learning and memorization of what was learned, which he attached great importance to.

The teachings of Aristotle found continuation and further development in the teachings of the Stoics, although in some respects there are also significant differences. The Stoics, in particular the ancient Greek philosopher Chrysippus (280-206 BC), for the first time give a definition of instinct. Instinct is understood by them as an innate, purposeful attraction that directs the movements of the animal to the pleasant, useful and takes it away from the harmful and dangerous. Indicative were experiments with ducklings hatched by a hen, which, nevertheless, at the moment of danger tried to hide in the water. As other examples of instinctive behavior, Chrysippus referred to nest building and caring for offspring in birds, the construction of honeycombs in bees, the ability of a spider to weave a web. According to the Stoics, animals perform all these actions unconsciously, since they simply do not have a mind. Animals carry out instinctive actions without understanding the meaning of their activity, on the basis of purely innate knowledge. It is especially significant, according to the Stoics, that the same actions were performed by all animals of the same species in the same way.

Thus, already in the writings of ancient thinkers, the main problems of animal behavior were touched upon: issues of innate and acquired behavior, instinct and learning, as well as the role of external and internal factors in the mental activity of animals were discussed. Oddly enough, the most accurate concepts were born at the junction of two diametrically opposed areas of philosophy, such as the materialistic and idealistic understanding of the essence of mental activity. The teachings of Aristotle, Plato, Socrates and other thinkers of antiquity were ahead of their time in many ways, laying the foundation for zoopsychology as an independent science, although it was still very far from its true birth.

Animal psychology in the 18th-19th centuries. The next significant research in the field of animal psychology was made only after a thousand-year gap, when a revival of scientific creativity began during the Middle Ages, but only in the 18th century. the first attempts are being made to study animal behavior on a solid foundation of reliable facts obtained as a result of observations and experiments. It was at this time that numerous works of outstanding scientists, philosophers and naturalists appeared, which had a great influence on the further study of the mental activity of animals.

One of the first zoopsychologists can rightfully be considered a French materialist philosopher, a doctor by education J.-O. Lamettry (1709-1751), whose views later had a great influence on the scientific work of J-B. Lamarck. According to Lamettry, instincts are a set of movements performed by animals forcibly, regardless of thought and experience. Lamettry believed that instincts are primarily aimed at the survival of the species and have a strict biological fitness. He did not dwell on the study of the instinctive activity of certain types of animals, but tried to draw parallels, comparing the mental abilities of different mammals, as well as birds, fish, and insects. As a result, Lamettri came to the conclusion about the gradual increase in mental abilities from simpler creatures to more complex ones, and placed man at the top of this peculiar evolutionary ladder.

In the middle of the XVIII century. saw the release of the "Treatise on Animals" by the French philosopher and teacher E.B. Condillaca (1715-1780). In this treatise, the scientist specifically considered the question of the origin of animal instincts. Noticing the similarity of instinctive actions with actions that are performed out of habit, Condillac came to the conclusion that instincts arose from rational actions by gradually turning off consciousness. Thus, in his opinion, at the basis of any instinct lies rational activity, which, through constant exercise, became a habit, and only then turned into an instinct.

This point of view on the theory of instincts has caused heated debate. One of the ardent opponents of Condillac was a French biologist Sh.Zh. Leroy. In his work “Philosophical Letters on the Mind and the Capacity of Animals for Improvement” (1802), which was published 20 years later than Condillac’s main work, he put forward the task of studying the origin of the mind from the instinct of animals as a result of the repeated action of sensation and memory exercises. Leroy's treatise was based on many years of field research. Being a keen natural scientist, he persistently argued that the mental activity of animals and especially their instincts can be known only with comprehensive knowledge of their natural behavior and taking into account their way of life.

Simultaneously with Leroy, another great French naturalist was studying the instincts of animals. J.L. Buffon (1707-1788, "Histoire naturelle des animaux", 1855). Having based his research on fieldwork experiences, Buffon for the first time was able to correctly interpret the results of his research, avoiding anthropomorphic interpretations of behavior. Anthropomorphic scientists tried to explain the behavior of animals, endowing them with purely human qualities. According to them, animals can experience love, hate, shame, jealousy and other similar qualities. Buffon proved that this is not so, and that many of the actions of animals cannot be found adequate "human" explanations. According to the teachings of Buffon, animals, in particular mammals, with which the naturalist mainly worked, are characterized by various forms of mental activity, such as sensations and habits, but not understanding the meaning of their actions. In addition, animals, according to Buffon, are able to communicate, but their language expresses only sensory experiences. Buffon insisted on the connection between environmental influences and the internal state of the animal, seeing this as the determining factor in its behavior. He drew attention to the fact that the mental qualities of an animal, its ability to learn, play the same, if not more important, role in the survival of the species, as well as physical qualities. All Buffon's concepts, built on real facts, entered the unified system of natural science he created and became the basis of the future science of the behavior and psyche of animals. In his later treatises, Buffon argued that the complex actions of animals are the result of a combination of innate natural functions that give the animal pleasure, and habits. This concept, which was based on numerous field observations and experiments, largely anticipated the development of zoopsychology, giving food for thought to future researchers.

The further development of zoopsychology as a science is closely connected with another area of ​​biology - the theory of evolutionary teaching. The urgent task of biologists was to identify which traits are inherited in behavior, and which are formed as a result of environmental influences, which trait is universal, species, and which is individually acquired, and also what is the significance of different components of animal behavior in the process of evolution, where line between man and animal. If until that time, due to the dominance of metaphysical views in biology, the instincts of animals seemed to be in an unchanged state from the moment they arose, now, based on evolutionary theories, it was possible to explain the origin of the instincts and show their variability using specific examples.

The first evolutionary doctrine was proposed at the beginning of the XNUMXth century. French naturalist J.-B. Lamarck (1744-1829, "Philosophy of Zoology"). This doctrine was not yet a holistic, complete study and in many respects lost to the later concepts of Charles Darwin, but it was it that served as a new impetus for the further development of zoopsychology. Lamarck based his evolutionary conception on the idea of ​​the guiding action of the mental factor. He believed that the external environment affects the animal organism indirectly, by changing the behavior of the animal. As a result of this influence, new needs arise, which in turn entail changes in the structure of the body through greater exercise of some and non-exercise of other organs. Thus, according to Lamarck, any physical change is based primarily on behavior, that is, he, following E.B. Condillac defined mental activity as the basis of the very existence of an animal.

Lamarck argued that even the most complex manifestations of mental activity developed from simpler ones and should be studied precisely in a comparative evolutionary plan. Nevertheless, he was a strict materialist and denied the existence of any special spiritual principle, not related to the physical structure of the animal and not amenable to natural scientific study. All mental phenomena, according to Lamarck, are closely connected with material structures and processes, and therefore these phenomena are cognizable by experience. Lamarck attached particular importance to the connection of the psyche with the nervous system. According to many psychologists, it was Lamarck who laid the foundations of comparative psychology, comparing the structure of the nervous system of animals with the nature of their mental activity at different levels of phylogenesis.

Lamarck also gave one of the first definitions of instinct, which for a long time was considered classical: “Animal instinct is an inclination that attracts (animal - Author), caused by sensations based on the needs that arise due to their needs and compels them to perform actions without any participation of thought, without any participation of the will." [1]

Lamarck argued that the instinctive behavior of animals is changeable and closely related to the environment. According to him, instincts arose in the process of evolution as a result of long-term effects on the body of certain agents of the environment. These directed actions led to the improvement of the entire organization of the animal through the formation of useful habits, which were fixed as a result of repeated repetition. Lamarck talked about the inheritance of habits, and often even habits acquired within the same generation, since no one could yet give an exact answer how long it takes for an animal to form one instinct or another under the influence of certain exercises. But at the same time, Lamarck argued that many instincts are extremely tenacious and will be passed on from generation to generation until any cardinal change occurs in the life of the population. Lamarck saw in the instincts of animals not manifestations of some mysterious supernatural force lurking in the body, but the natural reactions of the latter to environmental influences formed in the process of evolution. At the same time, instinctive actions also have a pronounced adaptive character, since it is precisely the components of behavior that are beneficial to the body that are gradually fixed. However, the instincts themselves were considered by Lamarck as the changeable properties of the animal. Thus, Lamarck's views compare favorably with the views on instinct that are encountered to this day as the embodiment of some purely spontaneous internal forces that initially have an expedient direction of action.

Despite numerous shortcomings and errors, Lamarck's theory is a completely finished work, which later served as the basis for the largest studies of the human and animal psyche, carried out by both Lamarck's followers and his antagonists. It is difficult to overestimate the role of this great natural scientist as the founder of the materialistic study of the mental activity of animals and the development of their psyche in the process of evolution. In many ways, he was ahead of his time and laid the foundation for further study of the evolution of mental activity, continued after some time by Charles Darwin.

The development of zoopsychology and the evolutionary teachings of Charles Darwin. The development of zoopsychology as a science cannot be imagined without the concepts of evolutionary teaching developed by Charles Darwin (1809-1882). Only after the recognition of Darwin's teachings did the idea of ​​a single pattern of development in living nature, of the continuity of the organic world, become firmly established in natural science. Darwin paid special attention to the evolution of mental activity in humans and animals. Thus, for his main work “The Origin of Species” (1859), he wrote a separate chapter “Instinct”, at the same time the fundamental work “The Expression of Emotions in Man and Animals” (1872), a series of separate articles on animal behavior, was published.

Darwin used a comparison of instincts in animals and humans, trying, on the basis of this comparison, to prove the commonality of their origin. He was the first among biological scientists to separate rational actions associated with the experience of individuals from instinctive actions transmitted by inheritance. Although Darwin avoided giving a detailed definition of instincts, he nevertheless emphasized that instinct is an action that is performed without prior experience and equally by many individuals to achieve a common goal. Comparing instinct with habit, Darwin said: “It would be a great mistake to think that a significant number of instincts can arise from the habit of one generation and be hereditarily transmitted to subsequent generations.” [2]

Darwin emphasized the great role of natural selection in the formation of instincts, noting that during this process there is an accumulation of changes beneficial to the species, which continues until a new form of instinctive behavior arises. In addition, based on the study of the external manifestations of the emotional state of a person, he created the first comparative description of the instincts inherent in both animals and humans. Although the constant comparison of the feelings of man and animals from the outside looks like anthropomorphism, for Darwin it was a recognition of the commonality of the biological foundations of the behavior of animals and humans and provided an opportunity to study their evolution.

In his research, Darwin paid little attention to individual learning, since he did not recognize its essential significance for the historical process of the formation of instinctive behavior. At the same time, in his works, he often referred to the highly developed instincts of working individuals of ants and bees, which are not capable of reproduction and, consequently, of transferring accumulated experience to offspring.

In his works "The Origin of Species", "The Expression of Emotions in Man and Animals", Darwin gave a well-founded natural-scientific explanation of the expediency of animal instincts. He classified instincts in the same way that he classified animal organ systems, emphasizing that natural selection preserves beneficial changes in innate behavior and eliminates harmful ones. This is because any changes in behavior are associated with morphological changes in the nervous system and sensory organs. It is these features of the structure of the nervous system, for example, changes in the structure of the cerebral cortex, that are inherited and subject to variability along with other morphological features. The expediency of instincts, according to Darwin, is the result of natural selection.

Darwin spoke in his works about the hierarchy of instincts. He believed that in the process of evolution, certain parts of the brain responsible for instinct have lost the ability to respond to external stimulation in a monotonous, i.e., instinctive manner, and such organisms exhibit more complex forms of behavior. Instinctive actions, according to Darwin, dominate to a greater extent in animals that are on the lower rungs of the evolutionary ladder, and the development of instincts directly depends on the phylogenetic rank of the animal.

As later studies have shown, such an interpretation of Darwin is not entirely correct, and the division of mental activity into monotonous and variable components is very arbitrary, since in more complex forms of behavior, any elements of behavior appear in a complex. Accordingly, at each phylogenetic level, these elements will reach the same degree of development. But it took more than a decade to figure it out. And Darwin's teaching itself is a landmark in the development of zoopsychology: for the first time, on the basis of a huge amount of factual material, it was proved that the mental activity of animals is subject to the same natural-historical patterns as all other manifestations of their life activity.

The evolutionary teachings of Darwin were positively received by many major scientists of that time: the German biologist E. Haeckel (1834-1919), English biologist and educator T.G. Huxley (1825-1895), German physiologist, psychologist and philosopher W. Wundt (1832-1920), English philosopher and sociologist G. Spencer (1820-1903). Darwin's views on instinct as an innate form of behavior were supported by an American geneticist T.Kh. Morgan (1866-1945), D.

romens (1848-1894, "The Mind of Animals", 1888) and many other researchers who continued to develop this theory in their works.

Animal psychology in Russia. One of the major Russian evolutionists who worked on the doctrine of instinct simultaneously with Charles Darwin was a professor at Moscow University K.F. steering wheel (1814-1858). He was one of the first Russian scientists who spoke out against the notions of the supernatural nature of instinct. Roulier argued that instincts are an integral part of animal life and should be studied along with anatomy, ecology and physiology. Roulier especially emphasized the relationship of instincts with the habitat of animals, he believed that their emergence and development is closely connected with other manifestations of life, therefore, the study of instincts is impossible without a comprehensive study of all its main manifestations.

The origin of instincts and their further development, according to Roulier, were subject to a general biological pattern and were the result of material processes and the influence of the external world on the body. He believed that instinct is a specific reaction developed by living conditions to manifestations of the environment that has formed over the long history of the species. The main factors in the origin of instincts, according to Roulier, are heredity, variability and an increase in the level of organization of the animal in the historical process. Roulier also believed that the instincts of highly developed animals could change in the process of gaining new experience. He especially emphasized the variability of instincts along with the bodily qualities of animals: “Just as cattle degenerates, just as the qualities of a pointer dog, unexercised, become deaf, so the need to fly away for birds that for some reason have not flown away for a long time may be lost: domestic geese and ducks have become sedentary, while their wild relatives are constantly migrating birds. Only occasionally does a domestic drake stray from home, while his girlfriend is sitting on the eggs, begin to run wild, take off and attach itself to wild ducks; only occasionally does such a wild drake fly off in the fall with its relatives to a warm country and next spring he will again appear in the yard where he was hatched." [3]

As an example of a complex instinct that changes throughout the life of an animal, Roulier cited the flight of birds. At first, birds fly only because of the instinctive processes that they learned from their parents, and, focusing on adults in the flock, they fly away even before the onset of cold weather, but gradually, accumulating knowledge, they can already lead the birds themselves, choose the best, most calm and feeding places of flight.

It should be especially noted that Roulier tried to fill each example of the use of instinct with specific content, he never used this term unfounded, without the application of scientific evidence, which scientists of that time often sinned. He obtained this evidence during numerous field studies, as well as experiments in which he emphasized the role and interaction of environmental factors and physiological processes. It was thanks to this approach that the works of Roulier took a leading place among the works of natural scientists in the mid-XNUMXth century.

Further work on the study of instincts, which served to form zoopsychology as a science, dates back to the beginning of the XNUMXth century. It was at this time that the fundamental work of the Russian zoologist and psychologist saw the light V.A. Wagner (1849-1934, "Biological Foundations of Comparative Psychology", 1910-1913). The author, based on a huge amount of material obtained both in the field and in numerous experiments, gave a deep analysis of the problem of instinct and learning. Wagner's experiments affected both vertebrates and invertebrates, which allowed him to draw conclusions about the emergence and development of instincts in different phylogenetic groups. He came to the conclusion that the instinctive behavior of animals arose as a result of natural selection under the influence of the external environment and that instincts cannot be considered immutable. According to Wagner, instinctive activity is a developing plastic activity subject to changes under the influence of external environmental factors.

As an illustration of the variability of instinct, Wagner cited his experiments with nest-building in swallows and weaving of trapping nets in spiders. Having studied these processes in detail, the scientist came to the conclusion that, although instinctive behavior is subject to change, all instinctive actions occur within clear species-type limits, it is not the instinctive actions themselves that are stable within the species, but the radius of their variability.

In the following decades, many Russian scientists conducted research on the variability of the instinctive behavior of animals and its relationship with learning. For example, the Russian physiologist, student of I.P. Pavlova L.A. Orbeli (1882-1958) analyzed the plasticity of animal behavior depending on the degree of their maturity. Russian ornithologist A.N. Prompts (1898-1948), who studied the behavior of higher vertebrates (birds and mammals), singled out integral conditioned reflex components in their instinctive actions that are formed in the process of ontogenesis, i.e., the individual development of an individual. It is these components, according to Promptov, that determine the plasticity of instinctive behavior (for more details, see 2.1, p. 27). And the interaction of innate components of behavior with conditioned reflexes acquired on their basis during life gives species-typical features of behavior, which Promptov called the "species stereotype of behavior."

Promptov's hypothesis was supported and developed by his colleague, a Russian ornithologist E.V. Lukina. As a result of experiments with passerine birds, she proved that young females nesting for the first time in their lives build nests characteristic of their species. But this stereotype may be violated if environmental conditions are atypical. For example, the gray flycatcher, which usually builds nests in semi-hollows, behind loose bark, in the absence of shelters of this kind can build a nest on a horizontal branch, and even on the ground. Here the modification of the nest-building instinct can be traced in relation to the location of the nest. Modifications can also be observed in the replacement of nest-building material. For example, birds living in large cities can use quite unusual materials as nest-building materials: cotton wool, tram tickets, ropes, gauze.

Employees of the laboratory of the Polish zoopsychologist R.I. Voytusyaka K. Gromysh and M. Berestynskoy-Vilchek conducted research on the plasticity of the building activity of insects. The first research results were published in the 1960s. Their objects were caterpillars of the species Psyche viciella, in which the process of building a cap was studied, and of the species Autispila stachjanella, in which the plasticity of instinctive behavior was studied when making passages in leaves and cocoons. As a result of numerous experiments, scientists have discovered a huge adaptive variability of instinctive actions, especially when repairing the structures of these insects. It turned out that when repairing houses, the instinctive actions of caterpillars can differ significantly depending on changes in environmental conditions.

Promptov's studies, despite their scientific significance, did not provide an objective understanding of such a complex process as the instinctive activity of animals. Promptov was certainly right when he emphasized the importance of the fusion of innate and acquired components in all forms of behavior, but he believed that the plasticity of instinct is ensured only by individual components of a behavioral act. In fact, as Wagner noted, here we are dealing with categories of instinctive behavior that are different in size and significance. In this case, there is a change in innate components, which manifests itself in individual variability of behavior typical of the species, and modification of instinctive behavior under extreme conditions. In addition, there are also acquired and, therefore, the most diverse forms of behavior, in which different forms of learning already play a dominant role, closely intertwined with innate components of behavior. Wagner described all this in detail in his writings, while Promptov's experiments only illustrated the complexity and ambiguity of the formation and development of instinctive behavior in animals.

Another major Soviet zoopsychologist of the early XX century. was an academician A.N. Severtsov (1866-1936). In the works "Evolution and the Psyche" (1922) and "The Main Directions of the Evolutionary Process" (1925), he deeply analyzed the fundamental difference between the variability of instinctive and acquired behavior (for more details, see 2.1, p. 28).

In the 1940-1960s. Zoopsychology, along with genetics, was declared a pseudoscience in Russia: numerous laboratories were closed, scientists were subjected to mass repression. Only since the mid-1960s. its gradual revival began. It is associated primarily with the names of such major animal psychologists as N.N. Ladygina-Cots (1889-1963) and her student K.E. Fabry (1923-1990), who developed a course of lectures on zoopsychology and ethology for the Faculty of Psychology of Moscow State University. The main theme of Fabry's work is related to the study of the ontogeny of the behavior and psyche of animals, the evolution of the psyche, the mental activity of primates, and the ethological and biopsychological prerequisites of anthropogenesis. Fabry is the author of the first and still practically unsurpassed textbook on zoopsychology, which has gone through three reprints since 1976. It was thanks to K. Fabry that numerous works on zoopsychology and ethology were translated into Russian, including the classical works of K. Lorenz and N. Tinbergen, the founders of modern ethology.

In 1977, a small zoopsychology laboratory was organized on the basis of the Faculty of Psychology of Moscow State University. At present, several dissertations have been defended at the faculty on the orientation and research activities of animals, the study of the motivation for animal games, a comparative analysis of the manipulative activity of different mammalian species, and the ontogeny of the intelligence of anthropoids (great apes). Classical studies are being carried out on anthropogenesis and the evolution of the psyche of great apes and humans. Applied research is also being carried out, the beginning of which was laid by K. Fabry. This, for example, has already become a classic study of the psychology of fish, which for the first time made it possible to change the traditional attitude towards fish - the object of fishing. This study showed that fish are animals with a fairly high level of development of the perceptual psyche, and are able to subtly adapt to the conditions of fishing.

The faculty continues teaching activities, publishing textbooks and anthologies - practically the only teaching aids on zoopsychology in Russia.

1.2. Subject, tasks, methods and significance of zoopsychology

Zoopsychology is a science that studies the mental activity of animals in all its manifestations. The subject of zoopsychology can be defined as the subject of the science of the manifestations, regularities and evolution of mental reflection at the level of the animal, the origin and development in the onto- and phylogenesis of mental processes in animals and the prerequisites and prehistory of human consciousness. In addition, the subject of zoopsychology is the origin and development of mental activity in animals, and as a result of this, the prerequisites for the emergence and development of human consciousness.

The object of zoopsychology is the behavior of animals. In addition to zoopsychology, animal behavior is also studied by other sciences, such as ethology, neurophysiology, physiology of higher nervous activity, and bionics. Animal behavior is understood as a set of manifestations of the external, mainly motor activity of the animal, aimed at establishing the vital connections of the organism with the environment. A zoopsychologist studies the whole complex of manifestations of the behavior and mental activity of an animal, considering the process of mental reflection as a product of its external activity. When studying this process, he is never limited only to the behavior of the animal, trying to consider the entire mental aspect of the emergence and development of this particular type of behavior.

Considering the object of zoopsychological research - the behavior of animals, it should be pointed out that zoopsychology, in contrast to classical psychology, where there is only one object of research - a person, has a huge number of objects, the number of which is still not known for certain. Hundreds of new species of animals are described every year in the world. Each species has its own biological and, consequently, mental characteristics, therefore, in order to create a more or less complete database of zoopsychological data, it is necessary to carefully study at least one representative of the family, and at best, the genus. However, modern zoopsychologists are very far from this goal, since only a few dozen species of insects, fish, birds and mammals have been thoroughly and reliably studied, and there is only fragmentary information about the behavior of the vast majority of species. In modern psychology, the term "animal" can only be used in a comparative psychological sense, when it comes to low levels of organization of the psyche as a whole.

It is necessary to dwell in more detail on the concepts that zoopsychologists often operate on, namely: the psyche, behavior and mental activity of animals.

The psyche is born only at a certain stage in the development of the organic world and is the highest form of reflection of objective reality. The psyche is expressed in the ability of highly organized living beings to reflect the surrounding world with their state. The emergence of the psyche is directly related to the emergence of the animal form of life, since with a change in the conditions of life, there was a need for a qualitatively new reflection of objective reality. The psyche allows a living organism to correlate its activity with the components of the environment, therefore, in order to ensure normal life in changing environmental conditions, the vast majority of animals have a single center for controlling the nervous activity of the body - the brain.

The psyche of animals is closely connected with behavior, which refers to all manifestations of external (motor) activity aimed at establishing connections with the environment. Mental reflection is carried out on the basis of this activity during the animal’s influence on the surrounding world. Not only the components of the environment are reflected, but also the behavior of the animal itself, as well as changes in the environment produced as a result of this influence. In the psyche of higher vertebrates, the most complete and profound reflection of surrounding objects occurs precisely as a result of their changes under the influence of the animal’s behavioral acts. As K. Fabry wrote, “it is fair to consider the psyche as a function of the animal organism, consisting in the reflection of objects and phenomena of the surrounding world in the course and result of activity directed towards this world, i.e. behavior. External activity and its reflection, behavior and psyche constitute an inseparable organic unity and can only be conditionally dissected for scientific analysis. As I.M. Sechenov showed, the psyche is born and dies with movement and behavior." [4]

Behavior is the root cause of psychic reflection, but although the psyche is a derivative of behavior, it is it that, correcting itself, directs the external activity of the organism in the right direction of interaction with the environment. Adequately reflecting the surrounding world with the help of the psyche, the animal acquires the ability to navigate in it, to build its relationship with the components of the environment.

The unity of psyche and behavior is usually expressed by the concept of “mental activity.” Here is what K. Fabry wrote about this: “By the mental activity of animals we understand the entire complex of manifestations of behavior and psyche, a single process of mental reflection as a product of the external activity of the animal. Such an understanding of mental activity, the inextricable unity of the psyche and behavior of animals opens the way for zoopsychology to "true knowledge of their mental processes and to a fruitful study of the paths and patterns of evolution of the psyche. Therefore, taking into account the primacy of behavior in mental reflection, when discussing certain aspects of the mental activity of animals, we will proceed primarily from the analysis of their motor activity in the specific conditions of their life." [5]

It was the appearance of behavior together with the animal form of life that caused the transition from unmediated (pre-psychic) ​​external activity to mental reflection, i.e., activity mediated by the reflection of objective activity. The field of activity of the zoopsychologist is at the junction of prepsychic and psychic reflection, at the level of the first manifestations of sensitivity expressed in the most primitive organisms. Further, investigating the mental activity of animals at different stages of evolution, the zoopsychologist reaches the boundary of human consciousness. The human psyche, in comparison with the psyche of animals, is a qualitatively different category, genetically related to the psyche of animals. Although biological factors common with animals continue to play an important role in human behavior, its essence is also significantly influenced by social and labor activity, articulate speech, and some other factors that are absent in animals.

Simultaneously with the psychological study of animal behavior, its general biological foundations and regularities are being studied quite widely, especially in recent decades. The science that studies these phenomena is called ethology. Ethologists are primarily interested in the behavior of animals as a factor in their adaptation to environmental conditions in the course of individual development and in the process of evolution. In addition, ethologists are trying to identify patterns of behavior change in the course of phylogenesis and the emergence of new forms of behavior. Thus, ethologists primarily pay attention to the biological roots of behavior and its adaptive significance in the evolutionary process. Zoopsychology and ethology complement each other: the first science studies the mental aspects of animal behavior, the second - the biological ones. These two aspects cannot be separated, since the psyche of animals is a necessary component of both ontogenesis and phylogenesis, regulating the relationship of the organism with the environment.

The links between zoopsychologists and neurophysiology and the physiology of higher nervous activity are very significant. Unlike the zoopsychologist, the physiologist does not study the mental reflection itself, but the processes in the body that determine its occurrence. When studying behavior, a physiologist first of all pays attention to the functions of the nervous system, in particular the brain, his main task is to study the activity of systems and organs involved in the behavior of an animal as an integral organism.

Basic methods of zoopsychological research. A psychological analysis of an animal’s behavior is carried out by a zoopsychologist during a detailed study of the movements of an experimental animal in the process of solving specific problems. Tasks should be selected in such a way that the movements of the animal can be used to most accurately judge a specific mental quality. We must not forget about the physiological state of the animal, the conditions of the experiment, as well as any external factors that could distort the purity of the experiment. It is also necessary to use direct observations of animal behavior in natural conditions. In this case, it is important to trace the changes that occur in the behavior of the animal during certain changes in the environment, which allows us to judge both the external causes of mental activity and the adaptive functions of the latter.

When studying animal behavior, it is also extremely important to carry out quantitative assessments of both external environmental factors and animal behavior. It is also necessary to take into account the biological adequacy of the experimental conditions and the applied methodology. As a rule, a certain technique is selected for conducting experiments with one or another type of animal. Otherwise, if the experiment is carried out without taking into account the specific features of the biology of the species under study and the natural behavior in the experimental environment, the results obtained during the work may not correspond to reality.

Methods of zoopsychological research are varied, but they all boil down to setting certain tasks for animals. Most of these methods were developed at the beginning of the XNUMXth century and have since been successfully used in most zoopsychological laboratories.

labyrinth method. The main task for an animal is to find a way to a goal that is not directly perceived by it. The ultimate goal can serve as a food bait, as well as a shelter, a sexual partner. In case of a noticeable deviation from the goal, punishment of the animal can be applied. The simplest maze looks like a T-shaped corridor or tube. With the correct choice of turn, the animal receives a reward, with an incorrect turn, it is punished. More complex labyrinths are made up of various combinations of T-shaped elements and dead ends, entry into which is regarded as animal errors. The results of the animal are evaluated by the number of mistakes made by him and the speed of achieving the final goal. The labyrinth method is very popular in zoopsychological research. With its help, one can study both issues related to the ability of an animal to learn, and problems of spatial orientation, in particular, the role of skin-muscular and other forms of sensitivity, memory, the formation of sensory generalizations, and many others.

Another equally popular method of zoopsychological research is called the detour method. Here, in order to achieve the goal, the animal needs to bypass one or more obstacles. In contrast to the labyrinth method, the final goal is directly perceived by the object throughout the entire path. The assessment takes into account the speed and trajectory of the animal when bypassing the obstacle. The famous Russian zoopsychologist L.V. Krushinsky (1911-1984, "The Formation of Animal Behavior in Norm and Pathology", 1960; "Biological Foundations of Reasoning Activity", 1979; "Problems of Animal Behavior", 1993) slightly modernized this method and successfully used it in studying the ability of different animal species to extrapolation (see the following sections).

The method of differentiation training is aimed at revealing the animal's ability to distinguish between several objects or features. The correct choice is rewarded, in case of an error, punishment is applied. Gradually reducing the differences between objects, it is possible to reveal the limits of their differentiation by one or another animal. Using this method, it is possible to obtain information characterizing the features of vision in animals of the species under study. This method is used to study the processes of formation of skills, memory, ability to communicate. In the latter case, by increasing the differences between sequentially presented objects, the ability of the animal to orientate itself by certain common features of these objects is revealed.

The sample selection method is one of the varieties of the above method. The animal is invited to make a choice among various objects, focusing on a certain sample. The right choice is rewarded. The method is used to study the sensory sphere of animals.

Problem box method (problem cell). During the experiment, the animal must, with the help of various devices (levers, locks, pedals, latches, etc.), leave the closed cage or, on the contrary, penetrate into it. Sometimes locked boxes are used, inside of which there is a treat: the animal is invited to extract it by unlocking the locks. The experiment can be complicated - in this case, the locks open in a strict sequence that the animal must learn. With the help of this method, complex forms of learning and motor elements of the intellectual behavior of animals are studied. Most often it is used in studying the behavior of animals with developed grasping limbs, such as rats, monkeys, raccoons. Experiments serve primarily to reveal the higher mental abilities of animals.

In a number of experiments, the use of various types of tools by animals (especially monkeys) is being studied. For example, with the help of a stick, the animal must pull a delicacy towards itself, move an inaccessible valve, or activate some mechanism. In a number of experiments with great apes, boxes and other objects are used, from which they must build "pyramids" in order to reach a high-hanging fetus. And in this case, the analysis of the structure of the objective activity of the animal in the course of solving the problem is of the greatest importance.

In addition, in zoopsychological research, an analysis of the usual manipulation of various objects is used, which is not supported by any reward. The study of such behavior makes it possible to draw conclusions about the play behavior of animals, orienting and research activities, abilities for analysis and synthesis, and some other factors that make it possible to shed light on the early stages of human evolution.

The importance of animal psychology. Data obtained in the course of zoopsychological research are important for solving fundamental problems of psychology, in particular for identifying the roots of human psychological activity, the patterns of origin and development of his consciousness. In child psychology, animal psychological research helps to identify the biological foundations of the child’s psyche, its genetic roots. Animal psychology also makes its contribution to educational psychology, because communication between children and animals has great educational and cognitive significance. As a result of such communication, complex mental contact and interaction are established between both partners, which can be effectively used for the mental and moral education of children.

In medical practice, the study of disorders of the mental activity of animals helps to study and treat nervous and mental diseases in humans. The data of zoopsychology are also used in agriculture, fur farming, and hunting. Thanks to zoopsychological research, it becomes possible to prepare these industries for the ever-increasing human impact on the natural environment. So, in fur farming, using data on animal behavior, it is possible to reduce the stress of animals when kept in cages and pens, increase productivity, and compensate for various unfavorable conditions.

The data of zoopsychology are also necessary in anthropology, especially when solving the problem of the origin of man. The study of the behavior of higher primates, data on the higher mental functions of animals are extremely important for clarifying the biological prerequisites and foundations of anthropogenesis, as well as for studying the prehistory of mankind and the origin of labor activity, social life and articulate speech.

Topic 2. Instinct

2.1. The concept of instinct. Modern ideas about instinct

The history of the study of instinctive behavior dates back several centuries, but a clear, unambiguous and universally accepted definition of instinct has not yet been developed.

Since the time of Charles Darwin's work, instinctive behavior has been understood as that part of animal behavior that is species-typical and fixed hereditarily. One of the first attempts to give an accurate interpretation of the concept of "instinct" was the definition of the German zoologist G.E. Ziegler ("Instinct", 1914). He singled out five points on which "instinctive" behavior differs from "rational" behavior.

instinctive behavior:

▪ hereditarily fixed;

▪ no additional training is required for its development;

▪ is the same for all individuals of a given species, i.e. species-typical;

▪ optimally matches the organization of the animal, its physiology;

▪ optimally adapted to the natural living conditions of animals of a given species, often even associated with cyclical changes in living conditions.

This definition of instinct has its drawbacks, for example, it does not take into account the possible variability of instinctive behavior.

The famous Russian physiologist I.P. Pavlov (1849-1936, "Conditioned reflexes: a study of the physiological activity of the cerebral cortex", 1925), one of the founders of the reflex theory, proposed to consider the concepts of reflex and instinct identical. In this case, instinctive behavior corresponds to an unconditioned reflex. This idea significantly narrowed the concept of instinct, but it was very convenient for studying the acquired components of behavior, higher nervous activity.

Dutch scientist N. Tinbergen (1907-1988) defined instinct as “a hierarchically organized nervous mechanism which responds to certain proposed and permissive impulses (external and internal) with fully coordinated, vital and species-specific movements.” [6]

Tinbergen created a hierarchical theory of instinct, which we will get acquainted with later.

Soviet physiologist HELL. Slonim gives the following definition: “Instinct is a set of motor acts and complex forms of behavior characteristic of an animal of a given species, arising in response to stimuli from the external and internal environment of the body and occurring against the background of high excitability of the nerve centers associated with the implementation of these acts. This high excitability is the result of certain changes in the nervous and hormonal systems of the body, the result of changes in metabolism." [7]

Slonim drew attention to the fact that instincts can appear and disappear during the life of an organism. For example, the instinctive behavior associated with the sucking reflex in young mammals disappears over time, but instincts related to reproduction and nest-building activity appear. HELL. Slonim points to constancy as the main property of instinctive behavior. In his opinion, insignificant individual differences cannot refute this property, but are only fluctuations in its manifestation.

Plasticity of instinctive behavior. This question is one of the key ones in animal psychology. To understand an animal's behavioral responses, it is important to determine whether innate behavior is constant or whether it can be modified. Currently, scientists have come to the conclusion that individual instinctive actions are not inherited; only the framework within which the development of instinctive reactions occurs is genetically fixed.

The Russian biologist and psychologist made an invaluable contribution to the development of this problem. V.A. Wagner (1849-1934). In the book "Biological Foundations of Comparative Psychology" (1913-1919), he came to the conclusion that instinctive behavior develops under the influence of external influences of the environment, therefore it cannot be invariable. This is a plastic and labile system that develops under the influence of natural selection. Only species-typical frames are stable, which determine the amplitude of the variability of instinct.

Subsequently, other scientists continued to develop questions of the variability of instinctive behavior. So, L.A. Orbeli revealed the dependence of the degree of plasticity of animal behavior on maturity.

A.N. Promptov pointed out that individual conditioned reflex components acquired during life make instinctive behavior plastic. As mentioned above, Promptov introduced the concept of "species stereotype of behavior", that is, behavioral features typical of a given species. They are formed by a combination of innate species-typical instinctive reactions and conditioned reflexes acquired on their basis in ontogenesis. These representations of A.N. Promptov were illustrated by the observations of E.V. Lukina for nest-building activities (see topic 1.1, p. 16).

Promptov's ideas about the combination of innate and acquired components in the behavior of animals are very important for a correct understanding of the problem of instinctive behavior. However, according to these ideas, the instinctive actions themselves are not subject to variations, their variability is ensured only by acquired components.

At present, it is believed that instinctive behavior is subject to changes within the limits of the hereditarily fixed norm of response. These limits are species-typical; outside of them, instinctive behavior under normal conditions cannot change. At the same time, in extreme conditions that go beyond the norm of response, an important role is played by the individual experience of the animal. It allows instinctive behavior to change quite a lot. Moreover, in addition to the highly conserved innate mechanisms, there is a variable component of behavior.

A.N. Severtsov in his writings gave an analysis of the variability of the instinctive and acquired components of behavior. Severtsov showed that in mammals, adaptation to changes in the external environment is carried out in two ways: through a change in organization, that is, the structure and functions of the body, and through a change in behavior. Changes in the organization allow only slow changes in the environment to be adjusted, because they require a long period of time. Changes in behavior do not require a restructuring and functioning of the animal's body, therefore, they occur at a fairly high speed. Such changes arise due to acquired, individual forms of behavior and allow the animal to adapt to drastic changes in the environment. In this case, the greatest success will be achieved by animals that can quickly develop plastic skills, whose behavior is flexible, and whose mental abilities are quite high. With this, Severtsov connects the progressive development of the brain of vertebrates that occurs in evolution.

According to Severtsov, instinctive behavior is not sufficiently changeable; therefore, its significance in evolution is roughly comparable to changes in the structure of the animal's body. Changes in innate behavior can also allow an animal to adjust to slow environmental changes. However, the role of such changes should by no means be underestimated.

According to Severtsov, "instincts are species adaptations, useful for the species to the same extent as certain morphological characters, and just as constant."

The ability to learn, according to Severtsov, depends on the hereditary height of mental organization. Actions in this case are not instinctively fixed. And in instinctive behavior, both action and the level of mental organization are hereditarily fixed. In other words, instinctive behavior is an innate program of actions that is realized in the course of the accumulation of individual experience.

Thus, the instinctive, innate, behavior of animals is determined by a genetically fixed program of actions, which is realized in the course of acquiring individual experience. Instinctive behavior must be sufficiently unchanging and stereotyped, because it concerns vital functions for the animal. If the instinct depended on the conditions in which the development of each member of the species takes place, individual individuals would not be able to benefit from the experience of the species. The slight plasticity of instinctive behavior is designed only for extreme changes in conditions. The ability to survive in all other changing conditions of existence is provided by the acquired components of behavior, learning processes. These processes make it possible to adapt a fairly rigidly fixed innate program of behavior to specific environmental conditions. With all these changes, the hereditary program itself must remain unchanged in order to ensure the performance of vital functions.

2.2. Instinct as the basis for the formation of animal behavior

Any behavioral act is a combination of interrelated components: instinct and learning. They cannot determine the behavior of an animal separately from each other. At every moment, one component prevails, but they do not exist in their pure form. Separation of instinct and learning in behavioral responses is rather conditional, therefore, it is often difficult to implement, although each of these components has its own characteristics.

Instinctive behavior can be divided into a number of instinctive actions, or instinctive acts, which, in turn, are made up of instinctive movements (separate postures, sounds, etc.).

The instinctive component of behavior determines both the very functioning of the animal's organs and the orientation of this functioning in time and space. Thus, not only how these organs will be used, but also when and in what direction are hereditarily fixed.

Learning as a plastic component of behavior cannot change the functioning of organs, but it can affect the orientation of their functions. For example, an animal that does not have flexible fingers cannot be trained to hold a glass. It does not have morphological and functional preconditions for this, it can perform only those actions for which its organs are adapted. However, training (i.e., artificial learning) can make an animal use its limbs at a certain time in a certain way. The main thing is that the very method of using the limbs should be natural for this animal. Consequently, learning can influence the orientation of the animal's functions in time and space, but the functions themselves are determined by instinctive movements.

Thus, the life process of the organism is based on instinctive reactions, and the elements of learning are completed on their basis. Congenital reactions provide all vital functions, the metabolic process, as well as such important aspects of an animal's life as reproduction and care for offspring. The development of the mental component of animal behavior is necessary in the process of evolution in order to adapt instinctive reactions to environmental conditions, to ensure the adaptation of the animal to these conditions. Hereditary behavioral responses cannot take into account the entire variety of conditions that each member of the species will encounter. In addition, instinctive behavior includes the basic mechanisms for regulating functioning and its orientation in space and time, and the learning process complements this regulation and orientation.

2.3. Internal and external factors. Structure of instinctive behavior

Internal factors of instinctive behavior. For a long time it was believed that learning was determined by external factors, and instinctive behavior - exclusively by internal factors, and the nature of these factors was unknown. A search and clarification of the internal factors of instinctive behavior would make it possible to answer the question of what determines the motivation of behavior.

Internal factors undoubtedly influence the instinctive behavior of animals. In the middle of the XX century. the American biologist P. Whit conducted experiments with spiders, during which he studied the weaving of the web when various chemicals entered the animal's body. The desired substance was applied in the form of a drop directly onto the web or injected with a syringe into the victim. Each substance stimulated the spider to weave a web of a certain type, while the very reaction of weaving a web is hereditary in a spider. So, caffeine made spiders weave a shapeless web of randomly tangled threads, while the spider experienced a semblance of neurosis. When pervitin entered the body, the spider became very restless and did not weave the entire web. The hydrochloride caused the spider to become numb, and he did not finish the web. And lysergic acid helped increase the focus on weaving, and the spider wove the web very carefully and evenly, while its quality was superior to natural.

The internal environment of the body is constant, various regulatory processes are aimed at maintaining the physico-chemical composition of the environment. It is constantly updated, however, all its parameters are maintained at a certain level due to self-regulation, which ensures the flow of all biochemical reactions. The peculiarity of the internal processes of the animal organism is that they often proceed in the form of rhythms. In the 1930s Soviet zoopsychologist V.M. Borovsky put forward the assumption that it is the deviations of these internal rhythms of the body from the norm that are the primary motivation for behavioral reactions. Under certain conditions, the internal coherence of physiological rhythms is disturbed, and the former balance in the new conditions does not ensure the normal functioning of the organism. An internal impulse arises, aimed at restoring internal balance, i.e., a need appears. Instinctive behavior in this case will be aimed at satisfying this need.

The most important sources of internal stimuli for instinctive behavior are hormones and receptors. It is known that sex hormones and pituitary hormones stimulate a number of forms of behavior associated with reproduction - fights between males for a female and for territory, guarding the nest, mating games.

For internal motivation, rhythmic processes that occur in the central nervous system are of great importance, first of all. The rhythmic activity of its stem part in vertebrates and of the abdominal nerve structures in invertebrates ensures the orientation of behavior in time. It is known that animals have the so-called "biological clock" - autonomous oscillatory processes that regulate all the rhythms of the body's vital activity. The "biological clock" determines the fluctuations in the external activity of animal behavior, all actions that are repeated with a certain cyclicity. They, as it were, lay the foundation for the instinctive behavior of the animal, and environmental factors make their own adjustments to these rhythms. Changes may be associated with the action of various external stimuli (auditory, visual, etc.), and may also depend on the general physiological state of the animal at the moment. Most often, in the behavior of animals, circadian, or diurnal, rhythms are noted, the period of which is equal to the day.

It is interesting to note that the activity of an animal is subject to such rhythmic diurnal fluctuations even in conditions of complete isolation from all environmental factors. For example, an animal can be placed in conditions of full round-the-clock illumination and yet observe an alternation of periods of sleep and wakefulness close to natural. In addition, during the day, short-term rhythms can be noted in the behavior of animals. An example is the observations of the German ethologist W. Schleidt on turkeys. He noted that the clucking of a turkey during the day is repeated with a certain rhythm, which persists even when the bird is completely isolated and deaf.

In addition to the orientation of the animal's behavior in time, the "biological clock" orients it in space. For example, migratory birds, when orienting by the position of the sun, must at each moment of time correlate its position with the time of day. This happens when they correlate information about the position of the sun with internal circadian rhythms.

Internal factors create a state in the body that precedes the manifestation of one or another instinctive reaction. However, the start of this reaction may depend on the external environmental conditions. For example, a certain level of sex hormones and pituitary hormones stimulates various behavioral responses of an animal associated with reproduction, but the production of these hormones is timed to a certain time of the year. If an animal that lives in the temperate zone of the Northern Hemisphere is kept in short daylight conditions with the onset of spring, the activity of the glands will not appear. On the contrary, if in winter the conditions of a gradually increasing day are created for the animal, hormones will begin to be released, and sexual behavior will manifest itself in the winter season.

Internal factors ensure the readiness of the body to perform one or another instinctive movement, external stimuli may not be necessary for the manifestation of an instinctive reaction.

The German neurophysiologist E. Holst discovered several zones in the brainstem of a chicken. When these zones are affected by a weak electric current, instinctive movements arise corresponding to one or another zone. It was noted that if one and the same zone is affected for a long time, increasing the strength of irritation, one can observe a whole series of instinctive actions that will be performed in the same order as in natural conditions. For example, a chicken performed the movements that are made when a terrestrial predator approaches: it showed slight anxiety, then rose, flapped its wings, screamed, and then took off. At the same time, the irritant (predator) itself was not within its visibility. Thus, under the influence of exclusively internal factors, not only individual instinctive movements, but also entire instinctive actions can manifest themselves. However, one should not forget that under natural conditions instinctive actions are "triggered" by external factors. The approach of a terrestrial predator, which the chicken would see, would lead to the excitation of the corresponding zone of the bird's brain, which was artificially stimulated under the conditions of the experiment.

External factors of instinctive behavior. If the task of internal factors of instinctive behavior is primarily to prepare the body to perform a certain behavioral act, then external factors more often play the role of unique activators of this instinctive action.

All instinctive actions are blocked by a special system, which is called the "innate trigger". This is a certain set of neurosensory systems that ensure the confinement of behavioral instinctive acts to a situation in which such behavior will be the most biologically adequate, i.e., to the so-called "starting situation". The innate trigger mechanism reacts to certain external stimuli or their combinations; it is characterized by high selectivity. Each stimulus, signal (or their combination) will be specific to a certain instinctive reaction. The innate trigger recognizes them, analyzes them, integrates the information, and unblocks the response. At the same time, the threshold of irritability of the corresponding nerve centers decreases, and they are activated. Intrinsic motivation "finds a way out", and the instinctive reaction is carried out precisely in those conditions and in that situation when it is biologically significant. The Austrian ethologist K. Lorenz (1903-1989) called this mechanism of "unblocking" the instinctive response an innate response scheme.

Instinctive action manifests itself in response to its own set of external stimuli. These stimuli are called "key" or "sign". The external signal in this case is correlated with the key, which is ideally suited to the lock (an innate trigger). For example, during the breeding season for male birds, stimuli characteristic of females of the same species will be key, these stimuli will cause instinctive actions in males associated with courtship, mating, etc.

The key stimuli can be simple physical or chemical features, their spatial relationships (eg size correlation) or vectors.

Carriers of key stimuli can be not only other individuals, but also plants, as well as various objects of inanimate nature. The German ethologist F. Walter noted that in antelope cubs, any vertical object is the key stimulus that determines the choice of a resting place. The key stimulus performs a guiding function here.

Sign stimuli are also extremely diverse in nature: they can be visual, acoustic, chemical, etc. For example, in the sexual behavior of many insects, amphibians, and some mammals, chemicals (sex attractants, pheromones) serve as key stimuli. Sound stimuli include a variety of cries, songs specific to a certain type of animal. Visual key stimuli are called "releasers". These include various morphological features (body color features, crests, crests in birds, growths). For example, for mallard females, releasers are "mirrors" on the flight feathers of drakes. There are also specific species-specific sets of movements that can act as sign stimuli (postures of submission, threatening postures, greeting rituals, mating rituals).

The animal is able to recognize the key stimulus even at the first presentation. For example, a red spot on a gull's beak elicits a "begging" response in chicks. To explain the principle of operation of this stimulus, the analogy with a key and a lock is often used.

There are also tuning key stimuli. Their action is different from sign stimuli. These stimuli lower the irritability threshold of the nerve centers and direct key stimuli.

The existence of key stimuli and their role in the development of instinctive reactions have been proven by many observations and experiments. N. Tinbergen studied the food reaction of chicks of herring gulls and thrushes when a parent individual appeared by the method of mock-ups.

The natural reaction of a hungry gull chick to its parent is to peck at the red spot on the adult bird's (yellow) beak. Tinbergen used several layouts in his experiments. Only one model exactly repeated the appearance of the head of an adult herring gull. On the rest of the layouts, individual details were excluded, and gradually the layout became less and less like a seagull's head. The last layout was a flat red object with an oblong ledge. However, the reaction of the nestlings to this object was not only not weaker than the reaction to the first model, but even exceeded it. The reaction of the chick to the layout in the form of a thin white stick with transverse dark red stripes became even more intense. From this we can conclude that the key stimuli for the appearance of the "begging" reaction in herring gull chicks are the red color and the oblong shape.

In experiments with XNUMX-day-old thrush chicks, flat disks were used as models. If the thrush chicks were offered a circle, they reached for its upper part, where the head of the parent bird was assumed. If a small circle was attached to a large circle, the chicks began to reach for it, and when two small circles of different sizes were attached, the relative size of the circles became decisive. With a large body size, the chicks were drawn to an additional circle of large sizes, with a small one, to a smaller one. Thus, the key irritants in this case are the relative position and relative size of the layout details.

Experiments on studying key stimuli in birds were carried out by Russian ornithologists G.L. Skrebitsky and T.I. Bibikova. During the experiments, the relationship of a seagull to its eggs was studied. The researchers moved eggs from one nest to another, replacing them with eggs of other bird species, and other objects of different shapes, sizes, and colors. Seagulls willingly began to “hatch” other people’s eggs, as well as eggs of other birds, differently colored dummies made of different materials (glass, clay, etc.), and foreign objects (balls, potatoes, stones). The birds did not refuse to roll even heavy stone balls into the nest, i.e., this reaction was not determined by the weight of the “egg.” G.L. Skrebitsky wrote: “... seagulls sitting on such objects presented a very original picture, but the spectacle became especially extraordinary when a bird driven from the nest returned to it and, before sitting down, carefully adjusted multi-colored balls, pebbles or potatoes with its beak ". [8]

Birds refused to incubate objects that did not have a rounded shape, such as stones with sharp protrusions or cubes. The scientists concluded that the key stimuli for the seagull were the roundness of the object, the absence of protrusions and indentations on it.

If a seagull was offered two eggs of different sizes, it began to roll a larger one into the nest. The researchers even observed such a situation when a seagull tried to incubate a wooden model of an egg of such gigantic size that it could hardly climb onto it. In this situation, a super-optimal response takes place. The animal encounters a superstimulus that has superoptimal features of the key stimulus, and begins to react to it more strongly than normal. Thus, the key stimuli are subject to the law of summation: with an increase in the parameters of the stimulus, the instinctive reaction increases proportionally. This phenomenon may explain the increased reaction of herring gull chicks to a stick cross-striped with red stripes.

N. Tinbergen drew attention to the quantitative side of the action of sign stimuli when studying the female chasing reaction in male velvet butterflies during the breeding season. Observations have shown that the male takes off not only at the approach of individuals of his own species, but also at the sight of other flying insects, as well as small birds and even leaves falling from trees. The scientist concluded that for marigold in this situation, some visual key stimuli are of paramount importance. Chemical stimuli in this case cannot be symbolic, because the direction of flight of males is in no way connected with the direction of the wind, which means that they are not guided by smells. Tinbergen and his assistants made models of butterflies out of paper and fixed them on a thin line tied to a long fishing rod. Each series of layouts had only one characteristic external feature: color, size, a certain shape. When the rod was twitched, the model butterfly began to move, which evoked a persecution response in the male marigolds. The intensity of the reaction was recorded by observation.

The results of the experiment showed that the reaction of pursuit was caused by models of all colors, but the most active males followed black "butterflies" - the reaction to them was even more pronounced than when they saw brown models, which corresponded to the natural color of the marigold female. In this case, one should speak of an increase in the visual stimulus - a dark color.

A similar picture was obtained when comparing the intensity of the reaction to the size of the layout. The males were most active in pursuing models larger than the natural size of the female. Such a stimulus as the shape of the object's body turned out to be not so important for marigolds. Males responded to models of all shapes, with long rectangular models being the least effective. However, observations showed that this was due to a violation of the nature of the movement of such "butterflies": it became less "dancing".

Tinbergen also drew attention to another feature of the action of key stimuli, which he called the incentive adder. The scientist wrote: "...a weakly attractive white model will cause the same percentage of reactions as a black one if it is shown at a shorter distance than the black one. The effectiveness of the small white model is also noticeably enhanced if it is made to 'dance.' Thus, the insufficient effectiveness of one parameter can be compensated by the increased attractiveness of a completely different parameter... incentives are added up in a kind of “stimulation adder”, which forces the marigold to react accordingly.” [9]

In addition, Tinbergen noted that the state of the male determines which stimuli are currently included in this adder. For example, under normal conditions, the males reacted only to the color tone of the layout (dark or light), i.e., the colors themselves were not included in the adder. When feeding on mock-ups painted in bright colors, the males reacted exclusively to blue and yellow models, i.e., color became a sign stimulus.

The response to a key stimulus is not always adequate to the situation and may not lead to the desired result. Thus, Tinbergen describes a phenomenon called “misfire.” A misfire in an animal’s behavior occurs when it encounters a “super-stimulus.” An example of such a “failure” is the feeding of a cuckoo chick by songbirds. The key stimuli that cause the parent bird to feed the chick are the chick's large beak and brightly colored throat. Both of these signs in the cuckoo have a “supernormal” expression. Tinbergen writes: “It is quite possible that many songbirds not only feed the cuckoo chick, but also take pleasure in its huge and attractive mouth.” [10]

A misfire can also occur in the relationships between representatives of different classes of animals. A case is described when a cardinal bird fed insects to goldfish in a pool for several weeks. The bird reacted to the wide open mouth of the fish, which is a key stimulus for it when feeding the chicks.

In conclusion, it should be noted that the instinctive behavior of animals is most often determined not by individual factors, but by their complex. This requires a combination of external and internal factors. Pigeons, for example, feed their chicks by regurgitating "crop milk" rich in proteins. The very process of education in the goiter "milk" is stimulated by the release of the hormone prolactin (internal stimulus). However, the regurgitation reaction is caused not by the filling of the goiter, but by external stimulation from the side of the chick, which, with its weight, presses on the goiter of the parent. In winter, even with a goiter filled with food, the pigeon does not have such a reaction, because there is no external stimulation.

Structure of instinctive behavior. Back at the beginning of the 1918th century. American researcher W. Craig (“Attractions and aversions as components of instinct,” XNUMX) showed that any instinctive action consists of separate phases. Craig identified two phases, which were named: the search (preparatory) phase, or appetitive behavior, and the final phase (final act).

Craig showed that under natural conditions, animals look for those key stimuli or their combinations (starting situations) that are necessary for the implementation of a certain instinctive reaction. For example, animals are looking for food, individuals of the opposite sex during the breeding season, nesting places, etc. Craig called these search behaviors appetitive, and the state of the animal at that moment appetence. Intermediate stimuli perceived at the search phase of behavior are not a goal for the animal; they are necessary only to lead to the perception of the key stimuli of the final behavior. The final phase of instinctive behavior is the very consumption by the animal of the elements of the environment it needs, it is this phase that is directly instinctive behavior.

The final phase is hereditarily determined, species-typical, it contains the biological meaning of all instinctive action. This phase of behavior consists of a small number of movements, always performed in a clear sequence. It is stereotyped, determined by the structure of the body of the animal. In this phase, only minor individual variations in behavior are possible, which are determined genetically. The acquired components of behavior practically do not play a role in the final act, and most often they are completely absent. K. Lorentz called the final acts of behavioral reactions endogenous movements, they are species-typical, hereditary and do not require special training.

In contrast to the final act, the search phase is more variable and adaptive in relation to conditions, although it is also typical of the species. It intertwines innate and acquired forms of behavior, the individual experience of the animal. The exploratory activity of the animal is characteristic of search behavior. It is through changes in appetitive behavior that instinctive reactions can be plastic. The preparatory phase is always divided into several stages. Its end comes when the animal reaches a situation in which the next link in this chain of reactions can take place. For example, the choice of a nesting territory by a male sometimes requires only a return to the old, last year's territory, and sometimes it may require a long search and even a fight with other males. According to K. Lorenz, the search phase of a behavioral act should be referred to as goal-directed behavior. At this stage, various actions are performed, but they are all subordinated to a specific goal. The exploratory phase is very important and is for the animal the same vital necessity as the consumption in the final phase. It is appetitive behavior that is a means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the outside world.

Craig built the concept of two phases of instinctive behavior on data obtained as a result of studying the feeding behavior of animals. The predator, experiencing a feeling of hunger, begins to look for prey. However, at first he does not have information about her whereabouts and therefore his search activity is still undirected. Soon the predator sees a potential prey, from which comes the first key stimulus, such as size and color details, and its search behavior moves to the next stage, which already has a certain direction. The predator begins to specify the location, the speed of movement of the prey, while focusing on other key incentives. Then the predator pursues the prey or sneaks up on it unnoticed, after which it seizes and kills. If this is necessary, the victim is dragged to another place, where he is cut into pieces. Only after this, the behavior of the animal enters the final phase, which includes the direct consumption of prey. All actions of the animal related to the search, catching and killing of the victim are related to appetitive behavior. All of them have an instinctive basis, but to a large extent depend on the process of individual learning, the experience of the animal and the situation.

Each stage of search behavior has its own preparatory and final phases. The end of one stage is a signal for the beginning of the next, and so on. Successive stages often have several degrees of subordination, so a complex structure of animal behavior is formed. For example, search behavior may lead not to the final phase of a behavioral act, but to a combination of stimuli that stimulates the next phase of search behavior. An example is the search behavior of birds during the breeding season. The first step is to select a territory for the nest. When she is found, the next stage of search behavior begins - building a nest, then the next - courting the female, etc.

Animal behavior largely consists of cycles, which in turn consist of a series of repeated simple acts. For example, a bird busy building a nest does this according to a certain pattern. First, she goes in search of building material, then, having found it, evaluates its suitability. If the bird is satisfied with the material, it carries it to the nest, otherwise it throws it away and looks for a new one. Having flown to the nest, the bird weaves the brought materials into its structure with certain movements, forms the shape of the nest, and then flies off again in search. This cycle begins spontaneously and continues as long as the bird has a need to complete the nest. Switching to each subsequent stage of the behavioral reaction occurs upon perception of a certain external stimulus. N. Tinbergen gives an example with female hymenoptera insects - philanthus (bee wolves), who feed their larvae with honey bees. The wasp, in order to make supplies, flies to places where bees gather, where it flies randomly until it meets a suitable victim. Having noticed a flying insect, the wasp flies up to it from the leeward side and stops about 70 cm. If after this the wasp catches the scent of a bee, which will be the key stimulus for the transition to the next stage of the behavioral reaction, it will grab the bee. If a bee is deprived of its scent using ether, the wasp will not grab it. The next stage of the wasp's behavior will be to paralyze the victim with a blow from the sting. To begin this stage, a stimulus associated with touching the victim is required. If you present a wasp with a model of a bee that does not look like it to the touch, but has the same smell, the wasp will not sting such a model. Thus, as an animal goes through various stages of a behavioral reaction, the stimuli that are key for it at the moment change.

The state of appetence occurs under conditions of very high excitability of the nerve centers that coordinate certain physiological reactions. K. Lorenz introduced the concept of "specific potential (energy) of action". This potential is accumulated under the influence of a number of external (temperature, light) and internal factors (hormones) in the nerve centers. Having exceeded a certain level, the accumulated energy is released, after which the search phase of the behavioral act begins. With an increased accumulation of "specific energy of action", the final act can be carried out spontaneously, that is, in the absence of appropriate stimuli, this is the so-called "idle reaction."

To explain the neurophysiological mechanism of these phenomena, Lorentz proposed his own theory. The data of the German physiologist E. Holst served as the basis for this theory.

Holst focused his experiments on the rhythmic activity of the central nervous system. He noted that in the isolated abdominal nerve cord of an earthworm, rhythmic discharges of impulses could be observed that exactly corresponded to the contraction of the worm's segments. In further research, Holst studied the swimming mechanism of the eel. It fixed the middle segments of his body and prevented them from contracting. According to the reflex theory, in this case, the posterior segments of the body will not receive irritation, and therefore will also not be able to contract. However, in reality they begin to move after a certain period of time. If the dorsal roots of the eel's spinal cord are cut, thereby disrupting the transmission of sensory information, the eel will retain the ability to swim, and their coordination will not be impaired. Thus, the movements of the eel’s body are performed not according to the mechanism of a reflex arc (depending on external stimuli), but in accordance with the rhythmic discharges of impulses in the central nervous system. Experiments by other scientists have confirmed this. For example, it has been noted that in decerebrate (with the cerebral hemispheres removed) cats, antagonistic leg muscles may contract rhythmically, completely devoid of sensory nerves. Tadpoles and fish with one intact sensory nerve retain the ability to swim and normal coordination of movements. This means that the central nervous system is characterized by endogenous automaticity, which does not depend on external stimuli. In this case, a minimum level of afferent impulses is necessary to maintain excitation (“specific action energy”) in the corresponding nerve centers at a certain level.

The works of E. Holst and his colleagues confirmed that the level of excitation in the corresponding nerve centers affects the nature of the course of instinctive reactions. The experiments were carried out on chickens, which were irritated by the current of the brain stem structures. Depending on the localization of the irritated structure, researchers noted elementary behavioral reactions (turning the head, pecking) or complex acts of behavior (courtship). And if simple reactions always proceeded in approximately the same way, regardless of the parameters of irritation and environmental conditions, then complex behavioral reactions depended on these factors. So, with a weak current strength, a rooster pounced on a stuffed ferret, and with an increase in current, even on the researcher's hand (nonspecific stimulus).

Hydraulic model of K. Lorenz. Lorenz proposed a hypothetical model for the implementation of instinctive reactions. The scientist borrowed the general principles of its operation from hydraulics, which is why it was called the “hydraulic model.”

"Specific energy of action" is represented in this model by water, which gradually fills the reservoir (energy is accumulated) through an open tap, which indicates a continuous flow of potential energy generated during the life of the organism. Water (energy) enters the reservoir (organism) as long as the body feels the need for this form of behavior. The pressure of the liquid inside the tank is constantly increasing, creating a certain voltage in the system. The outflow of water from the tank, indicating the activity of the animal, occurs through pipes, it is prevented by a valve (central braking mechanism). The valve can open in two situations: with a large pressure of water that has accumulated in the tank, or under the influence of the weight of a load suspended from the valve. Load denotes the influence of external stimuli specific to a given behavioral act. Increasing water pressure (accumulation of specific energy of action) and the severity of the load (strength of external stimuli) summarize their effect on the valve. The stronger the stimulus, the less energy is required. And vice versa, the more energy accumulated, the less force of the external stimulus is needed for the realization of the instinctive reaction. If the energy level is very high, the valve may open without external stimulus due to water pressure. This corresponds to the "idle reaction" (according to Lorentz, "reaction in the void"). So, Lorentz described the behavior of a hungry starling, which, in the absence of any stimulus from the external environment, such as an insect, fixes it with a glance and “catches” it in the air. An inclined tray with holes located at different levels indicates different types of animal motor activity during a behavioral act. The lowest hole corresponds to motor activity with the lowest threshold, the remaining holes correspond to forms of activity with a higher threshold. If the valve is only slightly opened, water will flow out in a small amount and enter only the area of ​​​​the lower hole. If the valve opens more and the intensity of the water flow increases, it will also enter the following holes. When the reservoir is emptied ("specific energy of action" runs out), this behavioral act stops.

The Lorenz model well explains the situation when the threshold for performing an action decreases with a long non-performance (water accumulates in the reservoir to such a level that it takes a small load to open the valve), and the possibility of reactions to non-specific stimuli (accumulation of water in the reservoir to such a level, when no weight is needed to open the valve).

The hydraulic model has been repeatedly criticized because of the mechanistic construction and sketchiness. The concepts of "specific energy of action" and "key stimuli" in modern zoopsychology correspond to the concept of "specific motivation".

Hierarchical theory of instinct by I. Tinbergen. It is noted that stereotypical motor reactions are in a certain relationship. Sometimes instinctive movements appear together, and an increase in the threshold of one of them causes an increase in the threshold of the second. From this we can conclude that both of them depend on one functional “center”. There is some regularity in the sequence of manifestations of actions in complex instinctive reactions. An example would be aggressive fish collisions when dividing territory. In fish of the cichlid family, direct confrontations are preceded by a special display of intimidation. In some species of cichlids, demonstrations are short-lived, and the fish almost immediately proceed to attack. In other cichlids, collisions with wounds occur only when the males are equally strong, and demonstrations are very complex and lengthy. There are also cichlids in which there are no fights at all, and a ritualized ceremony of intimidating the enemy is performed until one of the males is exhausted and retreats. Such rituals are sequential actions, starting with the display of the sides of the body, then the dorsal fins are raised vertically, followed by tail strokes. Opponents can evaluate the strength of such a blow using the side line, which perceives the vibrations of the water. Then the opponents stand in front of each other, and in some species mutual thrusts begin with open mouths, while in others the opponents bite each other in the open mouths. The rituals continue until one opponent gets tired, in this case his color fades and he swims away. All motor reactions during a ritual demonstration are strictly stereotypical and clearly follow one another. Thus, tail strikes cannot begin before the dorsal fin rises, and mutual thrusts occur only after tail strikes.

On the basis of such facts, N. Tinbergen developed a hierarchical theory of instinct (a diagram of the organization (hierarchy) of instinct). This concept is based on the idea of ​​a hierarchy of centers that control individual behavioral responses. The concept of "center" in this case is not anatomical, but functional. Tinbergen interprets instinct as a complete hierarchical system of behavioral acts. This system responds to a specific stimulus with a well-coordinated set of actions. In this case, the change in the excitability of the centers under the influence of external and internal influences occurs in a certain order. A “block” is removed from each center, which protects this center from exhaustion. First of all, the excitability of the center of the search phase of behavior increases, and the animal enters the state of searching for stimuli. When the stimulus is found, the center that controls the implementation of the final act "discharges"; this center is at a lower level of the hierarchy. Thus, the main meaning of the scheme is that the block (inhibition) of impulses is removed in the centers in a certain sequence, which stimulates the next stage of the animal's behavior.

As an illustration, Tinbergen gives a diagram of the hierarchy of the reproductive instinct centers of the male three-spined stickleback. The higher reproductive center of the stickleback male is activated by an increase in day length, hormonal and other factors. Impulses from the higher center remove the block of appetitive behavior lying near the center. This center discharges, and the male begins to search for suitable conditions for building a nest (corresponding temperature, territory, necessary soil, vegetation, shallow water). After choosing such a territory, inhibition is removed from the subordinate centers, they are discharged, and the construction of the nest itself begins.

If another male penetrates the territory of this male, the excitability of the center of aggressive behavior increases (the block is removed from it), and an aggressive reaction begins in relation to the opponent. When the rival is expelled and the female appears, the block is removed from the center of sexual behavior, courtship of the female and mating (the final act) begin.

A contribution to the study of the hierarchical organization of instinct was made by the English zoopsychologist of the XNUMXth century. R. Hynd. Using the example of the stereotypical actions of the great tit, he showed that it is not always possible to arrange these actions in the form of a hierarchical scheme. Some actions may be characteristic of several kinds of instincts, and in some cases they will be the final acts, and in others they will only be a means to create conditions in which the final act can occur.

The hierarchy of instinctive behavior is finally formed only in an adult animal. In young individuals, isolated motor acts may appear, devoid of meaning at this age, which, at a more mature age, are integrated into a complex functional set of movements.

N. Tinbergen's scheme provides for the possibility of interaction between the "centers" of various types of behavior, for example, in a situation where one center suppresses another. Thus, if a male's hunger intensifies while courting females, he interrupts mating demonstrations and begins to search for food.

As a special case of the interaction of "centers" Tinbergen considers conflict behavior that arises in a situation of simultaneous tendency to different (often opposite) forms of behavior. At the same time, none of the forms completely suppresses the others, and the incentives for each of them are extremely strong. As an example, the scientist cites observations of male three-spined sticklebacks and different types of gulls.

In a situation where one male stickleback invades the territory of another male, the owner of the territory attacks. He pursues a stranger, and he quickly swims away. When the escaping male enters his territory, they will switch roles, now the pursuer will begin to flee. If the clash of males occurs at the border of their territories, both males will have elements of both attack and flight responses in their behavior. The closer the male is to the center of his territory, the stronger the elements of attack will be expressed in his behavior. When moving away from the center of the territory, these elements will be suppressed, and the elements of flight will intensify.

In male black-headed gulls, threatening behavior during a collision at the border of territories includes five postures. Each of them expresses a certain degree of internal conflict between opposite feelings: aggressiveness and fear.

Sometimes, in such conflict situations, animals exhibit so-called "replacement movements": there occurs, as it were, a shift in the animal's activity. For example, when a starling meets with an opponent, instead of attacking, it begins to intensively sort out its plumage with its beak. When meeting on neutral territory, male herring gulls assume a menacing posture, and then suddenly begin to clean their feathers. A similar reaction can be observed in other birds, for example, white geese in such a situation make movements, as when bathing, gray geese shake themselves off, and roosters peck grass. Activity shift responses are innate.

Another type of behavior in a conflict situation is "mosaic behavior". The animal begins to perform several actions at the same time, but does not complete any of them. For example, a seagull rises to its feet in front of an opponent, raises its wings to strike, opens its beak to peck, but freezes in this position and does not move further.

The third type of behavior in a conflict situation is "redirected reaction". The animal directs its actions not to the object that causes the reaction, but to another. For example, a thrush, at the sight of an opponent, begins to peck branches furiously. Sometimes an animal addresses the aggression of a weaker individual, for example, a gray goose attacks not its rival, but a young gosling.

Variability of the structure of an instinctive behavioral act. The structure of instinctive behavior is extremely complex. The search phase is not always a reaction to searching for any environmental agents; it can also be negative. In this case, the animal avoids certain stimuli and avoids them. In addition, certain stages of search behavior may drop out altogether, in which case this phase is shortened. Sometimes the search phase does not fully manifest itself because the final act comes too quickly. The direction of search behavior may go astray, and then an “alien” final act is possible. In some cases, the search phase takes the form of a termination phase, while the actual termination phase is also retained. Then the actions in both phases look the same, but have qualitatively different motivations. In a number of cases, the final phase is not achieved at all, then the instinctive act does not proceed to completion. In animals with a highly developed psyche, the goal of a behavioral act may be the search for stimuli itself, i.e., the intermediate stages of search behavior (complex exploratory behavior).

Instinctive behavior and communication. Communication is physical (biological) and mental (exchange of information) interaction between individuals. Communication is certainly expressed in the coordination of the actions of animals, therefore it is closely related to group behavior. When communicating, animals necessarily have special forms of behavior that perform the functions of transmitting information between individuals. In this case, some animal actions acquire signaling significance. Communication in this understanding is absent in lower invertebrates, and in higher invertebrates it appears only in rudimentary form. It is inherent in all representatives of vertebrate animals to one degree or another.

The German ethologist G. Tembrok studied the process of communication in animals and its evolution. According to Tembroke, it is possible to talk about real animal communities in which individuals communicate with each other only when they begin to live together. When living together, several individuals remain independent, but together they carry out homogeneous forms of behavior in different areas. Sometimes such joint activity involves the division of functions between individuals.

The basis of communication is communication (information exchange). To do this, animals have a system of species-typical signals that are adequately perceived by all members of the community. This ability to perceive information and to transmit it must be genetically fixed. The actions by which the transmission is carried out and the assimilation of information takes place are hereditarily fixed and instinctive.

Forms of communication. According to the mechanism of action, all forms of communication differ in the channels of information transmission. There are optical, acoustic, chemical, tactile and other forms.

Among the optical forms of communication, the most important place is occupied by expressive postures and body movements that make up "demonstrative behavior." This behavior consists of showing the animal certain parts of its body, which, as a rule, carry specific signals. These can be brightly colored areas, additional structures such as combs, decorating feathers, etc. At the same time, some parts of the animal's body can visually increase in volume, for example, due to ruffled feathers or hair. The signal function can also be performed by special movements of the body or its individual parts. By performing these movements, the animal can demonstrate colored areas of the body. Sometimes such demonstrations are carried out with exaggerated intensity.

In the evolution of behavior, special motor acts appear, which are separated from other forms of behavior by the fact that they have lost their primary function and acquired a purely signal value. An example is the claw movement of a fiddling crab, which it performs when courting a female. Such movements are called "allochthonous". Allochthonous movements are species-typical and stereotypical, their function is the transmission of information. Their other name is ritualized movements. All ritualized movements are conditional. They are very rigidly and clearly fixed genetically, they are typical instinctive movements. It is this conservatism of movements that ensures the correct perception of signals by all individuals, regardless of living conditions.

Most often and in the largest number of ritualized movements are observed in the field of reproduction (first of all, these are mating games) and struggle. They transmit information to one individual about the internal state of another individual, about its physical and mental qualities.

Instinctive movements and taxis. Taxis are innate, hereditarily determined reactions to certain components of the environment.

By their nature, taxis are similar to instinctive movements, but they also have a difference. Instinctive movements always arise in response to key stimuli, while taxises are manifested under the action of directing key stimuli. This special group of stimuli is in itself not capable of causing the beginning or end of any instinctive movement. Directing key stimuli stimulate only a change in the direction of this reaction. Thus the taxises produce a general orientation of instinctive movements. Taxis are closely related to innate motor coordinations and, together with them, constitute instinctive reactions or their chains.

In addition to taxis, there are kinesis. With kinesis, there is no orientation of the animal's body relative to the stimulus. In this case, stimuli either cause a change in the speed of movement of the animal, or a change in the frequency of body turns. In this case, the position of the animal relative to the stimulus changes, but the orientation of its body remains the same.

With taxises, the body of the animal takes a certain direction. Taxis can be combined with movement, in which case the animal will move towards or away from the stimulus. If the motor activity is directed towards favorable environmental conditions for the animal, the taxis will be positive (the animal's activity is directed towards the stimulus). If, on the contrary, the conditions are not valuable for the animal or are dangerous, taxis will be negative (the animal's activity is directed away from the stimulus).

Depending on the nature of external stimuli, taxises are divided into phototaxis (light), chemotaxis (chemical stimuli), thermotaxis (temperature gradients), geotaxis (gravity), rheotaxis (liquid flow), anemotaxis (air flow), hydrotaxis (environmental humidity) and others

There are several types of taxis (according to G.S. Frenkel and D.L. Gunn; Fraenkel GS, Gunn DL "The Orientation of Animals", 1940).

1. Clinotaxis. In clinotaxis, for the orientation of the body relative to the stimulus, the ability of the receptor to determine the direction of the source of irritation is not necessary. The animal compares the intensity of stimulation from different angles by simply turning the organs bearing the receptors. An example is the establishment of the trajectory of movement towards the light of a fly larva. The photoreceptors of the larva are located at the front end of the body; when crawling, it deflects its head first to one side, then to the other. Comparison of the intensity of stimuli on both sides determines the direction of her motor reaction. This type of taxis is characteristic of primitive animals that do not have eyes.

2. Tropotaxis. The animal compares two simultaneously acting stimuli. A change in the direction of movement in this case occurs at different intensities of stimulation. An example of tropotaxis is the orientation of aquatic animals when swimming with the dorsal side up.

3. Telotaxis. The animal chooses one of the two sources of irritation and moves towards it. An intermediate direction is never selected. Thus, the influence of one of the stimuli is suppressed. For example, bees from two light sources choose one, to which they move.

4. Menotaxis ("light compass reaction"). The animal is oriented at a certain angle to the direction of the source of irritation. For example, ants, returning to the anthill, are partially guided by the position of the sun.

5. A. Kühn ("Die Orientierung der Tiere im Raum", 1919) distinguishes, in addition, mnemotaxis. In this case, the animal is guided by the configuration of stimuli, their relative position. An example is the orientation of hymenoptera when returning to a burrow. Observations by N. Tinbergen and V. Kruyt showed that philantine wasps (bee wolves), when returning to the burrow, react to the position of the entrance to it relative to the surrounding elements of the terrain.

The degree of complexity of taxises and their functions depend on the level of evolutionary development of animals. Taxis are present in all forms of behavior: from the simplest instinctive reactions to complex forms of behavior. For example, when nestlings of songbirds are oriented in relation to the parent, the key stimulus will be the very appearance of the object (adult bird), the guiding key stimulus is the relative position of the object’s parts, and taxis is the spatial orientation of the chicks towards this stimulus.

As mentioned above, for the gull chick, the key stimulus is the red color of the beak of an approaching object, which elicits a "begging" feeding response. The guiding key stimulus will be the location of the red spot on the beak, this stimulus will direct the feeding reaction of the chick. The very orientation of the nestling towards the beak of the object will be a positive phototaxis.

K. Lorenz and N. Tinbergen carried out joint studies of the relationship between instinctive motor coordination and taxis. They studied the reaction of rolling eggs into the nest in the greylag goose. For this bird, the key stimulus is the sight of a rounded object without protrusions on the surface, which is located outside the nest. This stimulus causes the geese to roll in. This innate reaction is a repeated movement of the beak towards the bird's chest, which will stop after the object being rolled in touches it. If a cylinder was placed in front of the goose, she immediately rolled it into the nest. However, when an egg was presented to her, which rolled in different directions, the female began to make additional lateral movements of the head, which gave the movement of the egg the correct direction to the nest. The guiding stimulus for taxis lateral movements of the head is the type of deviation of the egg. Thus, in higher animals, taxises orient the instinctive movements of both individual parts of the body and organs of the body.

Taxis are observed in the animal's behavioral acts both at the stage of the final act and in search behavior. In the search phase, taxises are supplemented by various orienting reactions, thanks to which the body continuously receives information about the parameters and changes in all components of the environment.

Topic 3. Behavior

3.1. Basic forms of animal behavior

When studying unconditioned reflexes and instincts, it became necessary to create a classification of the main forms of animal behavior. The first attempts at such a classification were made back in the pre-Darwinian period, but they reached their greatest development at the beginning of the XNUMXth century. So, I.P. Pavlov divided the innate elements of behavior into indicative, defensive, nutritional, sexual, parental and childish. With the appearance of new data on the conditioned reflex activity of animals, it became possible to create more detailed classifications. For example, orienting reflexes began to be subdivided into actual orienting and exploratory reflexes, an orienting reflex aimed at searching for food was called an orienting reflex, etc.

Another classification of forms of behavior was proposed by A.D. Slonim in 1949 in the article "On the relationship of unconditioned and conditioned reflexes in mammals in phylogenesis". In his scheme, three main groups of reflexes were distinguished:

1) reflexes aimed at preserving the internal environment of the body and the constancy of matter. This group includes eating behavior, which ensures the constancy of the substance, and homeostatic reflexes, which ensure the constancy of the internal environment;

2) reflexes aimed at changing the external environment of the body. These include defensive behavior and environmental, or situational, reflexes;

3) reflexes associated with the preservation of the species. These include sexual and parental behavior.

Later, scientists of the Pavlov school developed other classifications of unconditioned reflexes and the conditioned reflexes formed on their basis. For example, the classifications of D.A. Biryukov, created in 1948 by N.A. Rozhansky (1957). These classifications were quite complex, they included both the actual reflexes of behavior and the reflexes of the regulation of individual physiological processes, and therefore did not find wide application.

R. Hynd gave several classifications of types of behavior based on certain criteria. The scientist believed that there are a lot of such criteria, and in practice, criteria are most often chosen that are suitable for the particular problem that is being considered. He mentioned three main types of criteria by which classification is carried out.

1. Classification for immediate reasons. According to this classification, the types of activity determined by the same causal factors are combined into one group. For example, all types of activity are combined, the intensity of which depends on the action of the male sex hormone (sexual behavior of the male), types of activity associated with stimuli "male-rival" (agonistic behavior), etc. This type of classification is necessary to study the behavior of the animal, it is convenient to apply it in practice.

2. The functional classification is based on the evolutionary classification of activities. Here, the categories are smaller, for example, such types of behavior as courtship, migration, hunting, and threat are distinguished. Such a classification is justified as long as the categories are used to study functions, but it is rather controversial, since identical elements of behavior in different species can have different functions.

3. Classification by origin. In this group, a classification according to common ancestral forms, based on a comparative study of closely related species, and a classification according to the method of acquisition, which is based on the nature of the change in a behavioral act in the process of evolution, are distinguished. As examples of categories in these classifications, we can distinguish the behavior acquired as a result of learning and ritualized behavior.

Hynd stressed that any classification systems based on different types of criteria should be considered independent.

For a long time, among ethologists, a classification has been popular, which is based on the classification of Pavlov's reflexes. Its formulation was given by G. Tembrok (1964), who divided all forms of behavior into the following groups:

1) behavior determined by metabolism (foraging and eating, urination and defecation, food storage, rest and sleep, stretching);

2) comfortable behavior;

3) defensive behavior;

4) behavior associated with reproduction (territorial behavior, copulation and mating, care for offspring);

5) social (group) behavior;

6) construction of nests, burrows and shelters.

Let's take a closer look at some forms of behavior.

Behavior determined by metabolism. Eating behavior. Eating behavior is inherent in all representatives of the animal world. Its forms are very diverse and species-specific. Eating behavior is based on the interaction of central mechanisms of excitation and inhibition. The constituent elements of these processes are responsible both for the reaction to various food stimuli and for the nature of movements when eating. The individual experience of the animal plays a certain role in the formation of eating behavior, in particular the experience that determines the rhythms of behavior.

The initial phase of eating behavior is a search behavior caused by arousal. Search behavior is determined by depriving the animal of food and is the result of an increase in reactivity to external stimuli. The ultimate goal of search behavior is finding food. In this phase, the animal is especially sensitive to stimuli that indirectly indicate the presence of food. The types of irritants depend on the availability and palatability of different types of food. Signs that serve as irritants are common to different types of food or characterize its specific type, which is most often observed in invertebrates. For example, for bees, the color of the corollas of a flower can serve as such an irritant, and for termites, the smell of rotting wood. All these stimuli cause different types of activity. Depending on the circumstances and the type of animal, this may be the capture of prey, its preliminary preparation and absorption. For example, wolves have a certain way of hunting for different types of ungulates, while a lynx hunts all types of prey in the same way (jumping from an ambush on the victim's scruff). Predatory mammals have certain "rituals" when eating prey. Weasel eats mouse-like rodents from the head, and when there is a lot of prey, it is content only with the brain of the victim. Large predators also prefer to eat the prey, starting with the muscles of the neck and entrails.

When the animal begins to satiate, feedbacks caused by irritation of the receptors of the mouth, pharynx and stomach shift the balance towards inhibition. This is also facilitated by a change in the composition of the blood. Usually, the processes of inhibition are ahead of the compensatory abilities of tissues and proceed at different rates. In some animals, the processes of inhibition affect only the final act of feeding behavior and do not affect the search behavior. Therefore, many well-fed mammals continue to hunt, which is typical, for example, mustelids, some large cats.

There are many different factors that determine the attractiveness of different types of food, as well as the amount of food consumed. These factors are best studied in rats. In these rodents, which are characterized by complex behavior, the novelty of food can serve as a factor contributing to both an increase in the amount of food eaten and a decrease in its amount. Monkeys often eat new foods in small doses, but if a monkey notices that his relatives eat this food, the amount eaten increases markedly. In most mammals, young animals are the first to try a new food. In some flocking mammals and birds, individual individuals more often try unfamiliar food, being surrounded by relatives, and are very cautious about it, being in isolation. The amount of food absorbed may also depend on the amount of food available. For example, in the autumn period, bears eat pears in gardens in much larger quantities than from isolated trees.

Such widespread behavior as food storage can be attributed to food. To provide food for insect larvae, it is reduced to the activity of laying eggs on living objects (gadflies), the manifestation of parasitism, and the activity of gravedigger beetles. Food storage is also widespread among mammals. For example, food is stored by many species of predators, and their forms of storage are extremely diverse. A domestic dog can simply bury a piece of meat left over from lunch, and an ermine, a marten arrange entire warehouses consisting of the corpses of small rodents. Many species of rodents also store food, some of them (hamsters, saccular rats) have special cheek pouches in which they carry food. For most rodents, food storage periods are strictly limited; in most cases, they are timed to fall, when seeds, nuts, and acorns ripen.

Indirectly, urination and defecation can be correlated with eating behavior, or rather, with behavior determined by metabolism. In most animals, urination and defecation are associated with specific postures. The mode of these acts and characteristic postures are observed both in animals and in humans. The latter has been proven by numerous experiments carried out during wintering in the Arctic.

The states of rest and sleep, according to Tembroke, are related to metabolic behavior, but many scientists associate them with comfortable behavior. It was found that the postures of rest and the postures taken by the animal during sleep are species-specific, as well as individual types of movement.

Comfortable Behavior. These are diverse behavioral acts aimed at caring for the animal’s body, as well as various movements that do not have a specific spatial direction and location. Comfortable behavior, namely that part of it that is associated with the animal’s care for its body, can be considered as one of the options for manipulation (for more details, see 5.1, 6.3), and in this case the animal’s body acts as the object of manipulation.

Comfortable behavior is widespread among different representatives of the animal world, from the most underdeveloped (insects that clean their wings with the help of their limbs) to quite highly organized ones, in which it sometimes acquires a group character (grooming, or mutual search in great apes). Sometimes an animal has special organs to perform comfortable actions, for example, the toilet claw in some animals serves for special hair care.

In comfortable behavior, several forms can be distinguished: cleansing the hair and skin of the body, scratching a certain part of the body on the substrate, scratching the body with the limbs, rolling on the substrate, bathing in water, sand, shaking the wool, etc.

Comfortable behavior is species-typical, the sequence of actions to cleanse the body, the dependence of a certain method on the situation are innate and manifest in all individuals.

Close to comfortable behavior are postures of rest and sleep, the whole range of actions associated with these processes. These postures are also hereditary and species-specific. Studies on the study of the postures of rest and sleep in bison and bison, conducted by the Soviet biologist M.A. Deryagina, made it possible to identify 107 species-typical postures and body movements in these animals, belonging to eight different spheres of behavior. Of these, two-thirds of the movements belong to the category of comfortable behavior, rest and sleep. Scientists noted an interesting feature: differences in behavior in these areas in young bison, bison and their hybrids are formed gradually, at a later age (two to three months).

sexual behavior describes all the diverse behavioral acts associated with the process of reproduction. This form is one of the most important forms of behavior, as it is associated with procreation.

According to most scientists, key stimuli (releasers) play an important role in sexual behavior, especially in lower animals. There are a great many releasers, which, depending on the situation, can cause either the rapprochement of sexual partners, or a fight. The action of the releaser directly depends on the balance of the totality of its constituent stimuli. This was shown in Tinbergen's experiments with a three-spined stickleback, where the red color of the fish's abdomen acted as an irritant. When using various models, it was found that male sticklebacks react most aggressively not to models that are completely red, but to objects that are closest to the natural color of the fish. Sticklebacks reacted just as aggressively to models of any other shape, the lower part of which was painted red, imitating the color of the abdomen. Thus, the response to the releaser depends on a combination of features, some of which can compensate for the lack of others.

When studying releasers, Tinbergen used the method of comparison, trying to find out the origins of mating rituals. For example, in ducks, the courtship ritual comes from movements that serve to care for plumage. Most of the releasers paraded during mating games resemble unfinished moves, which in ordinary life are used for completely different purposes. Many birds in mating dances can be recognized as threat postures, for example, in the behavior of gulls during mating games, there is a conflict between the desire to attack a partner and hide from him. Most often, behavior is a series of individual elements that correspond to opposing tendencies. Sometimes in behavior you can notice the manifestation of heterogeneous elements at the same time. In any case, in the process of evolution, any movements have undergone strong changes, ritualized and turned into releasers. Most often, the changes went in the direction of enhancing the effect, which may consist in their repeated repetition, as well as an increase in the speed of their execution. According to Tinbergen, evolution was aimed at making the signal more visible and recognizable. The boundaries of expediency are reached when the hypertrophied signal begins to attract the attention of predators.

To synchronize sexual behavior, it is necessary that the male and female be ready for breeding at the same time. Such synchronization is achieved with the help of hormones and depends on the time of year and the length of daylight hours, but the final "adjustment" occurs only when the male and female meet, which has been proven in a number of laboratory experiments. In many species of animals, the synchronization of sexual behavior is developed at a very high level, for example, in sticklebacks during the mating dance of the male, each of his movements corresponds to a certain movement of the female.

In most animals, separate behavioral blocks are distinguished in sexual behavior, which are performed in a strictly defined sequence. The first of these blocks is most often the appeasement ritual. This ritual is evolutionarily aimed at removing obstacles to the convergence of marriage partners. For example, in birds, females usually cannot stand being touched by other members of their species, and males are prone to fighting. During sexual behavior, the male is kept from attacking the female by differences in plumage. Often the female assumes the position of a chick begging for food. In some insects, appeasement takes on peculiar forms, for example, in cockroaches, glands under the elytra secrete a kind of secret that attracts a female. The male raises his wings and, while the female licks the secretions of the odorous glands, proceeds to mate. In some birds, as well as in spiders, the male brings the female a kind of gift. Such appeasement is essential for spiders, since without a gift, the male runs the risk of being eaten during courtship.

The next phase in sexual behavior is the discovery of a marriage partner. There are a huge number of different ways to do this. In birds and insects, singing most often serves this purpose. Usually the songs are sung by the male, his repertoire contains a wide variety of sound signals, from which male rivals and females receive comprehensive information about his social and physiological status. In birds, bachelor males sing most intensively. The singing stops when a sexual partner is found. Moths often use scent to attract and locate a mate. For example, in hawk moths, females attract males with the secret of an odorous gland. Males perceive this smell even in very small doses and can fly to the female at a distance of up to 11 km.

The next stage of sexual behavior is the recognition of a marriage partner. It is most developed in higher vertebrates, in particular birds and mammals. The stimuli on which recognition is based are weaker than the release stimuli, and, as a rule, they are individual. It is believed that birds that form permanent pairs distinguish partners by appearance and voice. Some ducks (pintails) are able to recognize a partner at a distance of 300 m, while in most birds the recognition threshold is reduced to 20-50 m. In some birds, a rather complex recognition ritual is formed, for example, in pigeons, the greeting ritual is accompanied by turns and bows, and the slightest change partner is anxious. In white storks, the greeting ceremony is accompanied by a clicking of the beak, and the voice of the bird's partner is recognized at a considerable distance.

As a rule, the mating rituals of mammals are less diverse than those of fish and birds. Males are most often attracted to the smell of females, in addition, the main role in finding a partner belongs to the vision and skin sensitivity of the head and paws.

In almost all animals, intimacy with a sexual partner stimulates numerous neurohumoral mechanisms. Most ethologists believe that the point of complex mating rituals in birds lies in the general stimulation of the mating mechanism. In almost all amphibians, in which mating rituals are rather poor, an important role in the stimulation of neurohumoral mechanisms belongs to tactile stimuli. In mammals, ovulation can occur both after mating and before it. For example, in rats, copulation does not affect the mechanisms associated with the maturation of eggs, while in rabbits, ovulation occurs only after mating. In some mammals, such as pigs, the presence of a male is enough for the female to puberty.

Defensive behavior in animals was first described by Ch. Darwin. Usually it is characterized by a certain position of the ears, hair in mammals, skin folds in reptiles, feathers on the head of birds, i.e., the characteristic facial expressions of animals. Defensive behavior is a reaction to a change in the external environment. Defensive reflexes can occur in response to any factors of the external or internal environment: sound, taste, pain, thermal and other stimuli. The defensive reaction can be either local in nature or take on the character of a general behavioral reaction of the animal. Behavioral reaction can be expressed both in active defense or attack, and in passive fading in place. Motor and defensive reactions in animals are diverse and depend on the lifestyle of the individual. Solitary animals, such as a hare, running away from the enemy, diligently confuse the trail. Animals that live in groups, such as starlings, at the sight of a predator, rearrange their flock, trying to occupy the smallest area and avoid attack. The manifestation of a defensive reaction depends both on the strength and nature of the acting stimulus, and on the characteristics of the nervous system. Any stimulus reaching a known strength can cause a defensive reaction. In nature, defensive behavior is most often associated with conditioned (signal) stimuli that have developed in different species in the course of evolution.

Another form of defensive behavior is represented by physiological changes during a passive-defensive reaction. In this case, inhibition dominates, the movements of the animal are sharply slowed down, and most often it hides. In some animals, special muscles are involved in the passive-defensive reflex. For example, a hedgehog curls up into a ball during danger, his breathing is sharply limited, and the tone of his skeletal muscles decreases.

A special form of defensive behavior includes avoidance reactions, due to which animals minimize getting into dangerous situations. In some animals, fear-inducing signaling stimuli generate such a response without prior experience. For example, for small birds, the silhouette of a hawk serves as a signal stimulus, and for some mammals, the characteristic color and smell of poisonous plants. Avoidance also applies to highly specific reflexes.

Aggressive behavior. Aggressive behavior is most often called behavior addressed to other individuals, which leads to damage and is often associated with the establishment of hierarchical status, gaining access to an object or the right to a certain territory. There are intraspecific collisions and conflicts that arise in a “predator-prey” situation. Most often, these forms of behavior are caused by various external stimuli, consist of different organized complexes of movements and are determined by different neural mechanisms. Aggressive behavior is directed at another individual; stimuli can be visual, auditory and olfactory. Aggression occurs primarily due to the proximity of another individual.

According to many researchers, aggression can manifest itself as a result of conflict between other types of activity. This has been proven in numerous laboratory experiments. For example, in domestic pigeons, aggressive behavior directly depended on food reinforcement: the hungrier the birds were, the more aggressiveness increased.

Under natural conditions, aggression is most often a reaction to the proximity of another animal, which occurs either when the individual distance is violated, or when approaching objects important for the animal (nest, individual territory). In this case, the approach of another animal can cause both a defensive reaction, followed by flight, and an aggressive one, depending on the hierarchical position of the individual. Aggression also depends on the internal state of the animal. For example, in many passerines, short-term skirmishes are observed in winter flocks, where birds, depending on their internal state, maintain an individual distance from several meters to several tens of meters.

In most animal species, aggressive conflicts occur in the spring, when the gonads are active. The intensity of conflicts directly depends on the stage of the marriage cycle. At the peak of mating activity in almost all birds, aggression is caused by a rival that has appeared in the immediate vicinity of the site. Similar phenomena are also observed in some territorial fish species.

As a result of numerous studies, it was found that external stimuli play a more important role than the internal state in causing aggression. The latter most often affects the selectivity of the perception of stimuli, and not the intensity of aggressive behavior. Most of these data were obtained in the study of the behavior of passerine birds, but a similar phenomenon was observed in hermit crabs, as well as in some territorial fish species.

Extensive studies of aggressive activity were carried out by K. Lorenz, who devoted a number of scientific works to this phenomenon. He conducted a large number of experiments to study the aggressive behavior of rats, which helped to derive the basic patterns of aggressive behavior in humans as a biological species.

Territorial behavior first appears in annelids and lower mollusks, in which all vital processes are confined to the area where the shelter is located. However, such behavior cannot yet be considered full-fledged territorial, because the animal does not mark the territory in any way, does not let other individuals know about its presence on it, and does not protect it from invasion. In order to be able to talk about fully developed territorial behavior, the development of a perceptual psyche in an animal is necessary, it must be able to give other individuals information about their rights to this territory. In this process, the marking of the territory becomes extremely important. The territory can be marked by applying odorous marks to objects along the periphery of the site, by sound and optical signals, and trampled grass patches, gnawed tree bark, excrement on shrub branches, and others can act as optical signals. Animals with true territorial behavior tend to actively defend their territory from other individuals. This reaction is especially manifested in animals in relation to individuals of their own species and the same sex. As a rule, such behavior is timed or manifests itself in a particularly striking form during the breeding season.

In a fairly developed form, territorial behavior is manifested in dragonflies. And Hamer conducted observations of the males of the beauty dragonflies. It was noted that the males of these insects occupy individual areas in which functional zones of rest and reproduction are distinguished. Egg laying takes place in the breeding zone, the male attracts the female to this zone with the help of a special ritualized flight. Males perform all their functions within their territory, except for evening rest, which takes place outside it. The male marks his territory, actively defends it from other males. It is interesting to note that the battles between them take place in the form of rituals, and as a rule, they do not come to a real collision.

Of great complexity, as shown by the studies of the Russian ethologist A.A. Zakharov, reaches the territorial behavior of ants. In these insects, there are two different types of use of feeding areas: shared use of land by several families and use of a site by the population of one nest. If the density of the species is low, the sites are not protected, but if the density is high enough, the feeding sites are divided into protected areas, between which there are small unprotected areas. The most difficult behavior is in red forest ants. Their territories, which are strictly protected, are very large, and an extensive network of trails runs through them. At the same time, each group of ants uses a certain sector of the anthill and certain paths that are adjacent to it. Thus, the total territory of the anthill in these insects is divided into the territories of separate groups, between which there are neutral spaces. The boundaries of such territories are marked with odorous marks.

Many higher vertebrates, in particular mammals, birds and fish, stay in the center of a well-known area, the boundaries of which they jealously guard and carefully mark. In higher mammals, the owner of the site, even being at a lower rung of the hierarchical ladder, easily drives away a relative who has violated the boundary. It is enough for the owner of the territory to take a threatening posture, and the opponent retreats. True territoriality is found in rodents, carnivores, and some monkeys. In species that are characterized by disorderly sexual relations, it is impossible to single out an individual territory.

Territoriality is also expressed in many fish. Usually their territorial behavior is closely related to the process of reproduction, which is typical for many cichlids, as well as sticklebacks. The desire to choose a territory in fish is innate, in addition, it is due to the system of landmarks used by the fish. Protection of the territory in fish is most pronounced during the sexual period.

In birds, territorial behavior has reached a high level of development. Some scientists have developed a classification of territories in different bird species according to the types of use. Such birds may have separate territories for nesting, mating dances, as well as separate territories for wintering or overnight. To protect the territory of birds, singing is most often used. Territorial behavior is based on intraspecific competition. As a rule, the more aggressive male chooses the site and attracts the female. The size of the territory of birds is species-specific. Territoriality in birds does not always preclude gregarious behavior, although most often these behaviors are not observed simultaneously.

Parental behavior. All animals can be divided into two groups. The first group includes animals whose females demonstrate parental behavior already at the first birth. The second group includes animals whose females improve their parental behavior throughout their lives. This classification was first developed in mammals, although various forms of parental behavior are observed in other groups of animals.

Mice and rats are typical representatives of animals of the first group; they take care of their offspring from the very first days, and many researchers have not noted significant differences in this between young and experienced females. Animals of the second group include apes, corvids. More experienced relatives help a young female chimpanzee to care for her cubs, otherwise the newborn may die due to improper care.

Parental behavior is one of the most complex types of behavior. As a rule, it consists of a number of interrelated phases. In lower vertebrates, the main thing in parental behavior is the recognition by the cubs of the parents, and by the parents - of the cubs. Imprinting in the early stages of caring for offspring plays an important role here. Fish fry instinctively flock and follow the adults. Adults try to swim slowly and keep the cubs within sight. In case of danger, adults protect the juveniles.

The parental behavior of birds is much more complicated. As a rule, it begins with the laying of eggs, since the nest-building phase is more related to sexual behavior and often coincides with the courtship ritual. The stimulating effect on oviposition is the presence of a nest, and in some birds, its construction. In some birds, a nest with a full clutch can stop further egg laying for a while, and vice versa, an incomplete clutch stimulates this process. In the latter case, birds can lay several times more eggs than under normal conditions.

The next phase of parental behavior in birds is egg recognition. A number of birds lack selectivity; they can incubate eggs with any color and even dummies that have only a distant resemblance to eggs. But many birds, in particular passerines, well distinguish their eggs from the eggs of relatives. For example, some warblers reject eggs of relatives that are similar in color but slightly different in shape.

The next phase of parental behavior in birds is incubation. It is distinguished by an exceptional variety of forms of behavior. Both the male and the female or both parents can incubate the eggs. Incubation can take place from the first, second egg or after the completion of the laying. An incubating bird may sit tightly on the nest or abandon the nest at the first sign of danger. The highest skill has been achieved in incubation by weed chickens, when the male monitors thermoregulation in a kind of incubator made of rotting vegetation, and its construction can take several months. In species in which the male incubates, his desire for this action is synchronous with the timing of egg laying. In females, it is determined by physiological processes.

The next phase of parental behavior occurs after the chicks hatch. Parents begin to feed them semi-digested food. The reaction of the chicks is innate: they reach for the tip of the parent's beak for food. The releaser in this case is most often the color of the beak of an adult bird, in some birds it changes at this time. Adult birds most often react to the voice of the chick, as well as to the color of the throat of a chick begging for food. As a rule, it is the presence of chicks that makes parents take care of them. Under experimental conditions, hens can be maintained in parental behavior for many months by constantly laying chicks in her.

Mammals also differ in complex parental behavior. The initial phase of their parental behavior is the construction of a nest, which is largely species-typical. In females, a certain phase of pregnancy serves as an incentive to build a nest. Rats may start building a nest as early as the early stages of pregnancy, but usually it is not completed to the end and is just a pile of building material. Real construction begins three days before birth, when the nest takes on a certain shape, and the female rat becomes less and less mobile.

Immediately before childbirth, female mammals change the order of licking individual parts of the body. For example, in the last week of pregnancy, they lick the perineum more often and less often - the sides and front paws. Female mammals give birth in a wide variety of positions. Their behavior during childbirth can change quite a lot. As a rule, females carefully lick newborns, bite their umbilical cord. Most mammals, especially herbivores, greedily eat the placenta.

The behavior of mammals when feeding their young is very complex. The female collects the cubs, exposes them to nipples, to which they suck. The lactation period varies among species, ranging from two weeks in rodents to one year in some marine mammals. Even before the end of lactation, the young make short forays out of the nest and begin to try additional food. At the end of lactation, the cubs switch to independent feeding, but continue to pursue the mother, trying to suckle her, but the female is increasingly less likely to allow them to do this. She presses her belly to the ground or tries to run sharply to the side.

Another characteristic manifestation of parental behavior is the dragging of cubs. If conditions become unsuitable, animals can build a new nest and drag their offspring there. The drag instinct is especially strong in the first few days after giving birth, when the female drags not only her own, but also other people's cubs, as well as foreign objects into the nest. However, this instinct quickly fades away, and after a few days, females well distinguish their cubs from strangers. Methods of transferring cubs in different species are different. The dragging itself can be triggered by various stimuli. Most often, this reaction is caused by the calls of cubs, as well as their characteristic smell and body temperature.

Special forms of parental behavior include punishment, which is expressed in some predatory mammals, in particular dogs. Domestic dogs may punish puppies for various misdeeds. The female growls at the cubs, shakes them, holding them by the collar, or presses them down with her paw. With the help of punishment, the mother can quickly wean the puppies from looking for her nipples. In addition, dogs punish puppies when they move away from them, they can separate those who are fighting.

Social (group) behavior. This type of behavior is represented in lower invertebrates only in rudimentary form, since they do not have special signaling actions to carry out contacts between individuals. Group behavior in this case is limited by the colonial lifestyle of some animals, for example coral polyps. In higher invertebrates, on the contrary, group behavior is already fully manifested. First of all, this applies to insects whose lifestyle is associated with complex communities that are highly differentiated in structure and function - bees, ants and other social animals. All individuals that make up the community differ in the functions they perform; food-procuring, sexual and defensive forms of behavior are distributed among them. Specialization of individual animals according to functions is observed.

With this form of behavior, the nature of the signal is of great importance, with the help of which individuals communicate with each other and coordinate their actions. In ants, for example, these signals are of a chemical nature, while other types of receptors are much less significant. It is by smell that ants distinguish individuals of their community from strangers, living individuals from the dead. Ant larvae release chemicals to attract adults who can feed them.

With a group way of life, great importance is attached to the coordination of the behavior of individuals when the community is threatened. Ants, as well as bees and wasps, are guided by chemical signals. For example, in case of danger, "alarm substances" are released, which spread through the air over a short distance. Such a small radius helps to accurately determine the place where the threat comes from. The number of individuals emitting a signal, and hence its strength, increase in proportion to the increase in danger.

The transfer of information can be carried out in other ways. As an example, we can consider the “dances” of bees, which carry information about food objects. The dance pattern indicates the proximity of the food location. This is how the famous Austrian ethologist Karl von Frisch (1886-1983), who spent many years studying the social behavior of these insects, characterized the dancing of bees: “... if it (the food object - Author) is located next to the hive (at a distance of 2-5 meters from it), then a “push dance” is performed: the bee runs randomly through the honeycombs, wagging its belly from time to time; if food is found at a distance of up to 100 meters from the hive, then a “circular” is performed, which consists of running in a circle alternately clockwise and counterclockwise. If nectar is found at a greater distance, then a “wagging” dance is performed. These are runs in a straight line, accompanied by wagging movements of the abdomen, returning to the starting point, either on the left or on the right. The intensity of the wagging movements indicates the distance of the find: The closer the food object is, the more intense the dance is performed.” [eleven]

In all the examples given, it is clearly noted that information is always transmitted in a transformed, conditional form, while the spatial parameters are translated into signals. The instinctive components of communication have reached the greatest development in such a complex phenomenon as the ritualization of behavior, especially sexual, which has already been mentioned above.

Social behavior in higher vertebrates is very diverse. There are many classifications of different types of animal associations, as well as features of animal behavior within different groups. In birds and mammals, there are various transitional forms of organization from a single family group to a true community. Within these groups, relationships are built mainly on various forms of sexual, parental and territorial behavior, but some forms are characteristic only of animals living in communities. One of them is the exchange of food - trophallaxis. It is most developed in social insects, but is also found in mammals, such as wild dogs, which exchange food by burping it.

Social behavior also includes group care for offspring. It is observed in penguins: young cubs gather in separate groups, which are looked after by adults, while the parents get their own food. In ungulate mammals, such as moose, the male owns a harem of several females who can jointly care for offspring.

Social behavior also includes the joint performance of work, which is controlled by a system of sensory regulation and coordination. Such joint activity consists mainly in construction that is impossible for an individual, for example, the construction of an anthill or the construction of dams by beavers on small forest rivers. In ants, as well as colonial birds (rooks, sand martins), the joint defense of colonies from predator attacks is observed.

It is believed that for social animals the mere presence and activity of a conspecific serves as a stimulus for the initiation of social activity. Such stimulation causes in them a set of reactions that are impossible in single animals.

Exploratory behavior determines the desire of animals to move around and inspect the environment, even in cases where they experience neither hunger nor sexual arousal. This form of behavior is innate and necessarily precedes learning.

All higher animals react to a source of irritation in case of an unexpected external influence, try to explore an unfamiliar object, using all available sense organs. Once in an unfamiliar environment, the animal moves randomly, examining everything that surrounds it. In this case, various types of behavior are used, which can be not only species-typical, but also individual. One should not identify exploratory behavior with play behavior, which it outwardly resembles.

Some scientists, such as R. Hind, draw a clear line between the orienting reaction when the animal is motionless, and active research, when it moves relative to the object being examined. These two types of exploratory behavior mutually suppress each other. It is also possible to distinguish between superficial and deep exploratory behavior, and to draw distinctions based on the sensory systems involved in it.

Exploratory behavior, especially at first, depends on the reaction of fear and on the experience of the animal. The likelihood that a given situation will elicit either a fear response or exploratory behavior depends on the animal's internal state. For example, if a stuffed owl is placed in a cage with small birds of the passerine order, at first they rarely approach it, experiencing a reaction of fear, but gradually reduce this distance and subsequently show only exploratory behavior towards the stuffed animal.

At the initial stages of the study of the object, the animal may also show other forms of activity, for example, feeding behavior, brushing wool. Exploratory behavior largely depends on the degree of hunger experienced by the animal. Usually, hunger reduces exploratory activity, but hungry mammals (rats) noticeably more often than well-fed ones leave their familiar environment and go to explore new territories.

Exploratory behavior is closely related to the internal state of the animal. The effectiveness of exploratory responses depends on what the animal, based on its experience, considers familiar. It also depends on the internal state whether the same stimulus will cause fear or an exploratory reaction. Sometimes other types of motivations come into conflict with exploratory behavior.

Exploratory behavior can be very persistent, especially in higher mammals. For example, rats can explore an unfamiliar object for several hours and, even when in a familiar environment, exhibit exploratory behavior that may give them the opportunity to explore something. Some scientists believe that exploratory behavior differs from other forms of behavior in that the animal actively seeks increased stimulation, but this is not entirely true, because both feeding and sexual behavior include the search for final stimuli, which brings these behaviors closer to exploratory ones.

Exploratory behavior is aimed at eliminating the discrepancy between the model of a familiar situation and the central consequences of perceiving a new one. This brings it closer, for example, to nest building, which is also aimed at eliminating the discrepancy between stimuli in the form of a finished and unfinished nest. But in exploratory behavior, the discrepancy is eliminated not due to a change in stimuli, but due to the restructuring of the nervous model, after which it begins to correspond to the new situation. In this case, stimuli lose their novelty, and exploratory behavior will be directed to the search for new stimuli.

The exploratory behavior inherent in highly developed animals is an important step before learning and developing the intellect.

Topic 4. Learning

4.1. learning process

The mental activity of any animal, the variety of forms of its behavior are inextricably linked with such a process as learning.

All components of behavior are formed under the influence of two aspects, each of which is undeniably important. First, species experience is of great importance, which is fixed in the process of evolution of the species, and is transmitted to a particular individual in a genetically fixed form. Such components of behavior will be instinctive and innate. However, there is a second aspect - the accumulation of individual experience of an individual in the course of its life. At the same time, the acquisition of experience by an individual occurs in a rather rigid species-typical framework.

G. Tembrok identifies two forms of accumulation of individual experience by an individual: obligate and optional. In the process of obligate learning, an individual acquires individual experience, which does not depend on the conditions of its life, but is necessary for the survival of any representative of this species. Optional learning includes individual adaptations that a particular individual acquires depending on the conditions of its existence. This component of the animal's behavior is the most flexible; it helps to rebuild species-typical behavior in the specific conditions of a given environment. At the same time, unlike obligate learning, facultative learning will differ in different individuals of the same species.

Instinctive behavior may be subject to changes in the effector sphere (motor responses), the sensory sphere (signal perception), or in both spheres of behavior at the same time (the latter option is most common).

If learning captures the working organ, most often there is a recombination of innate motor elements of behavior, but new motor elements may also arise. As a rule, such motor elements are formed at the early stages of ontogenesis, for example, the imitative singing of young birds. In mammals, such acquired reactions play one of the main roles in the process of cognitive and research activity, in the development of intelligence.

If learning takes place in the sensory sphere, the animal acquires new signals. Acquisition by an individual of such new significant signals makes it possible to expand its ability to orientate itself in the environment. Initially, these signals are practically indifferent to the animal, in contrast to biologically significant key stimuli, but over time, in the process of accumulation of individual experience by an individual, initially almost indifferent signals acquire a signal value.

In the process of learning, an individual selectively selects individual components from the environment, which, from biologically neutral, become biologically significant. The basis for this are various processes in the higher parts of the central nervous system, which are determined by the action of both internal and external factors. Afferent synthesis occurs (the synthesis of perceived stimuli), then the stimuli are compared with information that was previously perceived and stored in memory. As a result, the individual becomes ready to perform certain response actions to stimuli. After they are completed, information about the results of the actions performed is received in the central nervous system according to the feedback principle. This information is analyzed, on the basis of which a new afferent synthesis occurs. Thus, the central nervous system not only contains innate, instinctive programs of behavior, but also new, individual programs are constantly being formed, on which the learning process is based.

It follows from the foregoing that the learning process is very complex, it is based on the formation of programs for future actions. Such formation is the result of a complex of processes: a comparison of external and internal stimuli, species and individual experience, registration of the parameters of a completed action and verification of the results of these actions.

The importance of the learning process. Learning processes are most important for an animal in the early stages of search behavior. Hereditary behavioral programs cannot take into account the entire variety of situations in which an individual will find itself, so it can only rely on its own experience. In this case, the timely orientation of the animal in conditions of changing environmental factors is extremely important. It must quickly and correctly choose an effective method of action already at the initial stage of the behavioral act. The speed and ease of achieving the final phase of the act will depend on this. Thus, acquired elements must inevitably be built into the instinctive behavior of an individual.

Such embedding is hereditarily fixed, so we can talk about the species-typical limits of learning. The learning process has certain, genetically fixed, limits, beyond which an individual cannot learn anything. In higher vertebrates, these limits can be much wider than necessary in the specific conditions of their life. Thanks to this, higher animals have the ability to change their behavioral responses in extreme conditions, their behavior becomes more flexible. In contrast, in lower animals, the ability to learn is extremely small, mainly their behavior is determined by hereditarily fixed reactions. Thus, the breadth of the range of learning can be an indicator of the mental development of the animal. The wider the framework within which an individual can carry out behavioral reactions, the more it is capable of accumulating individual experience, the better its instinctive behavior is corrected, and the more labile the search phase of its behavior will be.

Innate behavior and learning abilities are evolutionarily linked. In the process of evolution, instinctive behavior is constantly becoming more complex, which requires a broad framework of learning ability. Expanding these limits makes it possible to make innate behavior more flexible, which means it raises instinctive behavior to a higher level of development. The process of evolution encompasses not only the very content of instinctive behavior, but also the possibility of enriching it with individual experience. At the lower stages of evolution, the possibility of learning is limited and manifests itself only in such phenomena as habituation and training.

In the process of habituation, the response to repeatedly repeated irritation that has no biological significance gradually disappears. This process is opposite to the process of training, during which the improvement of instinctive action occurs, due to the accumulation of individual experience.

Primitive, simplest forms of behavior do not disappear in the process of evolution, they are replaced by more complex forms. Often other forms of behavior are superimposed on elementary forms, as a result of which the former acquire complexity and lability. Thus, the process of habituation, which already manifests itself in protozoa, can be observed in a complicated form in higher vertebrates. For example, R. Hind's experiments on mice showed that the reactions of these animals to multiple unreinforced acoustic signals weaken at different speeds. However, differences in habituation are determined not only by different intensities of stimuli (as in lower animals), but also by individual variability in the very process of habituation in higher animals.

Skills. In the process of evolutionary development, a qualitatively new component of learning appears in the behavior of animals - skill. Skill is a central form of optional learning. According to Russian psychologist A.N. Leontyev (1903-1979, “Problems of psychic development”, 1959; “Development of memory”, 1931), if we consider skills as any connections that arise in the process of acquiring individual experience, this concept becomes too vague and cannot be used for strict scientific analysis. Thus, the concepts of “skill” and “learning” must be strictly distinguished.

The ability to develop skills is manifested at a certain level of evolutionary development of the animal. The success of the performed motor actions, as well as the reinforcement of these movements with a positive result, will be decisive for the formation of a skill. Learning can take place on the basis of information that an individual independently received during an active search for a stimulus or in the process of communicating with other individuals. The latter option includes the process of imitation and various learning processes.

It is important to note that the skill is formed as a result of the exercise. In order for it to be preserved, constant training is necessary, this will improve the skill. In the absence of systematic training, skills are gradually destroyed.

There are many special methods for learning skills: the labyrinth method, the problem box (problem cell) method, the detour method (for more details on these methods, see 1.2.) Their distinctive feature is that the animal must choose a certain signal or method to solve a certain problem. actions. When using the labyrinth method, the basis for the formation of a skill for the animal will be the memorization of the object and the path to it. With repeated repetition of the experiment under the same conditions, the animal will run the distance to the food object in a constant, short way. In this situation, the skill of finding food in the maze becomes stereotypical and reaches automatism. In general, stereotyping is generally characteristic of the most primitive skills. Greater plasticity is characteristic of such skills only at the initial stages of education. On the contrary, skills of higher orders are characterized by rather significant plasticity at all stages of formation.

Skill Development Methods. There are two methods of experimental development of skills: the method of the American behavioral psychologist B.F. Skinner ("Behavior of Organisms", 1938) (operant, or instrumental, conditioning) and the classical method of I.P. Pavlova.

When developing conditioned reflexes according to the Pavlov method, the animal is initially asked to perform certain movements that it must perform in order to receive reinforcement. In Skinner's method, the animal must find these movements on its own, perhaps through trial and error. An example is an experiment with a rat that is placed in a cage. She will receive nutritional reinforcement only if she accidentally presses a bar attached to the cell wall. In this case, a temporary connection is formed in the nervous system of the rat between the accidental pressing of the bar and the appearance of the feeder. It is possible to significantly complicate the experiment: to give the animal the opportunity to choose one of two actions that will lead to different, opposite, results. For example, pressing the pedal in the cage alternately turns on the heater and turns off the fan, or vice versa. The rat can thus regulate the temperature in the cage.

With the Pavlovian method, the response strictly follows the stimulus, and the unconditioned reinforcement is associated with the conditioned stimulus through the formation of a conditioned reflex connection. With instrumental conditioning, a response (movement) is initially produced, which is reinforced without a conditioned signal. The need for reinforcement induces the animal to a certain reaction to the pedal; it corrects its behavior in accordance with the perception of the pedal. It is this perception that plays the role of a conditioned stimulus, since the action of the pedal leads to food reinforcement (a biologically significant result). If such a temporary connection is not established, the pedal has no signal value for the animal.

When developing reflexes according to the Pavlov method, the initial phase of the behavioral act is taken into account - the orientation phase of the animal. The animal learns under what external conditions, when it must produce a certain movement, i.e., orientation in time is carried out. In addition, the animal must also orient itself in space: find the pedal, learn how to use it. All these points are not taken into account in instrumental conditioning.

Method I.P. Pavlova makes it possible to qualitatively analyze the orientation of the animal by the components of the environment. However, when studying learning processes, one should not be limited only to this technique, because the development of skills is not identical to the development of classical conditioned reflexes.

Training - one of the forms of developing skills in an animal. In contrast to the instrumental development of skills, when the animal has the maximum opportunity to show independence, during training, strict control over the formation of skills is exercised. The animal is no longer faced with the task of independently searching for a method of action to achieve a result - on the contrary, in the course of constant training, undesirable actions are eliminated, and the required movements are reinforced. The result of training are complex and durable motor reactions that are performed by the animal in response to a human command. Reinforcement during training can be either negative (pain for an incorrect action) or positive (food reinforcement). A mixed method can also be used, in which wrong actions are punished, and correct ones are encouraged. The use of training in the study of animal skills is determined by the clarity of the conditions in which it is placed, as well as the possibility of taking into account the signals given by the trainer as accurately as possible.

Training is a complex process; it is not a chain of conditioned reflexes. The greatest difficulty the researcher faces is getting the animal to understand what the trainer expects of it. The expected actions should be species-typical for the animal, but under the given conditions may be unusual for it.

The theory of training was developed by the Soviet zoopsychologist M.A. Gerd. The training process was proposed to be divided into three stages: pushing, working off and strengthening.

At the pushing stage, the trainer must force the animal to perform the required system of actions. An example is the well-known circus number, in which an animal (for example, a dog) rolls out a carpet. When training a dog, a person shows her a piece of delicacy, standing near a carpet rolled into a tube, but does not allow her to grab food. The animal becomes excited, starts jumping briskly on the spot, barking, touching with its front paws. At the same time, any accidental touches of the dog on the carpet are reinforced with small pieces of treats. Gradually, the dog will deliberately begin to touch the carpet with its paws in order to receive reinforcement, it will form the necessary paw movements on the carpet for the number. Subsequently, all these movements are carefully worked out, their orientation is specified.

At this stage of training, you can act in three ways. The first method is the method of direct pushing, when the trainer makes the animal move after an object that is attractive to him (for example, food). The second method is indirect pushing: the trainer provokes movements not directed at the bait, but caused by the excitation of the animal. This method forms the manipulation actions of the limbs: the transfer of objects, grasping, pushing, and others. In the complex prodding method, the animal first develops a skill, and then in another situation it is forced to use this skill in a different way. For example, a fur seal is first taught to drop a ball into the handler's hands. The trainer then hides his hands behind his back for a few moments. The cat is forced to hold the ball on the bow because it only receives reinforcement when the ball hits the handler's hands. Gradually, the duration of holding the ball by the fur seal increases, and as a result, a circus act with balancing the ball is created.

At the second stage of training - the stage of working out - the trainer focuses his efforts on getting rid of the extra movements of the animal that accompany the necessary actions. This is especially true of all kinds of orienting reactions caused by a situation that is new for the animal. When unnecessary movements are eliminated, the primary system of actions is "polished", the necessary movements are made sufficiently clear and long, and a convenient signaling is selected to control the actions of the animal. In this case, the reaction to food reinforcement should be replaced by a reaction to the trainer's signal (for example, the sound of a whistle).

At the development stage, pushing techniques are also used. For example, the correct posture of an animal standing on its hind legs can be fixed by raising the bait above its head. With the help of these techniques, the development of artificial signaling is carried out.

The last stage of training is the stage of hardening. At this stage, efforts are focused on consolidating the acquired skills, as well as ensuring their mandatory reproduction in response to a signal. Pushing is no longer used here. Food reinforcement is produced not after each skill, but at the end of the whole complex of actions. As a result, skills take the form of a stereotyped reaction when the end of one action is the beginning of the second, and so on.

Thus, the artificial development of skills in animals is a very complex process, although it is undoubtedly inferior in terms of the degree of diversity to the formation of skills in animals in natural conditions.

4.2. The role of cognitive processes in the formation of skills

Well-known zoopsychologists G. Spencer, C. Lloyd-Morgan, G. Jennings and E. Thorndike, as a result of research conducted in the middle of the XNUMXth century, came to the conclusion that the process of skills formation is carried out by "trial and error". This meant the formation of skills both in relation to orientation among the components of the environment, and in relation to the formation of new combinations of movements. Random actions that led to a successful result are selected from the animal and fixed. Actions that do not lead to success are gradually eliminated and not fixed, and "successful" actions, repeated many times, form skills. Thus, the concept of "trial and error" states that all actions are performed spontaneously and randomly, while the animal turns out to be as if passive in relation to the components of the environment.

However, the formation of skills as a process requires the activity of the animal, a selective attitude to the components of the environment. In the 1920s the concept of "trial and error" had opponents - the American neo-behaviorist psychologist E. Tolman (1886-1959; Purposeful Behavior in Animals and Man, 1932), the Russian physiologist V.P. Protopopov (1880-1957) and other scientists. They did not agree with the ideas about the randomness and non-direction of animal movements in solving problems. According to them, skills are formed in the process of active orienting motor activity of the animal. The animal analyzes the situation and actively chooses those actions that correspond to the achievement of the goal. In other words, the resulting movements are adequate to the goal. The decisive factor here will not be a random choice, but an active motor analysis. These views also have experimental confirmation. The experiments of scientists I.F. Dashiella, K. Spence and W. Shipley, conducted in the middle of the XNUMXth century, showed that when a rat enters a maze, it more often enters dead ends located towards the target (food bait) than those located in the opposite direction. First, the rat carries out the first motor orientation in the maze, and on its basis creates a scheme of movement, i.e., its actions are not random. Thus, during the development of skills as a result of primary active orientation, directed actions arise in animals.

These data allowed the Polish zoopsychologist I. Krechevsky to put forward the assumption that the animal is guided in solving various problems by a kind of "hypotheses". They are especially pronounced if the animal is faced with a task that is obviously not solvable for him. For example, an animal is placed in a labyrinth, the doors in which close and open without any system and sequence, chaotically. In this case, according to Krechevsky's ideas, each animal builds its own "hypothesis" and repeatedly tests it. If, after repeating the actions, the "hypothesis" does not lead to a solution, the animal abandons it and builds another one, which it also tests, and so on. In such a situation, each animal behaves in the same way, regardless of changing external conditions. In experiments on rats in a maze with randomly closing passages, each animal acted in accordance with its "strategy". Some rats alternated turns to the right and left with a clear regularity. Others first turned right at each fork, and when this did not lead to success, they began to constantly turn left. Thus, in constantly changing conditions, animals seemed to be trying to identify a certain principle and act according to it. Krechevsky came to the conclusion that this abstract "principle" is due to the internal "tuning" of the animal.

Krechevsky drew attention to the complexity of the behavioral reactions of the animal at the initial stage of solving the problem - during this period, the role of exploratory behavior is especially pronounced. In his concept, emphasis is placed on the active behavior of the animal, the manifestation of initiative by it. In addition, Krechevsky's concept emphasizes the role of internal factors, and above all the mental state of the animal, in the choice of actions to solve the problem facing him.

The concept of "trial and error" is refuted by some experiences and experiments. For example, experiments with the use of "latent learning" are known. Their essence lies in the fact that the animal is given the opportunity to get acquainted with the device of the experimental setup before the start of the experiment. In this case, it is especially important that familiarization takes place actively, i.e., the animal has the opportunity to run a little in the installation. It should be noted that the orienting behavior of a rat that first entered the setup, in the absence of reinforcement, serves only to accumulate experience. When the rat is placed in the setup before the start of the experiment, it still does not see any goal in front of it, since there is no food reinforcement (positive stimulus) or pain (negative stimulus). With such a primary examination of the labyrinth, the nature of the perception of objects can differ significantly in different animals: some rats will use mainly visual stimuli, others - olfactory ones. Individual characteristics as a whole are a sign of the learning process, since the instinctive components of behavior are conservative and unchanged. If we compare the rate of skill formation in an animal that is placed in the maze immediately before the experiment, and in an animal that has actively familiarized itself with the maze, it will be much less in the second animal.

All these experiments convincingly prove that for the successful emergence of a habit, active cognitive activity of the animal is necessary as a prerequisite. It is this cognitive process that determines the nature of the skill.

A.N. Leontiev proposed a criterion for separating habit from other forms of learning. He called this most important criterion "operation". An operation is a component of an animal's activity that meets the conditions in which the object that stimulates this activity is given. Leontiev suggested that only fixed operations be considered skills. Highlighting an operation in the animal's motor activity indicates that this is a real skill.

An example of the selection of an operation can serve as an experiment using the workaround method, which was carried out by A.V. Zaporozhets and I.G. Dimanstein. In an aquarium with water, a transverse partition made of gauze was installed, and a narrow passage was left on the side through which the experimental fish could swim. A fish was placed in one part of the aquarium, and a food bait for it (for example, a bloodworm) was placed in the other part, separated by a partition. The fish could capture the bait only by going around the partition, this happened after it could not swim directly to the food. During the search for the path that led her to the bait, the experimental animal performed certain motor actions.

In this locomotor activity, Leontyev suggests seeing two components. The first is directed activity, which arises under the influence of the properties of the object itself that stimulates the activity, i.e. the smell of bloodworms, its type. The second component of the locomotor reaction is activity associated with the influence of an obstacle, i.e., with the conditions in which the object stimulating the activity is given. This activity will be, according to Leontiev’s terminology, an operation.

After the experimental fish learns a detour to the bait, i.e., a motor skill is developed, the barrier is removed from the aquarium. However, the fish will exactly repeat its path around the obstacle. Over time, the path will straighten out. Thus, the effect of the obstacle in this experiment is strongly connected with the effect of the bait, both of these components act together and inseparably, the bait does not separate from the partition, and vice versa. Consequently, in this situation, the operation can be distinguished only conditionally, it is not yet separable from other components of the motor reaction.

This fish example is an example of an automated skill - a skill that is still at a very low level of development. In this case, the cognitive aspect of skill formation is extremely weak, so the trajectory of the path to the bait becomes so strongly fixed that it persists even after the obstacle is removed. In order for a complex skill to be formed, its cognitive component must be very large. The characteristic feature of higher-level skills in higher vertebrates is that operation is clearly distinguished and plays an extremely important cognitive role. However, this does not mean at all that they lack primitive skills and are not important for the accumulation of individual experience. The level of the developing skill depends on the biology of the species and on the situation in which the animal faces the task.

The cognitive aspect of a skill is manifested in ways of overcoming an obstacle. When analyzing the formation of skills, an obstacle is understood not only as a direct physical obstacle that blocks the path to the stimulating object. An obstacle to solving the problem facing an animal is any obstacle on the way to the goal, regardless of its nature. This was experimentally proven by V.P. Protopopov. His research showed that absolutely any motor skills in animals are formed through overcoming a certain “obstacle”, and it is its character and nature that determine the content of the formed skill. According to Protopopov, the stimulus has only a dynamic effect on the formation of a skill, i.e., it determines the speed and strength of its consolidation. Overcoming an obstacle is an important element in the formation of a skill, not only when developing it using the workaround method, but also when using other methods for this purpose, for example, the labyrinth method and the problem box method.

The Hungarian zoopsychologist L. Kardos paid much attention to the cognitive aspects of skill formation. He especially emphasized that when an animal learns in a maze, it enriches its knowledge and accumulates a significant supply of useful information. Kardosh wrote about it this way: “...at the beginning of the labyrinth, the animal in memory... sees beyond the walls that cover its field of sensations; these walls become, as it were, transparent. In memory, it “sees” the goal and the most important ones from the point of view of locomotion ( movements. - Author) parts of the path, open and closed doors, branches, etc., “sees” exactly the same way and where and how it saw it in reality while walking around the labyrinth.”[12]

Along with this, Kardosh clearly defined the boundaries of the animal's cognitive capabilities in solving problems. Here, in his opinion, there are two possibilities: locomotor and manipulative cognition. In locomotor cognition, the animal changes its spatial position in the environment without changing the environment itself. With manipulative cognition, an active change in the environment of animals occurs.

Manipulative cognition is carried out during the formation of instrumental skills. Kardosh conducted studies in which he showed that an animal (in the experiment, a rat) can be taught to choose different paths in a maze that lead to one point, and then move on in different ways, for example, in one direction or the other. This may serve as an example of locomotor cognition. However, according to Kardosh, no animal (except apes) can be taught that, depending on the choice of one of two paths of movement, quite definite changes will occur in the environment. In the experiment, for example, food was replaced with another reinforcer - water. L. Kardosh writes: "... a person would be surprised to find different objects in the same place when he approached from the right and from the left, but he would learn after the first experience. It is here that development makes a leap... A person is completely can free itself from the directing influence of spatial order if temporal-causal connections require something else.” [13]

4.3. Learning and communication. Animal imitation

The role of imitation in the formation of behavior in higher animals cannot be overestimated. The phenomenon of imitation does not always belong to the process of learning, it can also belong to instinctive behavior. An example of such imitation is allelomimetic behavior (mutual stimulation), when the performance of actions (species-typical) by some animals induces others to perform the same actions (for example, the simultaneous collection of food). At the same time, a certain kind of action, inherent in all individuals of the species, is encouraged.

Learning by imitation is called "imitation learning". The essence of this process is that the animal individually forms new forms of behavior through direct perception of the actions of other animals. In other words, the basis of such learning is communication with other individuals. Simulation learning can be divided into obligate and optional.

In the process of obligate imitation learning, its result fits entirely within the framework of a certain species stereotype. Through imitation, individuals learn to perform vital actions. All these actions are inherent in the normal behavioral “repertoire” of the species. Obligate learning is most typical for young animals. An example is the formation of a defensive reaction to a predator in the form of flight in young fish of schooling fish species. At the same time, they imitate the behavior of adult fish, for example, when they see other members of the school being eaten by a predator. According to L.A. Orbeli, such imitative behavior is extremely important, “it serves as the main guardian of the species, for the enormous advantage lies in the fact that spectators present at the act of damage to a member of their own herd or their community develop reflexive protective acts and thus can avoid danger in the future.” . [14]

Obligate imitation learning also serves as an important element in the reaction of following and recognition of food objects by young mammals. Young individuals of animals such as birds and great apes (chimpanzees) acquire nest-building experience through obligate imitation learning.

The simplest optional imitation learning is manifested in the imitation of movements that are not inherent in this species. In this case, imitation occurs on the basis of allelomimetic stimulation. For example, when keeping great apes in conditions where animals can constantly contact people, monkeys begin to perform various actions with household items, imitating human actions. This behavior will no longer be species-typical: new methods of manipulative activity are being formed. Such actions are called "non-species imitation manipulation."

With optional simulation learning, problem solving occurs in a more complex form. One animal performs certain actions to solve the problem, the other (the spectator animal) only observes its actions, and the skill is developed in it in the course of observation. The ability for such learning has been noted in various mammals: rats, dogs, cats, lower and great apes, but it plays a particularly important role in the latter. Based on observations in nature, A.D. Slonim concluded that the formation of conditioned reflexes in a herd of monkeys occurs mainly on the basis of imitation.

But not all skills can be formed in animals through optional imitation learning. Instrumental skills are not formed in this way. This is confirmed by the experiments of the American researcher B. B. Beck. In his experiments, baboons were used, which observed the use of tools by relatives in solving problems. The baboons-viewers did not acquire instrumental skills, but they more often and more intensively than before these experiments performed manipulations of the tools, the use of which was observed. This example proves that allelomimetic behavior and non-species imitative manipulation play an important role in the development of complex skills in social conditions.

Imitation also captures the field of signaling and communication. An example is the onomatopoeia of birds. In this case, the stimulation of species-typical acoustic signaling occurs (for example, phenomena such as "choirs" of birds). The imitation of birds to other people's sounds and songs can be defined as non-species imitative manipulation. The assimilation of species-typical sounds by chicks by imitating the singing of adults refers to obligate imitation learning.

Two fundamentally different approaches can be applied to the study of the process of imitation in animals.

1. When studying amelomic behavior, animals are isolated from each other and trained separately, only then they are brought together. Animals can be trained to respond to the same signal in different ways, while achieving the opposite response. After bringing the animals together and presenting this signal to them, one can find out what prevails in a given group of animals: mutual stimulation or the results of the usual learning of each animal. The results will make it possible to judge the strength of the allelomimetic reaction in these animals, i.e., the strength of imitation.

2. If imitation learning is being studied, the animals are provided with communication from the beginning of the experiments. In this case, one individual (animal-actor) is trained by the researcher for a certain reinforcement in front of the other individuals (animal-spectators). We can talk about optional imitative learning if individuals who were not trained by the experimenter and did not receive rewards for solving the problem learn to solve this problem correctly and without their own exercises, based only on observation. For example, when one banana is thrown to the monkeys, the leader of the pack always gets it. However, soon all individuals of the flock begin to gather at a certain signal, although only the leader still receives the banana. In this way, the skills of all animals ("spectators") are formed, which helps to solve the problem even in the absence of a leader ("actor").

The phenomena of imitation in the natural environment are quite closely and intricately intertwined with the intragroup relations of animals. So, in communities, in addition to mutual stimulation for the joint performance of certain actions, there is also an opposite factor - the suppression by "dominant" individuals of the actions of other members of the community. For example, in the experiment described above, the monkeys were afraid even to approach the installation in which they put a banana, and even more so did not dare to take it. However, the monkeys also have special, as it were, "pacifying" signals. The purpose of these signals is to notify the dominant individual (leader) of the readiness of the rest of the pack members only to observe. This possibility provides the implementation of allelomimetic behavior and imitative learning.

Learning at different stages of a behavioral act. Any behavioral response of an animal begins with an internal stimulus (need). This stimulus activates the animal, prompts it to start an active search activity. The initial phase, the search behavior itself, and the final phase are always clearly genetically fixed, but the path along which the animal reaches the final phase of behavior may change. It depends on the learning process, on how variable the animal's behavior is, to what extent it is capable of correct orientation in a changing environment.

In higher animals, the main means of achieving the final phase of a behavioral act is facultative learning. Its success depends on the perfection of the mechanism of orientation of the animal in space and time. The more perfect this orientation, the more successful will be the overcoming of the obstacle, i.e., the conditions in which the object is given. The perfection of the animal's orienting reactions directly depends on the level of its mental activity. The most important here are the higher mental functions - intellectual capabilities. They give the animal's behavior flexibility and variability, thereby providing adaptive opportunities for behavioral responses.

Topic 5. Development of mental activity of animals in ontogenesis

5.1. The development of mental activity in the prenatal period

One of the central problems of animal psychology is the question of the innate and acquired components of animal behavior. This question is closely related to the study of the ontogenesis of behavior. It is important to assess which elements of behavior are inherited by an individual (and therefore genetically fixed), and which are acquired during individual development. Many animal psychologists worked on this problem, all of them expressed different opinions about the relationships between elements of behavior during ontogenesis. Thus, the famous English zoopsychologist K. Lloyd-Morgan wrote that “the activity that is the result of coordinating 10% of initially incoherent movements is a new product, and this product is the result of assimilation, acquisition, and is not inherited as a specific, coordinated action. Like a sculptor creates a statue from a piece of marble, so assimilation creates an action from the mass of given random movements. A specific, coordinated, reactive or response action is acquired. But there are certain actions that are determined from the very day of birth, which are inherited ready-made and the combination or coordination of which immediately after birth is already "is distinguished by complete perfection. Determination and coordination of actions in this case are not individual, but borrowed from ancestors." [15]

The scientist points to the fact that many actions of animals can be performed by them without additional information. For example, a waterfowl chick boldly enters the water for the first time. There was also an opposite opinion, according to which only one of the factors (internal - instinct or external - learning) influences the development of behavior. Adherents of mechanistic views on the development of behavior (without the action of internal factors) were G.E. Coghill and Qing Yang Kuo, in Russia - V.M. Borovsky. They believed that all behavior is the result of only learning that occurs in an animal, starting from the embryonic period of development. This concept was formed in opposition to the theory of the initial programmed behavior.

Currently, there is an understanding of the ontogenesis of behavior as a set of interacting external and internal factors, a combination of unconditioned and conditioned reflex activity. L.V. Krushinsky proposed the term “unitary reaction” to designate acts of behavior that have a similar external expression under different methods of formation. A unitary reaction combines conditional and unconditional behavioral elements. Such behavioral acts are aimed at “performing a specific act of behavior that has different ways of implementation and at the same time a certain pattern of final execution.” [16]

Thus, unitary reactions are aimed at performing a single action, which has an adaptive value. In this case, the unconditional and conditional components can be in different proportions.

The ontogenesis of behavior is closely related to the morphofunctional changes in the body, since innate movements are a function of "working" organs. Zoologist B.S. Matveev showed that in the course of ontogenesis the attitude of the organism to environmental factors changes. This causes various forms of adaptation of individuals to the environment in the process of individual development. In the early stages of ontogeny, adaptations can lead to changes in the morphological (body structure) and functional (body functions) spheres. In this case, first of all, the "working" organs change, and then changes occur throughout the body.

The course of ontogenesis of behavior is influenced by the degree of maturity of the animal. These features are closely related to the historical development of the animal species, their habitat and lifestyle. Depending on this, newborns have different degrees of independence immediately after birth.

In addition, such features of animal development as the presence or absence of a larval form in their life cycle also influence the ontogeny of behavior. Often, the larva differs from the adult in its way of life, in the features of movement, nutrition, etc. Particularly clear differences can be observed in invertebrates, although certain differences are also noted in vertebrates. During metamorphosis (the transformation of a larva into an adult animal), the most complex morphological and functional rearrangements of the body occur, which inevitably lead to changes in behavior.

K. Fabry proposes the following periodization of the ontogeny of behavior:

▪ early postnatal period;

▪ juvenile (play) period (distinguished only in animals that exhibit play activity).

The prenatal (embryonic) period is the time of development of an animal from the moment the embryo is formed to birth (or hatching from an egg). The behavior of the animal in this period is of great importance for the development of behavior in general. Embryos of both vertebrates and invertebrates produce a number of movements in the prenatal period of ontogeny ("embryonic movements"). At this stage of development, they do not yet have a functional significance, since the organism is not associated with the environment during this period. However, it was noted that embryonic movements are a kind of elements of future motor acts that the body performs at later stages of ontogenesis - it is then that these movements will acquire an adaptive (adaptive) value.

According to A.D. Slonim, embryonic movements can affect the physiological processes associated with the animal's muscular activity. They allow even in the intrauterine period of development to prepare the animal for environmental conditions. Such "training" movements are typical, for example, for young ungulate mammals, which, immediately after birth, are able to rise to their feet and move quickly, following the herd. The ability of cubs to carry out vigorous activity immediately after birth is determined by motor exercises in the prenatal period. It is noted that the embryos of these animals make foot movements resembling walking. By the time of birth, the animal develops good coordination of all physiological functions, including vegetative ones (for example, the regulation of respiratory rate).

The formation of behavior is determined by complex and diverse morphofunctional correlations. Russian zoologist and morphologist, known for his work in the field of comparative anatomy of vertebrates, I.I. Schmalhausen (1884-1963, "Paths and Patterns of the Evolutionary Process", "Factors of Evolution") singled out the so-called "ergonic correlations", i.e., relationships between organs due to functional dependencies between them. This refers to the typical functions of organs, such as the functions of the liver or heart of the animal. Schmalhausen cites as an example of ergonic correlations the relationship between the development of the nervous system and the sense organs. If any sense organs are removed from the embryo, then the elements of the nervous system that receive information from them do not develop fully.

Soviet physiologist P.K. Anokhin (1898-1974) drew attention to the mutual consistency of morphofunctional changes (changes in structure and function) in ontogenesis. He wrote: “The development of a function always proceeds selectively, fragmentarily in individual organs, but always in extreme coordination of one fragment with another and always according to the principle of the final creation of a working system.” [17]

When studying the embryonic development of mammals, the scientist noted that individual structures of the body develop asynchronously. At the same time, “in the process of embryogenesis, there is an accelerated maturation of individual nerve fibers that determine the vital functions of the newborn, because for his survival, the “system of relations” must be complete at the time of birth.” [18]

The concept of the significance of the embryonic behavior of animals for their behavior in the adult state is relative. The general patterns and direction of development of body functions are limited by historically established and genetically fixed factors. However, the development of the embryo and its behavioral reactions are also influenced to a certain extent by the living conditions of an adult animal.

embryonic learning. AT as a result of studying the behavior of animals in embryogenesis, it was noted that it may include fragments of movements that affect the development of the animal. Related to this is the concept of embryonic learning. As an example, consider the work of Qing Yang Kuo. This scientist studied the development of behavior in chicken embryos. He showed that in the process of embryogenesis in animals there is an accumulation of motor "embryonic" experience. Experience is accumulated by exercising the rudiments of future organs. During such exercises, motor functions are improved and further developed.

Kuo developed a method that allowed him to observe the movements of the embryos without disturbing their natural development. The scientist made a hole in the egg shell, inserted a window into it and observed the embryo. Kuo noticed that the chick embryo is exposed to various factors both from the outside and arising inside the egg due to the activity of the embryo itself. The initial movements of the fetus are passive, such as head movements due to rhythmic contractions of the heart. The embryo begins to carry out the first active movements on the third or fourth day of development. These are movements of the head to and from the chest, which are accompanied by vigorous opening and closing of the beak. Some researchers believe that in this way the chicken embryo learns pecking movements. On the sixth or ninth day, such movements are replaced by new ones: now the head turns from side to side. Such a change in movements may be associated with a lag in the growth of the neck muscles from the growth of the size of the head, as well as with the position of the head of the embryo in relation to the shell, the location of the yolk sac, the heartbeat, and even the movements of the toes.

As a result, after hatching, the chick has a number of behavioral responses that were developed in him in the process of prenatal development. In this case, reactions are developed not to a specific stimulus, but to a whole group of stimuli that cause one behavioral reaction. The movements of individual parts of the body have not yet been developed, basically the whole body moves, and the movements are very uneconomical. Thus, according to Kuo's conclusions, for the normal manifestation of all behavioral reactions, the animal must undergo a learning process, and therefore, innate behavior does not exist. There are only certain hereditary prerequisites for the formation of behavioral reactions, but these reactions develop depending on external conditions.

The innate component of behavior cannot be completely ignored. In the process of phylogenesis, a grandiose experience of a species is accumulated, and it is realized in the ontogeny of a particular individual through learning. Learning is necessary because the ontogeny of behavior cannot proceed only in a species-typical direction. It should be biologically useful for any animal and meet the conditions of its life.

Some elements of behavior, however, appear in the animal without embryonic learning. In this case, the possibility of improving the function of the organ through exercises is excluded, and the movement itself develops solely through the implementation of the innate program. An example of such a response that does not require learning is the nipple-seeking response in young mammals and subsequent sucking movements.

Immature babies (such as a baby kangaroo) also exhibit innate behavioral responses. A newborn kangaroo is at a stage of development that can be roughly compared with the embryo of a higher mammal. However, a newborn kangaroo already shows a whole range of motor reactions and orientation abilities. At the same time, he performs a whole sequence of innate movements, which are always performed one after another. The kangaroo independently rises to the mother's bag, crawls into it, finds the nipple, grabs it with his lips. Since the embryonic period of a kangaroo is extremely short, he could not even learn individual acts from this chain of behavioral reactions, not to mention the entire sequence of actions. There is an assumption that when looking for the mother's bag, the cub is guided by the dryness of the wool, on which he must crawl. On the opposite side, the kangaroo hair, moistened with birth waters, is wet. The kangaroo shows negative hydrotaxic behavior. This behavior could not have formed in him inside the birth membranes, since the embryo was in a humid environment there.

There were assumptions according to which all the behavior of an animal is only the result of the maturation of innate elements of behavior. In this case, the exercise of the organs is completely excluded. This point of view had its adherents, for example, the American scientist L. Carmichael, who considered behavior to be almost completely innate. However, at present, innate and acquired elements in the ontogenesis of behavior are not opposed, but are perceived as interrelated elements.

Below is an overview of the prenatal development of locomotor activity in embryos from different groups of animals.

Invertebrates. It is known that the embryos of cephalopods in the early stages of embryogenesis rotate inside the egg around the axis at a speed of one revolution per hour. In addition, they move between the poles of the egg. All movements are carried out with the help of cilia. This mode of locomotion is widespread among the larvae of marine invertebrates.

By the end of embryogenesis in invertebrates, some vital instinctive reactions are finally formed. So, mysids (crustaceans) by the time they hatch from eggs already have an avoidance reaction from adverse influences. At the same time, initially, reflex "shudders" are observed in the embryo in response to touching the egg.

In sea goats (marine crustaceans), spontaneous and rhythmic movements of parts of the embryo are observed from the 11th to the 14th day of embryonic development. Subsequently, specific motor reactions are formed on the basis of these movements.

In adult daphnia, antennas are used for swimming. The antennae of the embryo begin to move in the middle stages of embryogenesis. Closer to its end, they rise and take the position necessary to perform swimming movements, and then begin to move especially intensively. Thus, the reflex response is gradually formed on the basis of movements due to internal processes, and then associated with external stimuli.

Fish. Similarly, there are motor reactions of fish. They develop on an endogenous basis (that is, they depend on internal processes in the body). The movements of fish develop depending on the maturation of the corresponding neural connections. After the development of the sense organs, the behavior of the embryo begins to be influenced by external factors that are combined with innate movements.

By the time of the end of embryogenesis in bony fish, trembling, twitching of individual parts of the body, serpentine bending of the body and rotation can be noted. Immediately before hatching, the fish develop peculiar "pecking" movements and bending of the body, facilitating the exit from the ovoid shell.

Amphibians. The embryonic behavior of amphibians is broadly similar to that of fish embryos. First, bending movements of the body appear, then swimming movements and movements of the limbs are formed on this endogenous basis.

The development of the toad Eleutherodactylus martinicensis is interesting. Its larva develops inside the egg shells, but performs all the movements characteristic of the tadpoles of other tailless amphibians. At first, she develops general bending movements of the body, then swimming movements are formed on their basis. Initially, they are still connected to the general bending of the body, but after a day it is already possible to cause single reflex movements of the limbs, regardless of the movements of the muscles of the body. Later, coordinated movements of all four limbs appear in strict sequence and coordinated swimming movements develop. It is also curious that at this stage the larva has never been in an aquatic environment, because it is enclosed in egg shells.

For tailed amphibian embryos (by the example of ambystoma), it has been shown that they perform swimming movements long before hatching from eggs. Then leg movements appear, typical of the land movement of an adult ambistoma. L. Carmichael proved that this mechanism matures without learning. The ambystoma embryo was grown in an anesthetic solution, the embryo was completely immobilized, but grew and developed normally. Embryonic training under such conditions was impossible, but the locomotor abilities of the grown ambistoma were normally developed. This allowed Carmichael to conclude that the formation of the ability to swim depends only on the anatomical development of the animal and does not need to be learned. This conclusion was disputed by the Polish zoopsychologist J. Dembowski. He argued that in experimental embryos the possibility of accumulating motor embryonic experience was suppressed, but the corresponding processes in the nervous system still proceeded. Its functioning served as a kind of exercise for the development of the behavior of the embryo.

To prove the influence of internal factors on the formation of the motor activity of embryos, experiments were carried out on salamander embryos. They transplanted the rudiments of limbs, turned in the opposite direction. If the process were determined by embryonic learning, then in the course of embryogenesis there would be a correction that would restore the salamander's ability to normal forward movement. However, the hatched animals backed away from stimuli that, in normal individuals, elicit a forward movement response.

Thus, in lower vertebrates, the formation of locomotor movements (limb movements) in embryogenesis occurs not under the decisive influence of external factors, but as a result of endogenous maturation of internal structures.

Birds. Observations on the development of chicken embryos served as material for the study of the embryonic behavior of birds. Their incubation period lasts about three weeks, and motor activity begins around the fourth day of incubation. At first, it is represented by movements of the anterior end of the body of the embryo, gradually the place of motor activity shifts to the posterior end of the body. Somewhat later, spontaneous independent movements of the limbs, head, beak, tail and eyeballs begin.

The works of Ts.Ya. Kuo, who established the importance of embryonic learning for the development of bird behavior, while denying the innate component of development. Kuo drew attention to the following pattern: the embryo shows maximum motor activity at the very moment when the amniotic membrane of the embryo begins to move. The scientist suggested that it is the pulsating movements of the amnion that determine the moment the embryo begins to move. R.V. Oppenheim, on the basis of experiments, showed that there is an inverse relationship here: the movements of the embryo determine the movements of the amniotic membrane.

Kuo also pointed to the important role of environmental changes in the development of embryonic behavior. For example, from the 11th day of incubation, the yolk approaches the ventral side of the embryo, preventing the movements of the legs, which become, as it were, fixed in a bent position, one above the other. After resorption of the yolk, the leg that is located above gets the opportunity to move, but the second one is still constrained and begins to show activity only after the first leg moves away. According to Kuo, this explains the fact that the hatched chick does not move by jumping, but by walking, moving its legs alternately.

Research into the development of the embryonic behavior of birds was also carried out by V. Hamburger and his collaborators. It was found that the first embryonic movements of chicken embryos are caused by spontaneous internal processes in the nervous structures. During the first two or two and a half weeks of development, tactile stimulation (touch) has practically no effect on the movements of the embryo. In other words, at the first stages of bird embryogenesis, motor activity does not occur in response to external factors, but is caused only by internal factors. These assumptions were confirmed by experiments. On the first day of incubation, the rudiments of the spinal cord were cut in the chicken embryo, thus the integrity of the nervous structures of the embryo was violated. After this operation, the chick embryo showed a mismatch in the movements of the rudiments of the fore and hind limbs, which normally should move synchronously. However, the rhythm of motor acts was preserved, which means that the processes of motor activity in certain parts of the spinal cord are autonomous.

The course of the embryonic period in birds is greatly influenced by the biology of a particular species. It is especially important to note the differences between chicks and brood birds. If in chicks hatching occurs at early stages of development, then in broods it occurs at later stages, therefore, when comparing chicks of the same age, it may turn out that in brood birds this is still a process of embryonic development, and in chicks it is postembryonic. In brood birds, the processes of embryogenesis are longer, the formation of morphological structures and behavior begins even in the egg, and by the time of hatching, these parameters are already almost completely formed. The chick is forced to go through all these processes after hatching.

Mammals. The study of mammalian embryos is difficult due to the fact that the embryo develops in the mother's womb and observation of it is possible only if it is artificially removed from the mother's body. Such interference in development can adversely affect both the formation of the morphological structures of the embryo and the manifestations of motor activity.

The embryogenesis of the behavior of mammals has an important difference from the development of the behavior of the embryos of other vertebrates. Motor activity in other vertebrates (fish, amphibians, reptiles, and birds) is formed on the basis of the initial general movements of the entire embryo. In mammals, the movements of the limbs appear simultaneously with such movements or earlier. Thus, for the development of mammals, it is not endogenous stimulation from the nervous system that is of greater importance, but the early development of sensory pathways in it.

L. Carmichael observed the formation of motor activity in guinea pig embryos and established the following patterns. The first manifestations of motor activity are noted on the 28-29th day after fertilization and consist in twitching of the neck-shoulder area of ​​the body of the embryo. Motor reactions reach their maximum development a few days before birth. The embryo develops adequate reflex reactions to tactile stimuli, and these reactions can be modified. For example, a single touch to the area of ​​skin near the ear will cause a reflex twitching of the auricle in the embryo. If, however, such tactile stimuli are repeated many times, then at first the limb will be approached to the place of application of the stimulus, and then (if the stimulus is continued), the head and the entire body will begin to move.

Features of the development of mammalian embryos are also due to the presence of a placenta in them. Thanks to this organ, the development of the embryo is influenced by the maternal organism, primarily in a humoral way (due to the action of biologically active substances, primarily hormones). Experiments were carried out in which female fetuses of guinea pigs were exposed to the male sex hormone - testosterone. This exposure led to a change in their sexual behavior in adulthood: such females showed all the signs of sexual behavior characteristic of male guinea pigs. Interestingly, exposure of guinea pigs to testosterone in the postnatal period (after birth) did not have such an effect on their behavior. Thus, in the embryonic period, sex hormones influence the formation of behavior by influencing the central structures of the nervous system.

Another example of the influence of the maternal organism on the formation of behavioral responses in young mammals can be experiments with inducing a state of stress in pregnant rats. Such females gave birth to shy cubs, which showed such behavioral features, regardless of which female fed them.

The influence of sensory stimulation on the motor activity of the embryo. Despite the fact that motor activity in the embryonic period can be caused by endogenous processes (internal factors), sensory stimulation (exposure to stimuli from the external environment) is also of great importance for its development.

The presence in embryos, along with spontaneous movements (due to internal processes), of reflex movements (in response to external stimulation) was noticed as early as the 1930s. D.V. Orr and W.F. Windle. Already in the early stages of embryogenesis in the chicken embryo, general movements of the whole body are observed in response to tactile stimulation. However, such reactions appear later than spontaneous ones. This is due to the fact that motor pathways in the nervous system of the embryo are formed earlier than sensory (sensitive). Sensory stimulation reaches its greatest development in the last stages of embryonic development. V. Hamburger explains this fact by the fact that the development of behavior includes external factors that prepare the chicks for normal communication with their parents.

For bird embryos, acoustic (sound) contact with parental individuals, which is established immediately before hatching, is of great importance. At this moment, the organs of hearing and vision of the chick begin to function, it can send signals to the external environment, which will be perceived by the parent individuals. At the same time, the chick "learns" to recognize the voices of its parents, to distinguish them from other sound signals. It has been established that for this, the rhythm of the sound signals of the parent and the unhatched chick is coordinated. At the same time, the motor response of hatched chicks to a key stimulus (sound signal) is innate and is combined with embryonic learning. Such prenatal recognition of parental voices is noted in murre, razorbill, geese, waders, and many other birds.

German researcher M. Impekoven conducted experiments with gull chicks. She showed that the acoustic signals that the chicks emit before hatching cause the parents to switch from incubation to caring for the chicks. Conversely, parental individuals emit cries that stimulate the development of pecking movements in chicks, including the "begging" reaction (see Topic 2. Instinct). Thus, prenatal accumulation of experience takes place here.

5.2. The development of the psyche of animals in the early postnatal period

The postnatal period of development of an organism begins after its birth (hatching from an egg). Birth is a turning point in the development of an animal. However, there is continuity between the prenatal and postnatal periods, although after birth, new factors and patterns appear in the development of the organism. The organism is faced with a completely new environment for it. In such acute conditions, the acquisition of individual experience occurs, and the development of innate forms of behavior continues.

In the early postnatal period, the foundations of the behavior of an adult animal are laid, the body acquires the skills to communicate with other individuals, as well as with a changing environment. According to L.A. Orbeli, the early postnatal period is the most sensitive phase of an individual's ontogeny, when the organism actively reacts to all environmental influences.

The postnatal period is very specific. This is especially true for those species of animals in which newborn individuals differ in structure and lifestyle from adult animals. Such differences are observed in most invertebrates, as well as in a number of lower vertebrates that have larval forms (for example, cyclostomes - lampreys, hagfishes). In this case, the postnatal development of behavior is especially complex: on the basis of larval behavior, maturation of a qualitatively different behavior of an adult individual takes place. For example, in marine gastropods, young individuals lead a planktonic lifestyle, but after metamorphosis, adult animals show forms of near-bottom movement and feeding. Somewhat later, in these animals, in a fully formed form, a protective reaction is also manifested in the form of avoiding enemies. Experiments were carried out during which the mollusks were given the opportunity to gain preliminary experience. To do this, they were kept in the water in which the predator used to swim. The results of the experiments showed that in this case there is no accelerated maturation of the protective reaction. Thus, all reactions of the mollusk mature and manifest themselves depending on the development of the corresponding nervous structures.

The degree of maturity of animals is of great importance for the postnatal period. A.N. In this regard, Promptov introduced the concept of “early biostart”. By biostart he understood the moment when biological factors begin to influence the animal. The biostart will be early in an immaturely born baby, which cannot independently provide for its vital functions and depends on the parents for this. On the contrary, a mature-born baby has the ability to independently perform all functions immediately after birth. However, such “complete” maturity is rare; more often it is expressed to one degree or another. For example, fowl chicks need to be warmed by their parents for ten days after hatching, and their movements become coordinated only on the fourth day. But at the same time, from the first moments they can feed independently and exhibit a hiding reaction.

L.A. Orbeli, in his research, drew attention to differences in the development of behavior in mature and immature-born animals. Mature cubs are little affected by the influence of the environment, because they are born in an already formed state. Their conditioned reflex activity is already developed and is subject only to certain additional add-ons and complications. In immaturely born animals, at the time of birth not only are conditioned reflex forms of behavior not formed, but some congenital forms are not even developed. Such cubs are more vulnerable to the influence of the environment, however, according to Orbeli, the development of their behavior is more beneficial. They can adapt the development of behavior to changing environmental factors, so the formation of their behavior is often more adequate to the environment. Orbeli wrote: “These animals will be born with a nervous system so poorly formed that all further postnatal development represents a continuous processing of hereditary forms and newly emerging conditioned forms of behavior.” [19]

It has been noted that animals with the most highly developed psyche are, as a rule, among those born immaturely. They encounter the external environment in a state where their behavior has not yet been formed. The innate foundations of behavioral responses in such animals may be subject to change, so their behavior is more labile. However, the ability of animals of this species to accumulate individual experience is still of decisive importance in this matter. According to these indicators, mature and immature-born animals differ only in terms of acquiring this experience.

Instinctive behavior. In ontogenesis, instinctive movements go through stages of formation and improvement. This fact can be demonstrated experimentally by raising cubs in isolation from the moment of birth. Experiments on birds and rodents have shown that such animals have developed individual motor elements, but the behavioral acts themselves deviate from the norm: the duration, frequency of execution and coordination of behavioral reactions are disrupted. Vital movements are performed, but their coordination with each other is disrupted. Thus, instinctive movements in the behavior of animals in the early postnatal period are certainly present, but they require further development. For example, the American scientist E. Hess conducted experiments with chickens, which immediately after hatching were put on glasses that shifted the field of view by 7 degrees. After a short period of time, these chicks, like the chicks wearing glasses with regular lenses, pecked at the target more accurately than when it was first presented, but the chicks wearing distorting glasses continued to peck 7 degrees away from the target. It follows that the motor reaction associated with pecking movements is innate in birds, but the accuracy of pecking increases through the acquisition of individual experience. Similar data have been obtained for mammals, such as monkeys and guinea pigs.

Thus, the idea that innate behavior plays the greatest role in the early postnatal period is true only in relation to elementary motor reactions. Instinctive acts as a whole require the acquisition of individual experience for their normal formation.

Innate recognition. The importance of innate forms of behavior in the early postnatal period of animal development is primarily manifested in the phenomenon of innate recognition. An example is the nipple-seeking reaction in newborn mammals. They are able to orient their movements according to tactile stimuli, moving towards touch. For example, newborn puppies, when touched on their head, begin to crawl forward, and when touched on their side, they turn towards the stimulus.

The significance of this phenomenon for the life of the animal is very great. To be successful in its life, an animal must immediately after birth be able to orient its behavior in relation to environmental factors. This is especially true for such environmental factors as food, maternal or other parental individuals, other individuals of their species, enemies, etc. K. Fabry wrote: “Innate recognition manifests itself in the innate, species-specific selective attitude of animals to certain components of the environment, signs of objects, situations, in the ability of animals to biologically adequately respond to some signs of objects or situations still unfamiliar to them... we are dealing here with an innate form of orientation, manifested in reactions useful for the individual (and species) to signs of essential components environment without prior learning, with manifestations of “species memory.” [20]

An animal must recognize a biologically significant element of the environment at the first meeting with it and respond adequately to it. Taxis form the basis of innate recognition (see 2.3). Orientation of behavior is carried out according to key stimuli (individual features of biologically significant objects), and the direction of behavior is based on innate triggers. All this in combination ensures a high selectivity of innate recognition.

Along with processes that have an innate basis, early individual experience is of great importance for the behavior of an animal. The acquisition of experience during this period is closely related to the processes of postnatal learning. For example, if a stimulus for which there is innate recognition is often repeated, but has no biological significance for the animal, it gradually "gets used" to this stimulus and ceases to respond to it. Thus, nestlings of brood birds have an innate reaction of hiding to the approach of a predator. Initially, such a reaction follows when any moving object appears in the sky, but gradually the chicks begin to react selectively to objects and do not hide at the sight of a safe stimulus, for example, a leaf, a falling tree. In other words, innate recognition is refined through the acquisition of early individual experience.

In the course of early postnatal learning, the signal value of key stimuli may also change. Thus, in the first days of life, sturgeon juveniles exhibit negative phototaxis, i.e., they swim away from the illuminated areas of the reservoir and try to stay in the shade. However, during the transition to active feeding, conditioned reflexes to light are formed in fish. As a result, the fry show positive phototaxis.

Obligate learning. Sometimes innate recognition can change due to the inclusion of new senses. For example, blackbird chicks respond to nest shaking after hatching by extending their necks upward and opening their beaks. It does not matter what the source of irritation is. After the chicks’ visual organs begin to function, the same reaction begins to appear in them when a parent appears in their field of vision. And only a few days after this, the chicks learn to determine the exact location of the approaching bird and stretch their necks in this direction.

Thus, in addition to innate recognition, obligate learning, i.e., all forms of learning that are vital for an animal under natural conditions, is of great importance for the behavior of animals in the early postnatal period. Obligate learning is close to innate recognition, since it is also specific to a certain species; it forms an integral complex with innate recognition. Obligate learning is characterized by attachment to certain periods of ontogeny. Such periods are called sensitive, or sensible. These periods are usually very short. There are especially many sensitive periods in the early postnatal period, although some of them occur at later stages in the development of behavior.

One of the most important areas of behavior in which obligate learning is of great importance is the formation of eating behavior. First of all, through obligate learning, animals learn to recognize the distinctive features of food objects. If there is no preliminary contact of a newborn animal with an edible object, then in the future the recognition of suitable food for consumption will be difficult. In addition, food acquisition techniques are formed through obligate learning. These include motor reactions that are associated with the capture, capture, dismemberment and consumption of prey. These movements are innate, but without learning they appear in a primitive, imperfect form and must be completed on the basis of individual experience. For example, mongooses have a specific movement that allows them to crack hard-shelled eggs by throwing them under their own body. This is an innate movement; any mongoose is able to do it almost immediately after birth. However, in order for such movements to become synchronous and effective, some time of training and training must take place. The improvement of innate instinctive reactions in lower animals, which do not have a play period in ontogenesis, occurs entirely through obligate learning. In higher animals, there is a special period for such development of behavioral reactions - late postnatal (learning during games).

Obligate learning as the only way to improve innate behavior is characteristic of invertebrates. The observations of the ethologist V.G. Thorp and his staff. They found that if a larval stage insect is exposed to a scent, the adult insect will use that scent as a guide when looking for a place to lay eggs, for example. However, the taxis to normal odors is also preserved in insects. Thus, there is a combination of chemotaxis based on innate recognition (normal smells) and chemotaxis based on obligate learning (smell under experimental conditions).

Optional learning. In the early postnatal period, facultative learning plays a relatively small role; it only serves as an addition to obligate learning.

Experiments were carried out to determine the timing of the formation of the components of facultative learning in cubs of different species. In the course of the experiments, the animal was presented with an artificial stimulus that is not biologically significant for this species, or it was taught actions that were not typical for this species. For example, rat pups at 20 days old can be trained to press a lever to receive a food reward. Approximately at the same time, the abilities for optional learning appear in young carnivorous mammals. It has been established that these abilities depend on the development of short-term memory in them.

In other immature-born animals, facultative learning begins at an earlier date. For example, baby monkeys can develop a conditioned response to sound as early as three or four days of age. At the same time, it is important to remember that the first conditioned reflexes to thermal (temperature) and tactile stimuli begin to form in animals already in the first days after birth, especially for mature animals.

Manipulation. According to K. Fabry, manipulation is “the active handling of various objects with the predominant participation of the forelimbs, less often the hind limbs, as well as other effectors: the jaw apparatus, trunk (in the elephant), grasping tail (in broad-nosed monkeys), tentacles (in cephalopods mollusks), claws (in crayfish), etc." [21]

First of all, the manipulative activity of the animal is manifested in food-procuring and nest-building activity. In these processes, the animal actively interacts with various components of the environment, receives information about the external environment; there is an improvement in the motor reactions of the animal.

Manipulation is the highest form of orienting-exploratory activity of animals. It is fully manifested in the animal in the late postnatal period of ontogeny, however, the timing of the onset of manipulation and its form depend on the type of animal. In this case, the degree of maturity of the animal is of great importance.

In the early postnatal period, manipulation develops only in its simplest form, especially if the animal is immature. For example, puppies, before the organs of vision and hearing begin to work, spend all the time in a dream or in search of a nipple and sucking. Their first movements are manipulative in nature: they crawl, touch their parents and their fellows, make insufficiently clear movements of grabbing the nipple with their mouths, etc. According to the observations of the Soviet zoologist N.N. Meshkova, the fox cub develops manipulative activity of the jaws earlier, and later the motor activity of the forelimbs is formed. Thus, the relationship between various organs that can "replace" each other is clearly manifested.

The main direction of manipulative activity of an immature-born cub in the early postnatal period is the mother's body. Brothers and sisters are passively perceived by the cub, being biologically neutral for it during this period.

Thus, the cognitive value of manipulation in the early postnatal period in immature babies is small. In mature animals, the organs of vision and hearing function from the first hours of life. This allows them to actively interact with the environment.

Imprint. Imprinting is an important moment in the early postnatal period of ontogenesis. It refers to forms of obligate learning, but also includes elements of optional learning.

The first studies of imprinting were carried out by Spaulding. He observed the behavior of chickens in the first days after hatching. The scientist noted that at the age of two or three days, chickens begin to chase any moving object, that is, he first described the phenomenon of imprinting. However, the term "imprinting" and the first detailed descriptions of this phenomenon belong to another ethologist, O. Heinrot (1871-1945, Birds of Central Europe, 1912).

Heinroth conducted research on the behavior of newborn goslings and ducklings, thus laying the foundation for the comparative method in ethology. He noticed that if incubator goslings in an adult state were placed with other birds, and before that they were cared for by a person, such chicks ignored other geese and followed people everywhere. From these observations, Heinroth concluded that for the normal adaptation of the caterpillar to life among relatives, it must be protected from contact with people immediately after birth. For this, the caterpillar after the incubator must be placed in a bag, and then released to the birds. In this case, the chick will not imprint the appearance of a person, and its behavior will not be disturbed.

Heinrot's ideas were expanded and supplemented by the observations of K. Lorenz, who noted such an important quality of imprinting as irreversibility. Lorentz conducted research on the behavior of mallard chicks, pigeons, jackdaws and other bird species. He confirmed Heinroth's opinion that birds that have imprinted the appearance of a person will continue to direct their sexual behavior towards him. As evidence, Lorenz cites an example from the life of an Egyptian dove.

The bird was imprinted on a person, that is, it was imprinted on a person. After that, the dove began to show courtship behavior to the human hand. If the hand was positioned in a certain way, the turtledove made attempts to mate with it. Lorentz noted that the recognition of the object of imprinting has no innate basis, although the behavior itself in relation to the object is hereditarily fixed. So, in the given example, the ritual of courtship as an element of sexual behavior is innate, and the object of courtship depends on imprinting. According to Lorenz, imprinting is tied to a certain period of the animal's life - a sensitive one, and subsequently directs its sexual, "filial" and social behavior.

Parental individuals, other cubs of the litter, future sexual partners can act as objects of imprinting. In this case, typical signs of individuals of the same species are imprinted or, on the contrary, external signs of enemies. In the latter case, the reaction of defense is formed as a result of a combination of these signs and warning cries or other elements of the behavior of the parental individuals. Some scientists note that imprinting may contribute to the formation of a reaction to food objects and habitats characteristic of the species.

Lorentz believed that almost any object can be captured, even if it is very different in appearance from the animal itself. For example, a scientist cites the case of a parrot that captured a ping-pong ball. The adult parrot exhibited in relation to the ball all the same elements of behavior as in the case of a female of its own species. However, in reality, the range of objects that can potentially be imprinted is limited. For example, raven chicks will not show a follow response to a human because they do not have some of the specific features of an adult raven. Such features include the ability to fly and black coloration, possibly also the shape of the body.

The phenomenon of the so-called "multiple imprinting" is very interesting. R. Hind, W.G. Thorpe and T. Wiene describe such an imprint in coot and moorhen chicks. In these birds, at the age of three to six days, several very different objects can be imprinted. At the same time, the following reaction develops not in relation to any one object, but to any of them. But if during the first days of life the chicks did not see a moving object in order to follow it, subsequently the following reaction is disturbed in them. Such chicks run away at the sight of any moving model.

Observations show that certain details of an object, and not its entire appearance, can be imprinted on an animal. For example, observations are known of the behavior of a turkey that was fed by a male zoo attendant. Until the age of one year, this turkey did not see any birds. Already in adulthood, he began to show a sexual command, or rather a reaction of courtship in relation to the caretaker who raised him. It is curious that at the sight of women, as well as men in clothes with fluttering floors, the turkey ran away. Apparently, in addition to imprinting the appearance of the educator, such a reaction was due to the fact that the fluttering clothes evoked an innate defensive reaction in the bird, since it was similar to the pose that a turkey takes when threatened with an attack: it spreads its wings, spreads them on the ground and drags them along. In this example, one can trace the combination of an innate reaction and the imprinting of an unusual object.

Most often, imprinting occurs shortly after birth, while it is confined to a short period of time with clear time limits - sensitive, or sensible. Lorentz believed that the process of imprinting in this case is determined exclusively by internal factors (factors of an endogenous nature), but later it became known that the duration and time of the onset of the sensitive period depended on the life experience of the animal. It has been suggested that these periods are associated with the appearance of new movements in the animal, as well as with the maturation of the organs of vision and some areas of the brain.

Immediately after hatching from the egg, the chicks, as a rule, are not afraid of any objects that are new to them and tend to explore them. However, already after a few days, they begin to show a reaction of fear in such encounters and try to avoid unfamiliar objects. It is interesting that the time of occurrence of such a change in behavior depends on the conditions in which the chicks are kept. It is noted that chickens are less afraid of objects painted in the same colors as the walls of the incubator where they were kept. Thus, in the first days after hatching, when there is still no division into unfamiliar and familiar objects for the chicks, they distinguish any characteristics inherent in the environment from the environment. These characteristics help them to distinguish "familiar" from "unfamiliar". As a result, such a chick can already distinguish familiar objects, while avoiding unfamiliar ones. For example, when chickens are kept together with a hen, very soon both the parent and siblings become familiar objects, and the fear reaction to them does not develop.

The English biologist, anthropologist and philosopher G. Bateson (1904-1980) proposed an interesting model (the Bateson model), which is based on the analogy of the development of an organism with the movement of a train. The initial station from which the movement begins is associated with the moment of conception. Each compartment of this train represents a specific system of behavior. Open compartment windows indicate the sensitivity of behavior to environmental factors at a certain stage of development. At the beginning of the journey, the windows on the train are closed, there is still no connection with the outside world. Then the windows begin to open slightly, passengers can get acquainted with the outside world. The windows can then either close or remain open. At the same time, passengers themselves can change during the journey, and the external environment is constantly changing. Different systems of behavior that are formed in ontogenesis (compartment) can change their essence, their nature, i.e. passengers. These forms of behavior can be programmed to respond to external factors (to get acquainted with the outside world through open windows) at various moments of ontogeny (path).

There may be more than one sensitive period, the animal may go through several variants of the sensitive period. For example, experiments on chickens have shown that the sensitive periods for sexual and "filial" behavior do not coincide in time. Sexual imprinting occurs later. Experiments were carried out in which young cockerels at different ages were shown a moving model. Chickens at the age of 31-45 days, which were presented with such a model, showed sexual behavior towards it, while "filial" behavior was weak. On the contrary, chickens aged 1-30 days, imprinted on the same model, showed a strong "filial" behavior towards it.

K. Lorentz believed that imprinting refers to forms of behavior that are fundamentally different from other forms of learning. Most modern researchers consider imprinting to be a form of learning. Imprinting - teaching the body how it should react to the object that is imprinted. Imprinting is related to forms of perceptual learning. This statement is supported by experiments in which an animal receives a specific experience with the help of specific stimuli. As an example, consider the development of a song in a finch. In order for the song to form normally, it is necessary that the bird listen to it in early ontogenesis, and also have the opportunity to practice this in the later stages of development. The phase when the bird imprints a new song can be thought of as perceptual learning. Another example is the imprinting by zebra finches of the appearance of the bronze finches that raised them. In this case, zebra finches, after several years of isolation, will respond to her as a sexual partner. An example of the involvement of perceptual learning in the process of imprinting can also serve as observations of chickens, which more easily imprint those objects that they previously encountered.

In addition, imprinting is also related to instrumental learning. For example, ducklings were shown a moving toy train a day after hatching. Subsequently, such chicks could be taught to peck at posts if this train passes by immediately after pecking. It is important to note that during the demonstration of the train at the later stages of development, such a reaction was not developed in the ducklings. G. Bateson and K. Reese described observations of ducklings and chickens that can learn to press the pedal to turn on the flickering light. This training is important to take place in a sensitive period for imprinting.

Bateson and Wainwright studied the behavior of chickens in a special device that allowed them to quantify the degree of preference for certain stimuli. They experimentally showed that as the chick becomes familiar with the stimulus imprinted by it, it begins to give preference to other stimuli that are not familiar to it. Scientists have suggested that in natural conditions this helps the chicken to comprehensively study the mother, to get acquainted with all her signs. As a result, based on all the characteristics of the chick, its complex portrait is built.

It cannot be said unequivocally that imprinting is irreversible; it is probably reversible in some animal species. Thus, K. Lorenz gives an example of parrots, in which the appearance of the scientist himself was imprinted. The birds were kept in isolation from people for a long time; they normally mated with individuals of their own species and raised their chicks. However, two years later, finding themselves in the same room with Lorenz, the parrots immediately began to “court” him, abandoning the females of their species. Nikolai notes that a bullfinch, which was raised by a person, behaves with him as with a sexual partner, but in the fall or winter, when he meets a bullfinch of the opposite sex, he can communicate normally with him and does not show any reactions towards the person. However, if a bird does not see individuals of its own species, sexual imprinting on a person remains.

Following reaction. In this reaction, the imprinting manifests itself most clearly. Its essence lies in the fact that the young of mature animals, soon after birth, relentlessly move after their parents and at the same time - after each other. The following reaction is characteristic of both domestic and wild animals. For example, before the chicks hatch, a female goldeneye leaves her nest, which is located in a hollow tree at a height of about 15 m from the ground, and flies away. Upon returning, she, no longer flying into the hollow, emits calling cries, encouraging the chicks to leave the nest. The chicks approach the entrance and then rush down. They land, immediately begin to move actively and follow their mother. The mother waits until the entire brood is on the ground, after which she heads to the pond, the average distance to which is about 2 km. The chicks relentlessly follow her, moving at a fairly high speed. When the birds reach the pond, the mother enters the water and the chicks follow her. The same following reaction is characteristic of other birds. For example, shelducks, which nest in burrows at a height of 3-4 m from the ground, call chicks to them, which jump down to them from this height. Chicks of auks jump from their nesting sites (high cliffs) already at the age of 19 days.

The following reaction is also seen in mammals. It is well expressed in mature animals, especially in ungulates. Their cubs acquire the ability to move in a few hours or even less than an hour after birth. For example, a newborn baby camel already 10 minutes after birth makes the first attempts to stand up, and after 90 minutes it can already stand freely on its feet; the following reaction is formed in him during the day. Imprinting in mammals occurs both on optical and acoustic, and on olfactory signs - the smell of the parent. In mother-isolated cubs, imprinting can occur to their caretaker in captivity if the cub first sees it during the sensitive period. (However, it is believed that other factors besides imprinting underlie the formation of attachment to the mother.) The following reaction is expressed not only in ungulates, it is also well traced in rodents, for example, in mature guinea pigs. The following reaction has also been described in detail in other mammals, such as seals, as well as in fish.

The importance of the formation of the following reaction is great; it is oriented towards the parent individual and other young of the same brood. Thanks to the formation of this reaction, the cubs immediately after birth stay close to the parent, which in such a situation is easier to guide, control and protect them. The cubs learn to distinguish their mother from others and try to keep up with her. Thus, according to K. Fabry, “the rapid concretization of the instinctive behavior of the cubs on individually identifiable objects (parents, brothers) ensures the formation of vitally important adaptive reactions in the shortest possible time.” [22]

Like other cases of imprinting, the following reaction is timed to a sensitive period during which it is formed. For example, goldeneye chicks jump out of the hollow within 12 hours from the time of hatching, these are hours of the sensible period. In chicks of chickens and ducks, the sensitive period begins immediately after hatching and ends after about 10-15 hours. In some animals, this period is longer, for example, in guinea pigs, it stretches from the sixth to the 30-40th day of life. Imprinting occurs very quickly, often one meeting with the object is enough for this.

Reinforcement is not necessary to form a following reaction. E. Hess cites the results of his experiments, when the following of any object in chicks was artificially hampered, for example, by applying painful stimuli. In this case, the reaction not only did not disappear, but, on the contrary, became more intense.

Imprinting is an obligate form of learning, so it does not depend on any components of the environment, even those that could serve as a reinforcement of the reaction. Imprinting is too important for the individual, its life activity, it must be carried out in any conditions, even in the absence of the possibility of reinforcement. However, it is likely that "internal" proprioceptive reinforcement occurs during imprinting. In this case, the source of reinforcement is the sensations from the movements themselves.

Sexual imprinting. Imprinting can influence the choice of sexual partner manifested in an adult animal. This phenomenon is called sexual imprinting. It provides the individual with future communication with a sexual partner.

The difference between sexual imprinting and all other forms of imprinting is that its result appears much later. In this case, the animal learns to recognize the typical distinguishing features of the future sexual partner in the early stages of postnatal development.

Most often, sexual imprinting occurs in males, they "remember" the signs of the maternal individual as a model of an individual of their species. Thus, there is, as it were, a "clarification" of future sexual behavior. At the same time, the recognition of female species-typic characters is superimposed on the innate recognition of common species-typic characters.

Sexual imprinting has been established in various animals, but it is especially pronounced in birds. For example, Warriner and his co-workers conducted experiments with black and white varieties of domestic pigeons. The experiments used 64 pigeons that had not previously mated and were raised by either black or white parents. The results showed that in 26 cases out of 32 males mated with females of the same color as the adoptive parents. In the remaining five out of six cases, females preferred to mate with males that had the coloration of the adoptive parents. Thus, the results of the study showed that the preferences of males are more significant than the preferences of females.

The ethologist F. Schugz showed that in male wild ducks the optimal period of sexual imprinting is limited to 10-40 days. It was at this time that the duck family naturally breaks up. Schutz noted that male ducks choose a sexual partner who resembles the female who raised him in appearance. Females, on the other hand, prefer to mate with males of their own species, regardless of early experience. This has been confirmed experimentally. Of 34 mallard males reared by birds of a different species, 22 mated with females belonging to the species of adoptive parents, and 12 mated with females of their own species. In contrast, out of 18 mallard females reared by other bird species, all but three mated with males of their own species. At the same time, it is noted that males of species with sexual dimorphism (the difference between animals of different sexes in appearance) should rely more on early experience in order to recognize birds of their own species.

Sexual imprinting has also been studied in mammals, especially in ungulates and rodents. Olfactory stimuli play an important role in sexual imprinting. Experiments were carried out on mice: during the experiments they were sprayed with odorous substances. As a result, the cubs of such mice, when they reached sexual maturity, could not distinguish the sex of other individuals, so they did not find a sexual partner. Similar experiments have been done with other rodents, such as rats and guinea pigs. If male rodents are separated from their mothers during the first week of life and fed by individuals of another species, they can observe the effect of sexual imprinting on the alien species.

Sexual imprinting does not always occur in childhood, it can also be observed in adulthood. For example, according to the Swedish ethologists A. Ferne and S. Sjelander, male swordtail fish prefer females of the color that they saw within two months after the onset of maturity.

Thus, in the course of imprinting, there is a rapid postnatal completion of the innate mechanism of behavior due to individually acquired components. Due to this, instinctive behavior is refined, which ensures the effective execution of instinctive actions.

5.3. The development of mental activity in the juvenile (game) period. animal games

In the ontogenesis of higher animals, as a rule, such a period as juvenile or play is clearly distinguished. It is clearly traced in mature-born cubs, in which the maturation of behavior during games is carried out, and this happens long before the onset of puberty.

There are two main concepts to explain the nature of games and their significance in the ontogeny of behavioral responses. The first concept belongs to G. Spencer. Within the framework of this concept, play activity is presented as the consumption of some energy, which under the given conditions is excessive for the body. This energy is not needed to perform the actions necessary to ensure life. In this case, an analogy can be drawn with the so-called "idle actions" (see Topic 2. Instinct). In this situation, some instinctive movements are also carried out in the absence of key stimuli. However, Lorentz himself pointed out a number of significant differences between play activity and idle activities.

The second concept of play activity was formulated by K. Groos. The game is described by him as a kind of exercise of the animal in those areas of activity that are especially important for him, that is, as a kind of practice for the animal. Later, Lloyd-Morgan added that the advantage of learning the animal during the game is that in this case there is an opportunity to make mistakes. No mistake in action will be for the animal either harmful to it or deadly, at the same time, hereditarily fixed actions get the opportunity to improve.

It has now become clear that none of these hypotheses can fully describe the essence of play behavior. Both theories have both supporters and opponents. There is no agreement even on the importance of games for shaping the behavior of an adult animal. As an argument confirming that games have no functional significance for this, scientists cite the fact that normal behavior can be formed even in the absence of exercises in the juvenile period of ontogenesis. For example, the concept of the Dutch zoopsychologist F. Buytendijk is based on the fact that play behavior benefits the animal only in the emotional sphere at the time of the game, while instinctive behavior in any case matures as it is hereditarily fixed, exercises are not needed for this process. However, if the cubs are completely deprived of the opportunity to play in childhood, the psyche of an adult animal in most cases develops in a distorted form. For example, in guinea pigs, reactions to relatives become abnormal, and infantile features are observed in sexual behavior. Coyote puppies in the absence of play behavior in the juvenile period grow aggressive. These features are especially pronounced in monkeys. It is noted that if they are deprived of the opportunity to play with their peers, in adulthood they are not able to communicate normally with sexual partners, as well as perform maternal duties. At the same time, it is important that sexual behavior is formed properly if another animal or person was a partner in the game.

The ideas of another well-known ethologist G. Tembrok are also based on the idea of ​​the game as an autonomous, independent action. However, the scientist emphasizes that play behavior contributes to the fact that the number of behavioral options for an individual in relation to the factors and stimuli of the outside world increases. During the game, elements of learning are carried out, various actions are improved, new systems are formed in the motor sphere of behavior.

Tembrok points out the difference between play activity and "idle movements". Game reactions are quite variable in their manifestations and depend on both external and internal factors. On the contrary, “idle movements” arise under the influence of powerful internal motivation and always manifest themselves within clear limits, that is, they are absolutely unchanged. Tembroke considers games to be a kind of instinctive action with its own motivational mechanism. Like instincts, play actions have a preparatory phase of search behavior and key stimuli. However, unlike instincts, play actions can be performed repeatedly and are often directed to biologically neutral stimuli.

The Swiss scientist G. Hediger fundamentally disagreed with G. Tembrok's hypothesis. He believed that play activity is optional and different from instinct. An animal does not have any special working organs to perform play movements, as is the case with instinctive actions. To prove his assumptions, Hediger cited the results of experiments by the English physiologist V.R. Hess. This scientist, introducing microelectrodes into the cat's brain, did not find any structures responsible for game reactions in the animal.

HELL. Slonim suggested that in the postnatal period, due to the action on the animal's body of external or internal stimuli that do not reach the threshold value, instinctive reactions arise in it. It is this activity that manifests itself in the form of play activity.

Most scientists still hold views on play activity as an exercise in the sensitive and motor spheres, which helps the animal to prepare for adulthood. In this regard, feedback is of great importance. From the motor systems, information is constantly received about the success of the game behavior, it is corrected. Russian psychologist D.B. Elkonin suggested that play activity creates obstacles to the early fixation of instinctive reactions in finished form. This gives the animal the opportunity to orient itself in a changing environment, "tune" the systems of the sense organs and motor systems. V.G. Thorp sees play as an exercise in which the animal acquires useful skills and also expands information about the world. At the same time, according to Thorp, games related to the manipulation of various environmental objects are of particular importance.

The importance of play behavior for the formation and development of the behavior of an adult animal has been proven experimentally. G. Bingham in the 1920s showed that for normal mating of adult chimpanzees in childhood, they need sexual games. According to G. Harlow and S.J. Suomi other games in a similar way help the monkeys develop the ability to herd life.

Games are of such importance not only in monkeys, but also in other mammals. For example, it was noted that for the normal development of reproductive behavior in male minks, it is necessary that the animals receive appropriate play experience with sexually mature females.

D. Nissen together with K.L. Chau and J. Semmes conducted experiments on baby chimpanzees who were deprived of the opportunity to play with objects at an early age. In adults, such animals showed very poor coordination of movements of the forelimbs: chimpanzees could not accurately determine the place of touch with their hands, clumsily felt and took objects. Normal cubs willingly cling to a servant approaching them, but the cubs in the experiment not only did not grab his clothes, but also did not stretch out their hands to him. An important element of chimpanzee behavior - the "search response" - was also not manifested in such cubs.

According to the concept of K. Fabry, gaming activity simultaneously covers many functional areas and is constantly evolving. Fabry points out that “play activity fills the main content of the process of behavioral development in the juvenile period. Play is not represented as some special category of behavior, but as a set of specifically juvenile manifestations of ordinary forms of behavior... Play is a juvenile phase of behavioral development in ontogenesis.” [23]

Thus, in the juvenile period, the main way of behavior formation is games. However, those components of the ontogenesis of behavior that acted at earlier stages do not disappear. In the juvenile period, these factors also remain, but often in a modified form, merging with play activity. The game is carried out on an instinctive basis, it has elements of learning, both optional and obligatory. It is important to note that in the course of play behavior, it is not the adult behavioral acts themselves that are improved as a whole, but their individual components. In the process of play activity, the animal accumulates individual experience, which will be put into practice much later.

Manipulation games - these are games with objects, during which the objects of the environment are manipulated. K. Fabry described the manipulation games of young carnivorous mammals, on the example of which one can see what the game brings to the behavior of an adult animal.

A fox cub, up to twelve days of age, performs manipulative movements with two forelimbs. They are very primitive, they do not involve the jaws and there are no movements that are carried out only by one front paw. The play activity is manifested by the snails after their eyes are opened, at the age of about 16-23 days. After this, the intensive development of the motor sphere of behavior begins abruptly, the number of forms of manipulation increases, and the variety of environmental objects with which manipulation is carried out increases. The cubs have "toys", which can be various objects of the environment. Cubs are very active, mobile.

Fabry describes the typical movements of fox cubs as follows: “pick up an object with the nose (often followed by throwing), holding the object partially or entirely suspended in the teeth (in the first case, the object rests with one end on the substrate), holding the object with the mouth or nose on the forelimbs extended forward, which lie motionless on the substrate (the object rests on them, as if on a stand), raking the object towards oneself with the front paws, pressing the object to the body, lying on its back, while simultaneously biting, pushing and moving along the surface of the body with the nose or forelimbs. In other cases, the object is pressed against the substrate with its limbs, and at the same time part of the object is pulled upward or to the side by the teeth. Burrowing movements and others are often performed." [24]

It is at this age that fox cubs begin to behave in movements associated with manipulations with one limb (touching or pressing objects with one paw, stroking or touching objects with the edge of the hand simultaneously with abducting or adducting movements of the limb, pulling objects towards oneself with a paw with their simultaneous pinching with bent fingers or hooking them to the edges with claws).

Thus, motor activity in the juvenile period is sharply enriched. Primary actions change, due to the completion of construction, new actions are formed on their basis. Qualitative changes in behavioral reactions develop due to the maturation of the motor (motor) and sensory (sensitive) components of this primary manipulation. For example, at the beginning, in early ontogenesis, the reaction of grasping the nipple with the lips develops, and in the juvenile period, the ability to take toys with the mouth is formed on its basis. The primary functions of the mouth apparatus and forelimbs expand and intensify during play movements, that is, play is a developmental activity.

All the regularities described are manifested not only in the sphere of additional functions, but also in the sphere of the main functions of effector systems. This can be clearly seen in the development of food manipulation. The initial consumption of mother's milk requires the development of only one reaction from the cub - sucking. However, over time, food objects change, the sucking reaction can no longer ensure their consumption. The animal must master other, new for itself, forms of action that would allow it to adapt to such changes in food. These movements are formed and improved in the course of manipulation games. For example, a fox begins to lick and then grab various objects with its jaws. Initially, the grasping movements of the jaws served him exclusively for games, and their participation in the process of eating food is associated with a change in the functions of the behavioral reaction.

Manipulative games are observed not only in dogs, but also in other mammals. For example, in the course of play activities, badger cubs develop such actions as digging and carrying soil with the help of their forelimbs, as well as raking bedding material.

The manipulative actions of ungulates are extremely monotonous, because their locomotor apparatus is specialized mainly in support and motor functions, which minimizes the ability to manipulate. Ungulates lack manipulations that are performed jointly by the jaws and limbs, or simultaneously by both forelimbs, but do develop manipulation actions performed by the head or forelimbs, such as pushing objects with the nose and striking.

Manipulative games are developing very well in monkeys. In these animals, the forelimbs do not specialize in any one function, but perform many additional ones. That is why monkeys not only expand the range of possible manipulations, but they also acquire new forms.

Play activity is species-typical. For example, in the games of dingo puppies, actions associated with the pursuit of one individual by others predominate. This is in good agreement with the way of life of adult dingoes, which hunt by driving prey. Fox cubs often jump and hide during games. This is due to species-specific hunting techniques, such as "mouse".

Cooperative games. Often play activities are carried out by several animals at the same time, i.e., they take on the character of joint games. During such games, in addition to the functions already indicated, another very important function is performed - the formation of communication and group behavior of animals. Joint games are games in which there is a coordinated action of at least two partners. Group behavior is not only formed during games, but is also hereditary, i.e. it is instinctive. If an animal is isolated from other individuals from an early age, it will still exhibit some elements of group behavior as an adult.

Joint games can either be manipulative or non-manipulative, i.e., they can be performed in the complete absence of foreign objects. The second option is the most widely used. In joint games, the features of the life of animals of this species are manifested. For example, in guinea pigs, games are very active, they consist mainly of joint jogging and jumping, their games lack fighting techniques, which appear in the ontogeny of behavior only at the onset of puberty. In another species of rodents - marmots - the opposite situation is observed. The young of these animals have a favorite method of play - fighting, pushing and running together as part of the game. However, in general, their games are not as mobile as those of guinea pigs.

Games are very widespread among predatory mammals. In mustelids, for example, they often take on the character of game hunting and subsequent struggle, while the pursued animal changes places with the pursuer. As a result, each individual gets the opportunity to acquire motor skills. In bear cubs, play activity is also manifested in the fight, in addition, the cubs swim and run in a race, and also hide from each other, "rehearsing" and practicing hiding hunting techniques.

In the course of joint games, especially in the course of game struggle, the simplest hierarchical relations between individuals often develop. For the time being, the animals seem to be acquiring the skills of establishing such relationships, but they themselves do not establish direct relationships of subordination. For example, in dogs, the first mutual attacks appear at the age of less than a month, and at 1-1,5 months, subordinate relationships among puppies are already beginning to be established. At the same time, the cubs exhibit aggressive behavior that does not carry a ritualized character - scuttling and jumping on a partner. In contrast to these forms, which have a signal value, ritualized aggression, which serves to establish a hierarchy in adult canids, appears in their behavior much later.

During joint manipulation games, the animals do not communicate directly, because the joint actions of the young in this situation are directed not at each other, but at the objects of the environment. Such games are of great importance for the formation of animal communication, their ability to take joint actions to change the environment. Often joint manipulation games are in the nature of the so-called trophy games. The goal of such a game is to take possession of some object by taking it away from the partners in the game. In trophy games, elements of demonstrative behavior are clearly traced - the possession of an object is demonstrated, in addition, there is a game struggle, a comparison of forces, and the establishment of simple primary hierarchical relationships.

Of great importance in joint games is the coordination of the actions of animals, which is achieved by mutual signaling. Such signaling is innate, it is a kind of key stimulus for play activity, so it is understandable to every animal. Specific postures, movements or sounds can act as signals, they play a stimulating role. For example, canine cubs have a kind of “invitation to play” ritual: the puppy falls on its forelegs, makes sharp jumps to the side, wags its tail, barks briefly in a shrill voice, touches its partner with its front paw, while the corners of its mouth are stretched, ears are directed forward , and longitudinal folds appear on the forehead. The game also includes "appeasement" signals, which should show the partner that the activity is of a playful nature. Otherwise, as sometimes happens in adult animals, the game can turn into a real fight with severe injuries.

Game behavior in the sphere of communication is also characterized by a change of functions. Thus, signals stimulating a partner to play, outside the game situation, have the character of a genuine threat and signal aggressive behavior.

Playing activity is closely related to the exploratory activity of the animal. However, some scientists, such as L. Hamilton and G. Marler, believe that the similarity between the game and exploratory behavior is only external and is not essential. Most likely, research activities during this period are combined with play, during which information about the environment is also collected. Any game has an element of research activity, but research in a young animal does not always take place in the form of a game. The highest form of orienting-research activity is manipulation games with biologically neutral objects.

It is noted that play manipulation is especially intense when the animal is presented with unfamiliar or new objects. It is in such games that the animal actively influences the object. In games that do not have a manipulative nature, such as racing, exploratory activity is minimal. With joint trophy games, we can talk about the general research activity of animals, which is of great importance for the formation of communication.

In the process of individual development, the animal's cognitive and exploratory activity becomes more complex, i.e., the function of this form of behavior expands. After the animal leaves the nest, its exploratory activity is directed to qualitatively different objects, i.e., in addition to expanding functions, they also change.

In various games, the general physical abilities of the animal are developed, for example, eye, strength, dexterity, speed and other qualities. In addition, the elements of behavioral reactions related to nutrition, reproduction and other vital and biologically significant actions are improved, communication skills are formed, and a hierarchy is established.

A special kind of manipulative play can be observed in monkeys. This type of games is characterized by the complexity of the forms in which animals handle objects, and their mobility is low. The animal manipulates objects, remaining in one place for a long time, and its actions are mainly destructive. The animal performs such play actions alone. Such games, according to K. Fabry, should be classified as games of the highest rank. He writes: “During such complex games with objects, highly differentiated and subtle effector abilities (primarily fingers) are improved and a complex of musculoskeletal sensitivity and vision develops. The cognitive aspect here acquires special significance: the animal becomes thoroughly and in-depth acquainted with the properties of the object components of the environment, and Of particular importance is the study of the internal structure of objects of manipulation during their destruction. Of particular importance is the fact that the objects of manipulation are most often “biologically neutral" objects. Thanks to this, the scope of information received is significantly expanded: the animal becomes familiar with components of the environment that are very different in their properties and at the same time acquires a large stock of various potentially useful “knowledge.” [25]

Interesting data were obtained by comparing the play behavior of animals and children. Thus, in some games of young children, it is possible to clearly identify certain components that correspond to the forms of play activity of the young of higher animals. However, already at this stage of ontogenesis, socially determined content can be traced in children’s games. As the child gets older, this feature of games only intensifies, and the game becomes specific to the “human child.” Thus, Russian zoopsychologist A.N. Leontyev wrote that “the specific difference between the play activity of animals and the game, the rudimentary forms of which we first observe in preschool children, is primarily in the fact that the games of the latter represent objective activity. The latter, forming the basis for the child’s awareness of the world of human objects, determines the content child's games." [26]

In the games of children, as in the games of animals, a complex restructuring of connections with factors and stimuli of the external environment is carried out. In the course of ontogenesis, actions in relation to these stimuli also change. In both cases, during the transition from the pre-game period to the game period, motor activity changes dramatically, especially manipulative activity, the methods and objects of manipulation change. However, the formation and development of play activity in children is more complex than in animals, even higher ones.

Topic 6. General characteristics of the animal psyche. Evolution of the psyche

6.1. General characteristics of the mental activity of animals

The evolution of mental activity is an integral part of the process of evolution of the animal world and occurs according to the laws determined by this process. With an increase in the level of organization of animals, their interaction with the outside world becomes more complicated, there is a need for more intensive contacts with an increasing number of subject components of the environment, as well as for improving maneuvering between these components and active handling of them. Only in this case, the balance between the increasing consumption of vital components of the environment and the level of organization of the organism is restored, and more successful avoidance of dangers and unpleasant or harmful effects is carried out. But the process is extremely complex and lengthy, it requires the improvement of orientation in time and space, which is achieved primarily by the progress of mental reflection.

It can be considered that it was the various forms of movement that became the decisive factor in the evolution of the psyche. At the same time, there is an inverse relationship: without the progressive development of the psyche, the motor activity of organisms cannot be improved, biologically adequate motor reactions cannot be carried out, and the further evolutionary development of the organism slows down. The psychic reflection itself does not remain unchanged in the process of evolution, but undergoes profound qualitative transformations. Initially, primitive psychic reflection provided only an escape from unfavorable conditions. Then came the search for conditions favorable to the organism, not directly perceived. Such a search is now a permanent component of developed instinctive behavior.

At higher levels of development, when object perception already exists, and the sensory actions of animals ensure the development of images, mental reflection is able to completely orient and regulate the behavior of animals. First of all, reflection is necessary for an animal to overcome various kinds of obstacles, which is necessary for the emergence of labile forms of individual behavior in changing environmental conditions: in most animals, skills, and in highly developed animals, intellect. The most profound qualitative changes in the psyche in the process of evolution helped to identify several stages of evolutionary development. The clearest line runs between sensory and perceptual psyche.

According to the definition of the Russian zoopsychologist A. N. Leontyev, the elementary sensory psyche is the stage at which the activity of animals “responds to one or another individual influencing property (or a set of individual properties) due to the essential connection of this property with those influences on which the implementation of the main biological functions of animals. Accordingly, the reflection of reality associated with such a structure of activity has the form of sensitivity to individual influencing properties (or a set of properties), the form of an elementary sensation." [27]

Perceptual psyche, as defined by A.N. Leontyev, “is characterized by the ability to reflect external objective reality no longer in the form of individual elementary sensations caused by individual properties or their combination, but in the form of a reflection of things.” [28]

Within the elementary sensory psyche, as well as within the perceptual psyche, one can single out significantly different levels of mental development: the lower and the higher, and also, according to a number of scientists, some intermediate levels. Within large taxa, there are always animals at different stages of mental development, and all the qualities of a higher mental level are always laid down at the previous, lower level.

It should be remembered that innate and acquired behavior do not replace each other on the ladder of evolution, but develop together, as two components of a single process. There is not a single animal in which skills would completely replace all instincts. Progressive development of precisely instinctive, genetically fixed behavior corresponds to progress in the field of individually variable behavior. Instinctive behavior reaches its greatest complexity precisely in higher animals, and this progress entails the development and complication of forms of learning.

6.2. Levels of development of the sensory psyche

The lowest level of mental development characteristic of a fairly large number of animals. Among them, the most typical representatives are the simplest. However, this group also has exceptions. For example, ciliates, as rather highly organized protozoa, have reached a higher level in the development of the elementary sensory psyche than most other protozoa.

The behavior of animals that are at the lowest level of development of the sensory psyche can be extremely diverse, but all manifestations of mental activity in them are still primitive. Mental activity appears in them in connection with the emergence of the ability to feel, to feel. It is the sensation, the reaction to the surrounding world, its factors and stimuli, that is the elementary form of mental reflection, which is inherent in the simplest. These animals actively interact with the environment, react to its changes. It is important to emphasize that protozoa not only show certain reactions to changes in the environment that are biologically significant for them, but also react to biologically insignificant factors. In this case, stimuli that do not directly affect the success of the individual's life activity act as a signal that marks the appearance of changes in the environment that are vital for the simplest.

The lowest level of development of the sensory psyche is preceded by the level of prepsychic reflection, which is characteristic, for example, of plant organisms. At this stage of development, only the processes of irritability are inherent in the body. With the achievement of the lowest level of development of the sensory psyche, the prepsychic reflection in the simplest does not disappear, its elements are preserved. An example is the reaction of protozoa to such a vital component of the environment as the temperature regime. In this case, one can also talk about the identity of a vital factor and a factor that acts as an indirect signal about the presence of an important environmental factor. Protozoa do not have specific thermoreceptors responsible for the body's perception of the temperature regime. However, it has long been proven that they show reactions to temperature changes, and quite differentiated ones. So, at the beginning of the 24th century. M. Mendelssohn drew attention to the fact that the responses to temperature changes in ciliates, when approaching a certain thermal optimum, become more and more differentiated. For example, for ciliates-shoes, the optimal water temperature is 28-6 ° C. At a temperature of 15 to 0,06 ° C, the shoe reacts to a temperature difference of 0,08 to 20 ° C, and at 24-0,02 ° C, to a difference of 0,005 to XNUMX ° C. G. Jennings suggested that the sensitivity of ciliates-shoes to changes in temperature is associated with increased sensitivity to this factor of the anterior end of the protozoan body. However, experiments with cutting ciliates into two parts across the body showed that both halves of the body show the same response to temperature fluctuations. It is possible that the reaction of such protozoa to the temperature regime is determined by the properties of the entire protoplasm of the animal. In this case, the reactions can be similar to biochemical reactions, for example, with enzymatic processes. Thus, in protozoa, along with mental reflection, prepsychic reflection continues to exist, and this is characteristic of both highly organized representatives of the type (ciliates) and low-developed ones (for example, euglena).

Mental reflection and its qualities are determined by the degree of development of the animal's ability to move, as well as to orientation in space and time, to change innate behavior.

The modes of movement of protozoa are extremely diverse. So, they can passively soar in the water column, or they can actively move. This group of animals has specific modes of movement that are absent in multicellular organisms. Examples are movement by moving protoplasm and forming pseudopodia (typical for amoeba), as well as the "reactive" method of locomotion - mucus is released from the posterior end of the body under high pressure, which pushes the animal forward (typical of gregarines). In addition, the protozoa may have specialized structures for movement - cilia and flagella. These motor structures are plasma outgrowths that perform rotational, oscillatory and wave-like movements, and cilia are a more complex effector apparatus than flagella. Due to the specialization of the ciliary apparatus (the formation of an accumulation and fusion of several cilia, their grouping in certain areas of the body), the movements of protozoa can become more complex. For example, infusoria of the genus Stilonychia, along with swimming, can move along the bottom, while changing the direction of movement.

The motor apparatus of most protozoa is represented by myonemes - fibers consisting of myofibrils. Myonemas are located in the organism of the simplest in the form of rings, longitudinal threads or ribbons. They can have both a homogeneous (homogeneous) structure and transverse striation. Myonemes enable the simplest animals to carry out body contractions, as well as more complex specialized locomotor and non-locomotor movements. Myonemas are absent in such protozoa as amoeba, rhizopods, the vast majority of sporozoans, etc. These protozoa move due to contractile processes in the cytoplasm.

All forms of protozoan motor activity are at the level of instinctive behavior - kinesis (see also 2.3). At the same time, behavioral reactions are carried out in the form of positive or negative taxis that arise on the basis of sensation and allow the animal to adequately respond to environmental conditions - to avoid adverse conditions and move towards the action of positive and biologically favorable ones. The instinctive behavior of the protozoa is still very primitive, since it either lacks the exploratory phase or this phase is very poorly developed. Mental reflection at this stage is also extremely poor in content, since its content is determined by an active search and evaluation of stimuli in the search phase. Search behavior in protozoa exists at an embryonic stage. For example, predatory ciliates are capable of actively searching for prey. However, in general, it can be noted that at the lowest level of the sensory psyche, only, as a rule, negative components of the environment are recognized at a distance. Biologically neutral factors do not yet have a signal value, therefore they are not perceived by animals at a distance. It can be said that mental reflection at this level of development of the psyche performs exclusively the role of a "watchman": biologically insignificant components of the environment are perceived by the body only if they are accompanied by negative biologically significant components.

In the behavior of protozoa, integration in the motor and sensory spheres can be noted. An example is the phenomenon of a phobic reaction (fear reaction) in protozoa, for example, in Euglena. The simplest, having encountered an obstacle, stops and makes circular movements with the front end of the body. Then the euglena swims away in the opposite direction to the obstacle. Such integration can be carried out with the help of special functional structures, which would be similar to the nervous system of multicellular organisms. For the simplest, such structures were found only in ciliates. Perhaps, in addition to this, a system of gradients in the protoplasm is involved in the conduction of nerve impulses.

The simplest have a weakly expressed ability to learn. For example, if an infusoria floated along the walls in a triangular vessel for a long time, it retains such a trajectory of movement in a vessel of a different shape. As a result of N.A. Tushmalova discovered phenomena in the behavior of ciliates, which the researcher interpreted as examples of elementary trace reactions. So, ciliates, which were subjected to rhythmic vibration for a long time, initially reacted to this factor with a contraction, and after a while they ceased to show a reaction. Tushmalova suggested that such trace reactions represent the simplest form of short-term memory, which was formed on the basis of molecular interactions. The question of whether such a change in behavior is the simplest form of learning has been discussed by many scientists. Probably, in this case, such an elementary form of learning as habituation takes place. At the lowest level of development of the sensory psyche, addiction is based solely on sensations: the animal gets used to the effects of specific stimuli, which embody specific properties of the environment. At the same time, species-typical instinctive reactions cease to manifest themselves in the animal if their repetition does not produce a biologically significant effect.

Addiction in appearance is very similar to fatigue. In contrast to the latter, habituation is associated not with the waste of energy reserves, but rather with their saving, with the prevention of energy expenditure on the implementation of movements that are biologically useless for the animal. In experiments with ciliates, fatigue manifested itself in the fact that after the animal was irritated by strong stimuli for several hours, it completely ceased to respond to stimuli.

In highly developed representatives of protozoa, in addition to habituation, the level of development of the sensory psyche is also characterized by the beginnings of associative learning. In this case, temporary connections are established between a biologically significant stimulus and a biologically neutral stimulus. For example, in the experiments of the Polish scientist S. Vavrzhinchik, ciliates were taught to avoid swimming into a darkened area of ​​a glass tube with water, in which they were irritated by an electric current. Gradually, the protozoa stopped swimming into the shade even in the absence of electric shocks for 50 minutes. Such experiments were subsequently carried out by another Polish researcher, J. Dembowski, who suggested that in this case one could rather talk about the development of primitive conditioned reactions in ciliates, which is controversial.

As evidence of the ability of ciliates to associative learning, the results of experiments with placing ciliates in capillaries with a bent end were considered. A protozoan was placed at this end of the capillary, and then the time it took for the ciliates to exit it was recorded. It was noted that with repetition of the experiment, this time was significantly reduced. However, later F.B. Applewhite and F.T. Gardner repeated these experiments, and after each experiment, the capillary was thoroughly washed. In this case, the exit time after each repetition of the experiment did not decrease. The scientists concluded that the decrease in the exit time is associated not with the ability of ciliates to associative learning, but with their orientation in the capillary according to the metabolic products accumulated there.

In general, we can say that the behavior of the simplest is weakly plastic, because it is almost completely determined by instinctive components, and the possibility of modification lies in the phenomenon of habituation, which cannot yet be called a full-fledged form of learning. Habituation fully provides the lability of behavioral reactions necessary for the simplest. The habitat of the protozoa is quite stable, the accumulation of individual experience is not so important for them, because the life span of the protozoa is extremely short.

The highest level of development of the elementary sensory psyche achieved by most multicellular invertebrates. However, some of them (sponges, most coelenterates and lower worms) are an exception in this respect, their sensory psyche is comparable in terms of its level of development with the mental development of protozoa. Nevertheless, in general, for all multicellular invertebrates, fundamental changes in behavior can be noted due to the emergence of a special system for coordinating tissues, organs, and organ systems - the nervous system. In this case, first of all, the speed of conducting nerve impulses increases significantly: if in the protoplasm of the simplest it does not exceed 1-2 microns / s, then already in the primitive nervous system, which has a cellular structure, it increases to a speed of 0,5 m / s. The nervous system of lower multicellular organisms can have a different structure: reticulate (hydra), ring (jellyfish), radial (starfish) and bilateral.

In the process of phylogenetic development, the nervous system was immersed in the muscle tissue, and the longitudinal nerve cords became more and more pronounced, the process of cephalization of the nervous system was observed (the appearance of a separate head end of the body, and with it the accumulation and subsequent compaction of nerve structures in the head). In higher worms (annelids), the nervous system takes the form of a "nervous ladder". Their brain is located above the digestive tract at the anterior end of the body, there is a near-pharyngeal nerve ring and paired abdominal nerve trunks with symmetrically located nerve ganglia connected by transverse cords. It is in annelids that the signs of the highest level of the elementary sensory psyche are fully expressed. It is important to note that the level of mental development is determined not only by the development of the nervous system, but also by the complexity of the conditions for the existence of the organism.

The behavior of annelids (annelids) still does not go beyond the boundaries of the elementary sensory psyche, because it is composed of movements oriented only according to individual properties of objects based only on sensations. The abilities for objective perception, i.e., for perception, are still absent in the rings. It is possible that the beginnings of such abilities first appear in free-swimming predatory mollusks, as well as in some polychaetes. For example, a terrestrial mollusk may begin to bypass an obstacle even before it comes into direct tactile contact. However, such abilities of the mollusk are also limited: it does not react in this way either to small objects or to too large ones, the image of which occupies the entire retina.

As in the case of protozoa, avoidance of unfavorable environmental factors is of paramount importance in the behavior of lower multicellular animals. However, they also have signs of a higher level of sensory psyche, i.e., they are actively looking for positive stimuli. In the behavior of these invertebrates, along with kinesis and elementary taxises, there are the beginnings of complex forms of instinctive behavior (especially in some polychaetes, leeches, and also gastropods) and higher taxises appear. Higher taxises provide an increase in the accuracy and efficiency of the orientation of the animal in space, as well as the full use of trophic resources. The higher taxises include tropotaxis, telotaxis, menotaxis and mnemotaxis (for details on them, see 2.3, pp. 51-52).

In the behavior of the higher representatives of the group of multicellular invertebrates, a number of elements are noted that are characteristic of the behavior of more highly organized animals. In polychaetes, unlike other invertebrates, there are complications of species-typical innate behavior that already go beyond the elementary sensory psyche. Thus, marine polychaetes are able to carry out constructive actions, which are expressed in the fact that the worms actively collect material for future structures with the help of bristles, and then actively work on building "houses" from it. The construction process is a complex action consisting of several successive phases that can change, adapting the process to external environmental factors. For example, the structure of a house may change depending on the nature of the soil and the speed of the current, the topography of the bottom, the number of particles sinking to the bottom and their composition, and the material for construction may also change. Polychaete is actively looking for material for construction, and selects it according to size. For example, young worms choose smaller diameter granules for this purpose, while older animals prefer large particles.

In polychaetes, the beginnings of mating behavior and aggression are outlined, which means that communication appears. True mating behavior and aggression begin to develop only at the lowest level of the perceptual psyche (in arthropods and cephalopods) and are characterized by a certain degree of ritualization. However, even in polychaetes (in particular, in the sea worm Nereid), one can observe the struggle for the right to own a house. During such "battles" the animals usually do not cause severe damage to each other, but they bite and can drive the individual out of the house. At the same time, ritualization of behavior and any signaling are completely absent. The aggressive behavior of a polychaete male towards another male during pair formation was noted by SM. Evans and co-workers on Harmothoe imbricata. Mating behavior has been noted in gastropods and polychaetes. So, in grape snails, direct mating is preceded by long "nuptial dances", during which the partners prick each other with the so-called "love arrows" - lime needles. Thus, the higher forms of behavior appear in a primitive and rudimentary form even at the lower stages of the development of the psyche.

The nervous system of the lower multicellular organisms is still very primitive. Its primary and main function is the internal coordination of all vital processes of the organism. This becomes necessary in connection with the developed multicellular structure, the emergence of new structures that must function in concert, the "external" functions of the nervous system are "secondary" for it. They are determined by the degree of external activity of the animal, which is still very weak and rarely surpasses the activity of protozoa. Therefore, the "external" activity of the nervous system, as well as the structure and functions of its receptors, are significantly developed in invertebrates that lead an active lifestyle. As a rule, these are free-living forms capable of active movement in the environment.

The plasticity of the behavior of lower multicellular organisms, including annelids, still remains poorly expressed. Behavior is dominated by instinctive components, stereotyped reactions. Practically no individual experience is accumulated, and learning in these invertebrates is extremely weakly expressed. Its results are not able to persist for a long time, and it takes a long time to build associative links.

All rings are characterized by habituation: after repeated exposure to a stimulus that is not accompanied by a biologically significant effect, the innate species-typical reaction of the animal to this stimulus is lost. For example, earthworms, after repeated shading without adverse effects for them, cease to respond to this phenomenon by the desire to crawl away to a lighted place. Habituation is observed not only in physical activity, but also in the field of eating behavior. For example, experiments were carried out with predatory annelids, which were given pieces of paper soaked in the juice of the victim of the ring. Initially, the worm ate the offered paper several times, but after a series of repetitions it stopped accepting it. The experiment was complicated: the ring was given paper and a real victim alternately, in this case, after numerous repetitions, the worm learned to distinguish between objects, eating food and rejecting paper with the smell of the victim. The same experiments were carried out on animals with the lowest level of elementary sensory psyche (intestinal polyps). After several similar repetitions, the polyps also began to reject inedible objects even before they came into contact with the mouth opening. Thus, lower invertebrates have abilities that allow them to distinguish an edible object from an inedible object by secondary physical qualities. Note that the taste qualities (direct physical qualities) of both objects were the same. When determining the suitability for food of the proposed object, the animal is guided by its specific property. This property acts as a signal, and the sensitivity of the animal acts as an intermediary between the vital component of the environment and the organism itself. This indicates that already at the lowest level of development in animals a psychic reflection appears in its true form.

In flatworms (and more highly developed worms), learning through “trial and error” is manifested in a rudimentary form, as well as the formation of individual motor reactions. For example, if you put a strip of sandpaper in the path of a milk planaria, it will pause, but then crawl through the paper. If you shake the surface of the table while crawling, the worm will stop crawling through the paper even if the shaking is not happening at the moment. In this case, however, there is still no real, true association of the two stimuli, i.e., paper roughness and surface shaking. This effect is explained by a general increase in the excitability of the animal, which occurs as a result of a combination of two negative stimuli.

Planarians can also develop complex reactions to two stimuli, one of which is biologically neutral for the animal. For example, L.G. Voronin (1908-1983) and N.A. Tushmalov developed defensive and food conditioned reflexes in flatworms (milk planaria) and annelids. The conditioned reflexes of planarians were extremely primitive and did not persist for a long time, while in polychaetes they could independently recover after extinction and had sufficient stability. This testifies to the progressive phylogenetic development of the mental activity of animals (in particular, worms), which is accompanied by a complication of the morphological, anatomical, and functional features of the nervous system.

The plasticity of the behavior of oligochaetes (low bristle worms) was studied as early as the beginning of the 120th century. American zoopsychologist R. Yerks. He noted that in order to teach the earthworm to find a "nest" in the T-shaped maze, and to avoid electric shock at the other dead-end end of the maze, the experiment must be repeated for 180-XNUMX times. Worms can be retrained by swapping the dead ends of the labyrinth with current and "nest". Such experiments were also carried out with worms in which the anterior segments of the body were removed; in this case, the results of learning did not change. V.A. Wagner concluded that in annelids the ganglia of each segment of the body are capable of autonomous work to ensure the performance of elementary mental functions. The process of cephalization in oligochaetes has not yet reached such a development as to determine the behavior of the animal, however, already at this stage of development, the brain has a guiding effect on behavioral acts. So, if the earthworm is cut across the body, its rear end will not be able to move purposefully, while the front end will dig into the ground.

The associative links of polychaetes are much more pronounced. For example, experiments were carried out to change the sign of the behavioral response of polychaetes to lighting. Under normal conditions, it is negative, but with repeated combination with food reinforcement, it can be rebuilt into a positive one. In this case, when the house is illuminated, the polychaete does not hide in its depths, but, on the contrary, actively crawls out of the shelter.

6.3. Perceptual psyche. The problem of intelligence in animals

The lowest level of development of the perceptual psyche. The perceptual psyche is the highest stage of development of mental reflection. This stage of mental development is already characterized by the presence of genuine skills and perceptions. The components of the environment are reflected by the organism as integral units, whereas at the previous level of development only individual properties or the sum of the objective components of the environment were reflected. It is at this stage of mental development that sensory ideas appear. The perceptual psyche itself, which is observed in many living organisms, reveals great differences. Therefore, it became necessary to carry out a more detailed classification, according to which the first level of development of the perceptual psyche is called the lowest.

The lowest level of development of the perceptual psyche is characteristic primarily of higher invertebrates - cephalopods and arthropods. Among arthropods, the characterization of this level of mental development is best considered using the example of insects, the most numerous class of arthropods.

A specific lifestyle, various forms of motor activity, and a variety of qualitatively different environmental agents that control behavior determined the development of numerous and peculiarly arranged sense organs in insects. Among them, the most important is the visual apparatus, since it was well-developed vision that contributed to the optical perception of forms as a necessary component of the perceptual psyche. It should be remembered that at the level of the elementary sensory psyche it is still impossible for animals to distinguish between forms.

Until recently, it was believed that insects are capable of perceiving form, but only within specific limits. In the first experiments, it was shown that bees can perceive only those objects that remotely resemble a flower in their structure (circles, stars). But later, in the experiments of the Soviet zoologist Mazokhin-Porshnyakov, it was proved that bees can initially be trained to perceive shapes that are unusual for them, such as a triangle or a circle, as a result of which it was concluded that bees are able to recognize figures directly by their graphic features.

Similar experiments on single wasps were carried out by N. Tinbergen, one of the founders of modern ethology. He trained female wasps to recognize a circle of pine cones laid out around the entrance to a burrow. After the wasp flew away for prey, the circle moved 30 cm to the side. Returning, the wasp first looked for a hole in the center of the circle. In the following experiments (in addition to moving the circle), the cones were replaced with black pebbles, and a triangle or even an ellipse was built around the mink from these pebbles, but the wasp nevertheless flew into the circle, although it was known from previous experiments that it was quite capable of distinguishing pebbles from cones. Thus, spatial orientation was carried out here only according to the shape (circle).

The ability for object perception in higher insects is noticeably lower than in vertebrates, which can be explained by the specific structure of the organs of vision. In addition, insects are more oriented not by the subject components of the environment, but by their individual features, which is more typical for the elementary sensory psyche stage.

Perhaps more important than in insects, vision also plays in cephalopods. For them, vision is the leading reception, as indicated by the complex structure and large size of the eyes. The relative sizes of the eyes of squids exceed the relative sizes of the eyes of most aquatic mammals (whales, dolphins) by tens of times. The huge resolving power (vigilance) of the cephalopod eye is also striking: for 1 mm2, different representatives of cephalopods have from 40 to 162 thousand sticks, in humans - 120-400 thousand, in an owl with the most keen eye in the world - 680 thousand.

Cephalopods are capable of genuine object perception, which is expressed primarily in their discrimination of the shape of objects. This was proved in the experiments of B.B. Boycott and J. Z. Young. It turned out that octopuses can not only perceive the shape of objects, but also distinguish their relative size, as well as their position in space (for example, they distinguished a vertical rectangle from a horizontal one). In total, these cephalopods distinguished more than 46 different forms.

In higher invertebrates, the rudiments of communication already appear, which is especially developed in animals leading a group lifestyle (bees, ants). It was these insects that had the opportunity to transmit information using special signal actions. Very pronounced in invertebrates and territorial behavior. Its beginnings can be found already in earthworms. In higher invertebrates, the marking of an individual site, a peculiar combination of territorial behavior and information transfer, are well expressed.

Already at the lowest level of development of the perceptual psyche, all those progressive features that characterize the perceptual psyche in general are present, but in many respects the behavior of animals belonging to this category also bears primitive features that bring it closer to the behavior of lower animals. Behavior is still focused on individual properties of objects, object perception is poorly expressed. The behavior is dominated by hard-coded elements and has very little flexibility. At the same time, at this level of development of the psyche, a clearly expressed active search for positive stimuli appears, and taxis behavior develops powerfully. There are all kinds of higher taxis, including mnemotaxis. It is mnemotaxis that play an important role in spatial orientation, and in memorizing landmarks, the ability to change behavior, i.e., to learn, is already manifested.

Although in invertebrates, in particular insects, the accumulation of individual experience and learning play a significant role, there are also certain inconsistencies in learning processes, a combination of progressive and primitive features. The transitional stage between instinctive behavior and true learning is clearly visible, which places this level of development of the psyche between the elementary sensory and the developed perceptual psyche.

Instinctive behavior itself is represented by already developed new categories, such as group behavior, communication. At the present stage of the development of science, the language of bees has been best studied, it has been proved that complex forms of communication are well developed in these insects. The most complex forms of instinctive behavior are naturally combined in them with the most diverse and complex manifestations of learning, which ensures not only the exceptional coordination of the actions of all members of the bee colony, but also the maximum plasticity of the individual's behavior. The psychic abilities of bees (as well as some other higher insects) in some respects, obviously, already go beyond the lower level of the perceptual psyche.

At the lowest level of the perceptual psyche, there are also a number of representatives of the lower vertebrates. The main reason for this is their relatively small size. All invertebrates live in conditions (temperature, lighting) that are fundamentally different from those of large vertebrates. For this reason alone, the psychic reflection of reality in insects, like in most other invertebrates, cannot but be fundamentally different from that of vertebrates. According to the general signs of mental reflection characteristic of this level, we can conclude that insects have a typical manifestation of the lower level of the perceptual psyche, but in forms that correspond to those special conditions of life of these animals, which were mentioned above.

The highest level of development of the perceptual psyche. It has been proven that during the evolutionary process in the animal world, three separate peaks were formed: vertebrates, insects and cephalopods. All these groups dissociated themselves from the common evolutionary trunk quite early and independently reached the heights of development. It is in these animals that the most complex forms of behavior and mental reflection are observed, due to the high development of the level of structure and vital activity. Representatives of all these groups are capable of object perception, but only in vertebrates it has received full development. It is not surprising that only vertebrates, and even then not all representatives of this type, reached the highest level of development of the perceptual psyche in the course of evolution. Only in higher vertebrates are all the most complex manifestations of mental activity found in the animal world.

The high development of the mental activity of vertebrates is directly related to the complication of their organization, the variety of movements, the complication of the structure of the nervous system and sensory organs. All the main manifestations of mental activity characteristic of animals, described in other sections of the book, are characteristic of vertebrates. Let's consider the most important of these manifestations.

The first is manipulation. The limbs of animals, which initially performed only supporting and locomotor functions, acquired a number of additional functions as they developed, one of which is manipulation. For a zoopsychologist, of particular interest is the manipulation of the forelimbs, which ultimately led to the emergence of tool activity in primates and served as a biological prerequisite for the emergence of labor actions in ancient people. Manipulation is characteristic mainly of primates, much less often it is observed in representatives of other orders of mammals. When manipulating the animal comprehensively gets acquainted with the object, learns more about its properties. Under appropriate conditions, animals receive the most comprehensive and varied information necessary for the development of higher forms of mental activity. It turned out that bears have three ways of fixing an object on weight, raccoons - six, lower monkeys and semi-monkeys - three dozen such ways! In addition, only monkeys have different motor capabilities sufficient to produce a genuine destructive analysis (dismemberment) of an object in weight. A variety of manipulation is also comfortable behavior, which is well developed in many higher vertebrates.

At this stage in the development of the perceptual psyche, visual generalizations and the formation of representations also developed. It is known that the true perception of the subject components of the environment is possible only on the basis of the ability to analyze and generalize, since only in this way can constantly changing components of the environment be recognized. All vertebrates, starting with fish, are capable of object perception, in particular, of the perception of forms. Higher vertebrates are capable of generalization, that is, in experiments they recognize an object if it has not only changed its place, but also changed its position in space. For example, mammals can quickly recognize triangles of various sizes and orientations in a plane. With appropriate learning, higher vertebrates are able, even in very difficult situations, to isolate essential details in perceived objects and recognize these objects in a greatly altered form. This leads to the conclusion that vertebrates have rather complex general ideas.

The presence in vertebrates of representations expressed in delayed reactions and the ability to find detours (including extrapolation phenomena) gives their behavior exceptional flexibility and greatly increases the efficiency of their actions at the search stages of behavioral acts. However, the ability to generalize does not indicate a high level of mental development of the organism. This ability is primarily a prerequisite for the development of complex skills, which constitute the main content of the accumulation of individual experience not only in the sensory, but also in the effector sphere of the body's activity.

In higher vertebrates, the processes of communication are noticeably more complicated. They have a very diverse means of communication, which include elements of various modalities, such as olfactory, tactile. They inherited olfactory communication from territorial behavior, when animals actively marked the boundaries of their own territories.

The components of the instinctive behavior of vertebrates that serve for communication are ritualized to one degree or another. Optical communication is carried out with the help of characteristic postures, body movements, which are noticeably simplified and have a clear sequence of actions. First of all, they serve for the biological differentiation of species and are more pronounced in closely related species. The specific forms of optical communication in higher vertebrates are very diverse and differentiated. In mammals, optical communication is often combined with olfactory communication, and the allocation of communication systems according to individual modalities in these animals is largely arbitrary. To some extent, this also applies to acoustic signals, which in mammals are often accompanied by characteristic postures. The most developed sound signaling in birds, it covers almost all spheres of their life. Of great importance are not only clear interspecies differences in acoustic communication, but also individual differences, by which individuals recognize each other.

Thus, it can be said that at the highest level of development of the perceptual psyche, all the basic forms of animal behavior are formed, and the more ancient of these forms, which arose in the early stages of the evolution of the psyche, reach their highest development.

Complex skills are exclusively dynamic motor-receptor systems that ensure the development of very plastic motor programs on the basis of highly developed orienting activity. In higher animals, the orienting process merges with motor activity, and correct decisions are made in changing environmental conditions on the basis of a highly developed sensory generalization. Such complex skills, characteristic of higher vertebrates, have become prerequisites for the development of higher forms of animal mental activity - intellectual actions.

The problem of animal intelligence. It is generally accepted that intellectual behavior is the pinnacle of mental development in animals. Numerous experiments have proven that intellectual activity is characteristic only of higher vertebrates, but, in turn, is not limited to primates. It should be remembered that the intellectual behavior of animals is not something isolated, out of the ordinary, it is only one of the manifestations of a single mental activity with its innate and acquired aspects. According to K. Fabry, “...intellectual behavior is not only closely connected with various forms of instinctive behavior and learning, but is itself composed (on an innate basis) of individually variable components of behavior. It is the highest result and manifestation of individual accumulation of experience ", a special category of learning with its inherent qualitative features. Therefore, intellectual behavior gives the greatest adaptive effect... with sudden, rapid changes in the environment." [29]

The main prerequisite for the development of intelligence is manipulation. First of all, this applies to monkeys, for whom this process serves as a source of the most complete information about the properties and structure of the objective components of the environment. In the course of manipulation, especially when performing complex manipulations, the experience of the animal's activity is generalized, generalized knowledge about the subject components of the environment is formed, and it is this generalized motor-sensory experience that forms the main basis of the intelligence of monkeys. During manipulation, the animal receives information simultaneously through a number of sensory channels, but in monkeys, the combination of skin-muscular sensitivity of the hands with visual sensations is predominant. In addition, the examination of the object of manipulation involves smell, taste, tactile sensitivity of the perioral vibrissae, and sometimes hearing. Animals receive complex information about the object as a single entity with properties of different quality. This is precisely the meaning of manipulation as the basis of intellectual behavior.

Of primary importance for intellectual behavior are visual generalizations, which are also well represented in higher vertebrates. According to experimental data, in addition to primates, visual generalization is well developed in rats, some predatory mammals, and among birds - in corvids. In these animals, visual generalization is often close to the abstraction characteristic of mental processes.

Another element of intellectual behavior, directed to the motor sphere, is studied in detail in vertebrates using the problem box method. Animals are forced to solve complex subject problems, find the sequence of unlocking various locks and valves in order to get out of the cage or get to the treat. It has been proven that higher vertebrates solve objective tasks much worse than tasks based on the use of locomotor functions. This can be explained by the fact that the mental activity of animals is dominated by the cognition of spatial relations, comprehended by them with the help of locomotor actions. Only in monkeys and some other mammals, due to the development of manipulative activity, locomotor actions cease to dominate, animals abstract more easily and, accordingly, solve objective problems better.

An important prerequisite for intellectual behavior, according to K. Fabry, is the ability to widely transfer skills to new situations. This ability is fully developed in higher vertebrates, although it manifests itself in different animals to varying degrees. The main laboratory experiments in this direction were carried out on monkeys, dogs and rats. According to K. Fabry, “the abilities of higher vertebrates for various manipulations, broad sensory (visual) generalization, for solving complex problems and transferring complex skills to new situations, for full orientation and adequate response in a new environment based on previous experience are the most important elements of intelligence animals. And yet, these qualities in themselves are not yet sufficient to serve as criteria for the intelligence and thinking of animals." [thirty]

What are the main criteria for the intellectual behavior of animals? One of the main features of the intellect is that during this activity, in addition to the usual reflection of objects, there is also a reflection of their relations and connections. In its rudimentary forms, this was presented during the formation of complex skills. Any intellectual action consists of at least two phases: the action preparation phase and the action implementation phase. It is the presence of the preparation phase that is a characteristic feature of intellectual action. According to A.N. Leontiev, the intellect first appears where the process of preparing the possibility to carry out this or that operation or skill arises.

In the course of the experiment, it is possible to clearly distinguish between the main phases of intellectual action. For example, a monkey takes a stick and in the next moment with its help pushes a banana towards him, or he first builds a pyramid from empty boxes in order to pluck a bait suspended from the ceiling from a rope. N.N. Ladygina-Kots studied in detail in chimpanzees the process of preparing and even manufacturing tools needed to solve a technically simple task - pushing a bait out of a narrow tube. Before the eyes of the chimpanzee, the bait was placed in the pipe in such a way that it could not be reached simply with the fingers. Simultaneously with the tube, the animal was given various objects suitable for pushing food. After some improvement was made in the object used to get food, the experimental monkey completely (although not always immediately) coped with all the tasks assigned.

In all these experiments, two phases of intellectual action are clearly visible: the first, preparatory phase - preparing the tool, the second phase - getting the bait with the help of this tool. The first phase, out of connection with the next phase, is devoid of any biological meaning whatsoever. The second phase - the phase of the implementation of activities - as a whole is aimed at satisfying a certain biological need of the animal (in the described experiments - food).

Another important criterion of intellectual behavior is the fact that when solving a problem, the animal does not use one stereotypically performed method, but tries different methods that are the result of previously accumulated experience. Animals try to perform not different actions, but different operations, and in the end they can solve the problem in different ways. For example, you can build a pyramid out of boxes to pick a hanging banana, or you can take the box apart and try to knock down the delicacy with separate planks. The operation ceases to be fixedly connected with the activity that meets a specific task. This is what intelligence is noticeably different from any, even the most complex, skills. Since the intellectual behavior of animals is characterized by a reflection not just of the objective components of the environment, but reflects the relationship between them, here the transfer of the operation is carried out not only according to the principle of similarity of things (for example, barriers) with which it was associated, but also according to the principle of similarity of relations, connections. things she responds to.

Despite the high level of development, the intelligence of mammals, in particular monkeys, has a clear biological limitation. Along with other forms of behavior, it is entirely determined by the way of life and biological laws, beyond which the animal cannot step over. This is shown by numerous observations of great apes in nature. So, chimpanzees build rather complex wicker nests in which they spend the night, but they never build even the simplest canopies from the rain and get mercilessly wet during tropical downpours. Under natural conditions, monkeys rarely use tools, preferring, if necessary, to obtain more affordable food than to spend time and effort on the extraction of hard-to-reach ones.

The limitations of intellectual behavior were also shown in numerous experiments conducted by Ladygina-Kots on apes. For example, a male chimpanzee sometimes made stupid mistakes when using objects provided to him to push bait out of a pipe. He tried to push a piece of plywood into the pipe, despite the obvious discrepancy between its width and the diameter of the pipe, and began to nibble it only after a number of such unsuccessful attempts. According to Ladygina-Cotes, chimpanzees “are not able to immediately grasp the essential features in a new situation.” [31]

Even the most complex manifestations of monkey intelligence are ultimately nothing more than the application of a phylogenetically developed mode of action in new conditions. Monkeys are able to attract fruit to themselves with a stick only because in natural conditions they often have to bend down a branch with a fruit hanging on it. It is the biological conditionality of all the mental activity of monkeys, including anthropoids, that is the reason for the limitedness of their intellectual abilities, the inability to establish a mental connection between mere representations and their combination into images. The inability to mentally operate with representations leads monkeys to an inability to understand true cause-and-effect relationships, since this is possible only with the help of concepts that monkeys, like all other animals, completely lack.

Meanwhile, at this stage in the development of science, the problem of animal intelligence has not been studied enough. In essence, detailed experimental studies have so far been carried out only on monkeys, mainly higher ones, while the possibility of intellectual actions in other vertebrates is practically not confirmed by conclusive experimental data. However, it is a mistake to assume that intelligence is inherent only in primates. Most likely, objective research by future zoopsychologists will help shed light on this difficult, but very interesting question.

Topic 7. Human psyche

7.1. The evolution of the human psyche in phylogenesis. The origin of labor activity, social relations and articulate speech

At the earliest stages of evolution, man, paying attention to the differences and similarities in the behavior of animals, tried to realize his attitude to the animal world. This fact is supported by the special role that man assigned to the behavior of animals, reflecting it in various rituals, fairy tales, and legends. Legends and rituals of this type were created independently on different continents and were of great importance in shaping the consciousness of primitive man.

Much later, with the emergence of scientific thinking, the problems of animal behavior, its psyche, the search for a "soul" became an integral part of many philosophical concepts. Some ancient thinkers recognized the close relationship between man and animals, placing them on the same level of mental development, while others categorically denied the slightest connection between human mental activity and similar animal activity. It was the ideological views of ancient scientists that determined the interpretation of the behavioral and mental activity of animals for many centuries.

The subsequent surge of interest in the mental activity of man in comparison with the mental activity of animals was associated with the development of evolutionary doctrine. Ch. Darwin and his followers one-sidedly emphasized the similarity and kinship of all mental phenomena, from lower organisms to man. Darwin categorically denied the fact that there are any differences between the human psyche and the psyche of animals. In his works, he very often attributed human thoughts and feelings to animals. Such a one-sided understanding of the genetic relationship between the psyche of an animal and a person was criticized by V.A. Wagner.

Wagner insisted that it is not the psyche of man and animals that should be compared, but the psyche of the forms inherent in the previous and subsequent group of animals. He pointed to the existence of general laws of the evolution of the psyche, without the knowledge of which it is impossible to understand human consciousness. Only such an approach, according to this scientist, could reliably reveal the prehistory of anthropogenesis and correctly understand the biological prerequisites for the emergence of the human psyche.

At present, we can judge the process of anthropogenesis, as well as the origin of human consciousness, only indirectly, by analogy with living animals. But we should not forget that all these animals have gone through a long path of adaptive evolution and their behavior has been deeply imprinted by specialization to the conditions of existence. Thus, in higher vertebrates, in the evolution of the psyche, a number of lateral branches are observed that are not related to the line leading to anthropogenesis, but reflect only the specific biological specialization of individual groups of animals. For example, in no case should one compare the behavior of human ancestors with the behavior of birds or the behavior of many highly developed mammals. Even the living primates most likely followed a regressive path of evolution, and all of them are currently at a lower level of development than the human ancestor. Any, even the most complex, mental abilities of monkeys, on the one hand, are entirely determined by the conditions of their life in the natural environment, their biology, and on the other hand, they only serve to adapt to these conditions.

All these facts should be remembered when searching for the biological roots of anthropogenesis and the biological prerequisites for the emergence of human consciousness. From the behavior of the now existing monkeys, as well as other animals, we can only judge the direction of mental development and the general laws of this process on the long path of anthropogenesis.

Origin of work activity. It is well known that the main factors in the development of human consciousness are labor activity, articulate speech and the social life created on their basis. At the present stage, the most important task for animal psychologists is to study the ways of development of human labor activity using the example of the use of tool activity by higher animals. Labor has been manual since its inception. The human hand is primarily an organ of labor, but it also developed thanks to labor. The development and qualitative transformations of the human hand occupy a central place in anthropogenesis, both physically and mentally. The most important role is played by its grasping abilities - a phenomenon quite rare in the animal world.

All biological prerequisites for labor activity should be sought in the characteristics of the grasping functions of the forelimbs of mammals. In this regard, a reasonable question arises: why did monkeys, and not other animals with grasping forelimbs, become human ancestors? This problem was studied for a long time by K.E. Fabry, studying in a comparative aspect the relationship between the main (locomotor) and additional (manipulative) functions of the forelimbs in monkeys and other mammals. As a result of numerous experiments, he came to the conclusion that the antagonistic relationship between the main and additional functions of the forelimbs plays an important role in the process of anthropogenesis. The ability to manipulate arose at the expense of basic functions, in particular, fast running. In most animals with prehensile forelimbs (bears, raccoons), manipulative actions fade into the background, they are, as it were, an unimportant appendage, without which the animal, in principle, can live. Most of these animals lead a terrestrial lifestyle, and the main function of their forelimbs is motor.

The exception is primates. Their primary form of locomotion is climbing by grasping branches, and this form constitutes the main function of their limbs. With this method of movement, the muscles of the fingers are strengthened, their mobility increases, and most importantly, the thumb is opposed to the rest. This structure of the hand determines the ability of monkeys to manipulate. Only in primates, according to Fabry, the main and additional functions of the forelimbs are not in antagonistic relationships, but are harmoniously combined with each other. As a result of a harmonious combination of locomotion and manipulative actions, the development of motor activity became possible, which elevated monkeys above other mammals and later laid the foundation for the formation of specific motor capabilities of the human hand.

The evolution of the primate hand proceeded simultaneously in two directions:

1) increasing flexibility and variability of grasping movements;

2) increasing the full grasp of objects. As a result of this bilateral development of the hand, the use of tools became possible, which can be considered the first stage of anthropogenesis.

Simultaneously with progressive changes in the structure of the forelimbs, there were also profound correlative changes in the behavior of human ancestors. They develop musculoskeletal sensitivity of the hand, which after a while will acquire leading importance. Tactile sensitivity interacts with vision, there is an interdependence of these systems. As vision begins to partially transfer its functions to skin sensitivity, hand movements with its help are controlled and corrected, become more accurate. In the animal kingdom, only monkeys have a relationship between vision and hand movements, which is one of the most important prerequisites for anthropogenesis. Indeed, without such interaction, without visual control over the actions of the hands, it is impossible to imagine the origin of even the simplest labor operations.

The interaction of vision and tactile-kinesthetic sensitivity of the hands is concretely embodied in the extremely intense and diverse manipulative activity of monkeys. Many Soviet zoopsychologists (N.N. Ladygina-Kots, N.Yu. Voitonis, K.E. Fabry, and others) studied the labor activity of monkeys. As a result of numerous experiments, it was revealed that both lower and higher monkeys carry out a practical analysis of the object in the course of manipulation. For example, they try to break the object that fell into their hands, examine its various details. But in higher apes, in particular in chimpanzees, there are also actions for the synthesis of objects. They may try to twist individual parts, twist them, twist them. Similar actions are observed in great apes and in the wild, when building nests.

In addition to constructive activity, in some monkeys, in particular chimpanzees, some other types of activity are distinguished that manifest themselves when manipulating objects - these are orienting-observing, processing, motor-playing, tool activities, as well as preserving or rejecting an object. The objects of orienting-examining, processing and constructive activity are most often objects that cannot be used for food. Tool activity in chimpanzees is rather poorly represented. This separation of the forms of various activities can be explained by analyzing the characteristics of the life of these monkeys in natural conditions. The orienting-observing and processing activity occupies a large place in the behavior of chimpanzees, which is explained by the variety of plant foods and the difficult conditions in which one has to distinguish between edible and inedible. In addition, the food items of monkeys can have a complex structure, and in order to reach the edible parts (extract insect larvae from stumps, remove the shell from tree fruits), it will take effort.

The constructive activity of chimpanzees, in addition to nest building, is very poorly developed. In conditions of captivity, these monkeys can twist twigs and ropes, roll balls of clay, but this behavior is not aimed at obtaining the final result, but on the contrary, most often it turns into a destructive one, into the desire to break something, to unravel. This type of behavior is explained by the fact that under natural conditions the tool activity of a chimpanzee is extremely poorly represented, since the monkey does not need this type of behavior to achieve its goals. Under natural conditions, tools are used extremely rarely. Cases have been observed of extracting termites from their buildings with twigs or straws, or collecting moisture from depressions in a tree trunk with a chewed clump of leaves. In actions with twigs, the most interesting circumstance is that before using them as tools, chimpanzees (as in the experiments of Ladygina-Kots described earlier) break off the leaves and side shoots that interfere with them.

Under laboratory conditions, chimpanzees can form quite complex tool actions. This serves as proof that the data obtained under experimental conditions testify only to the potential mental abilities of monkeys, but not to the nature of their natural behavior. The use of tools can be considered an individual, and not a specific feature of the behavior of monkeys. Only under special conditions can such individual behavior become the property of the entire group or pack. One should constantly bear in mind the biological limitations of anthropoids' tool actions and the fact that here we are clearly dealing with the rudiments of former abilities, with an extinct relic phenomenon that can fully develop only in the artificial conditions of a zoopsychological experiment.

It can be assumed that the use of tools was much better developed among fossil anthropoids - the ancestors of man - than among modern anthropoid apes. According to the current state of tool activity in the lower and higher apes, we can judge the main directions of the labor activity of our fossil ancestors, as well as the conditions in which the first labor actions originated. The prerequisite for labor activity was, apparently, the actions performed by modern anthropoids, namely, the cleaning of branches from leaves and side knots, the splitting of a torch. But among the first anthropoids, these tools did not yet act as tools, but rather were a means of biological adaptation to certain situations.

According to K.E. Fabry, objective activity in ordinary forms could not go beyond biological laws and go into labor activity. Even the highest manifestations of manipulative (tool) activity in fossil apes would forever remain nothing more than forms of biological adaptation, if the immediate ancestors of man did not undergo fundamental changes in behavior, analogs of which Fabry discovered in modern apes under certain extreme conditions. This phenomenon is called "compensatory manipulation". Its essence lies in the fact that in a laboratory cage, with a minimum of objects of study, a noticeable restructuring of manipulatory activity is observed in monkeys, and the animal begins to "create" much more objects than in natural conditions, where there are plenty of objects for ordinary destructive manipulation. In cage conditions, when objects for manipulation are almost completely absent, the normal manipulation activity of monkeys is concentrated on those few objects that they can have (or are given to them by the experimenter). The natural need of monkeys to manipulate numerous diverse objects is compensated in an environment sharply depleted in subject components by a qualitatively new form of manipulation - compensatory manipulation.

Only as a result of fundamental restructuring of objective actions, in the process of evolution, could labor activity develop. If we turn to the natural conditions of the origin of mankind, it can be noted that they were actually characterized by a sharp depletion of the habitat of our animal ancestors. Tropical forests were rapidly shrinking, and many of their inhabitants, including monkeys, found themselves in sparse or completely open areas, in an environment that was more monotonous and poor in objects for manipulation. Among these monkeys were also forms close to the human ancestor (Ramapithecus, Paranthropus, Plesianthropus, Australopithecus), and also, obviously, our immediate Upper Pliocene ancestor.

The transition of animals, the structure and behavior of which was formed in the conditions of forest life, to a qualitatively different habitat was associated with great difficulties. Almost all anthropoids are extinct. In the new habitat conditions, those anthropoids gained an advantage, in which, on the basis of the original way of moving through the trees, bipedalism developed. Animals with free forelimbs found themselves in a biologically more advantageous position, since they were able to use their free limbs for the development and improvement of tool activity.

Of all the anthropoids of open spaces, only one species survived, which later became the ancestor of man. According to most anthropologists, he was able to survive in changing environmental conditions only through the successful use of natural objects as tools, and then the use of artificial tools.

It should not be forgotten, however, that tool activity was able to fulfill its saving role only after a profound qualitative restructuring. The need for such a restructuring was due to the fact that manipulative activity (vital for the normal development and functioning of the motor apparatus) in a sharply depleted environment of open spaces had to be compensated. Forms of "compensatory modeling" arose, which eventually led to a high concentration of elements of the psychomotor sphere, which raised the tool activity of our animal ancestor to a qualitatively new level.

The further development of labor activity cannot be imagined without the use of various tools, as well as the emergence of special tools. Any object used by an animal to solve a specific task can serve as a direct tool, but a tool must certainly be specially made for certain labor operations and requires knowledge of its future use. This type of tool is made in advance, before its use becomes necessary. The making of tools can only be explained by foreseeing the occurrence of situations in which it is indispensable.

In modern monkeys, any tool is not assigned its special meaning. The object serves as a tool only in a specific situation, and, losing the need for use, it also loses its significance for the animal. The operation performed by the monkey with the help of a tool is not fixed behind this tool; outside of its direct application, it treats it indifferently, and therefore does not permanently store it as a tool. The manufacture of tools and their storage presupposes the foreseeing of possible causal relationships in the future. Modern monkeys are not able to comprehend such relationships even when preparing a tool for direct use in the course of solving a problem.

Unlike monkeys, man keeps the tools he makes. Moreover, the tools themselves preserve human methods of influencing natural objects. Even when made individually, a tool is a public item. Its use was developed in the process of collective work and secured in a special way. According to K. Fabry, “every human tool is the material embodiment of a certain socially developed labor operation.” [32]

The emergence of labor radically rebuilt the entire behavior of anthropoids. From the general activity aimed at the immediate satisfaction of a need, a special action is singled out, not directed by a direct biological motive and gaining its meaning only with the further use of its results. This change in behavior marked the beginning of human social history. In the future, social relations and forms of action that are not directed by biological motives become fundamental for human behavior.

The manufacture of a tool (for example, hewing one stone with the help of another) requires the participation of two objects at once: the first, to which changes are made, and the second, to which these changes are directed and which as a result becomes a tool of labor. The impact of one object on another, which could potentially become a tool, is also observed in monkeys. However, these animals pay attention to the changes that occur with the object of direct influence (the tool), and not to the changes that occur with the processed object, which serves as nothing more than a substrate. In this respect, monkeys are no different from other animals. Their instrumental actions are directly opposite to the instrumental actions of a person - for him, the most important changes occur with the second object, from which, after a series of operations, a tool of labor is obtained.

Hundreds of thousands of years have passed from the creation of the first tools of labor like the hand ax of Sinanthropus to the creation of various perfect tools of labor of a modern type human (neoanthrope). But it should be noted that already at the initial stages of the development of material culture, one can see a huge variety of types of tools, including composite ones (heads of darts, flint inserts, needles, spear throwers). Later, stone tools appeared, such as an ax or a hoe.

Along with the rapid development of material culture and mental activity since the beginning of the Late Paleolithic era, the biological development of man has sharply slowed down. Among the most ancient and ancient people, the ratio was reversed: with an extremely intensive biological evolution, expressed in a great variability of morphological features, the technique of making tools developed extremely slowly. There is a well-known theory of Ya.Ya. Roginsky, which was called "a single jump with two turns". According to this theory, simultaneously with the emergence of labor activity (the first turn), the most ancient people developed new socio-historical patterns. But along with this, biological laws also acted on the ancestors of modern man for a long time. The gradual accumulation of a new quality at the final stage of this development led to a sharp (second) turn, which consisted in the fact that these new social patterns began to play a decisive role in the life and further development of people. As a result of the second turn in the late Paleolithic, modern man arose - a neoanthropist. After its appearance, biological laws finally lost their leading significance and gave way to social laws. Roginsky emphasizes that only with the advent of the neoanthrope do social patterns become a truly dominant factor in the life of human groups.

If we follow this concept, the first human labor actions were carried out in the form of a combination of compensatory manipulation and instrumental activity enriched by it, as mentioned in his works by Fabry. After a long time, the new content of objective activity acquired a new form in the form of specifically human labor movements that are not characteristic of animals. Thus, at first, the outwardly uncomplicated and monotonous objective activity of the first people corresponded to the great influence of biological laws inherited from the animal ancestors of man. Ultimately, as if under the cover of these biological laws, labor activity arose, which formed a person.

The problem of the emergence of social relations and articulate speech. Already at the very beginning of working life, the first social relations arose. Labor was originally collective and social. Since their appearance on earth, monkeys have lived in large herds or families. All biological prerequisites for human social life should be sought in the objective activities of their ancestors, carried out in conditions of a collective way of life. But it is necessary to remember one more feature of work activity. Even the most complex instrumental activity does not have the character of a social process and does not determine the relations between members of the community. Even in animals with the most developed psyche, the structure of the community is never formed on the basis of tool activity, does not depend on it, much less is not mediated by it.

Human society does not obey the laws of group behavior of animals. It arose on the basis of other motivations and has its own laws of development. K.E. Fabry wrote about this: “Human society is not simply a continuation or complication of the community of our animal ancestors, and social patterns are not reducible to the ethological patterns of life of a herd of monkeys. Social relations of people arose, on the contrary, as a result of the breakdown of these patterns, as a result of a radical change in the very essence herd life and emerging work activity." [33]

For a long time, N.I. Voitonis. His numerous studies were aimed at studying the peculiarities of the structure of the herd and the herd behavior of various monkeys. According to N.I. Voitonis and NA Tych, the need for monkeys in a herd way of life originated at the lowest level of primate evolution and flourished in modern baboons, as well as in great apes living in families. In the animal ancestors of man, the progressive development of gregariousness also manifested itself in the formation of strong intra-herd relationships, which, in particular, turned out to be especially useful when hunting together with the help of natural tools. From the direct ancestors of a person, adolescents obviously had to learn the traditions and skills that had been formed in previous generations, learn from the experience of older members of the community, and the latter, especially males, should not only show mutual tolerance, but also be able to cooperate, coordinate their actions. All this was required by the complexity of joint hunting with the use of various objects (stones, sticks) as hunting tools. At the same time, at this stage, for the first time in the evolution of primates, conditions arose when it became necessary to designate objects: without this, it was impossible to ensure the coordination of actions of herd members during joint hunting.

According to Fabry, a special phenomenon, which he called "demonstrative manipulation", played a great role in the early stages of the formation of human society. In a number of mammals, cases are described when some animals observe the manipulative actions of other animals. This phenomenon is most typical for monkeys, which in most cases react briskly to the manipulative actions of another individual. Sometimes animals tease each other with objects of manipulation, often manipulation turns into games, and in some cases into quarrels. Demonstration manipulation is characteristic mainly of adult monkeys, but not of cubs. It contributes to the fact that individuals can become familiar with the properties and structure of the object manipulated by the "actor" without even touching the object. Such acquaintance is made indirectly: someone else's experience is assimilated at a distance by observing the actions of others.

Demonstrative manipulation is directly related to the formation of "traditions" in monkeys, which has been described in detail by a number of Japanese researchers. Such traditions are formed within a closed population and cover all its members. In a population of Japanese macaques living on a small island, a gradual general change in eating behavior was found, which was expressed in the development of new types of food and the invention of new forms of its pre-processing. The basis of this phenomenon was the play of cubs, as well as demonstrative manipulation and imitative actions of monkeys.

Demonstration manipulation combines communicative and cognitive aspects of activity: observing animals receive information not only about the manipulating individual, but also about the properties and structure of the object being manipulated. According to K.E Fabry, “demonstrative manipulation served in its time, obviously, as the source of the formation of purely human forms of communication, since the latter arose along with labor activity, the predecessor and biological basis of which was the manipulation of objects in monkeys. At the same time, it was the demonstration manipulation creates the best conditions for joint communicative and cognitive activity, in which the main attention of community members is focused on the objective actions of the manipulating individual.” [34]

An important milestone in anthropogenesis, which largely changed the further course of evolution, was the development of articulate speech at a certain stage in social relations.

In modern monkeys, the means of communication, communications are distinguished not only by their diversity, but also by their pronounced addressing, an inciting function aimed at changing the behavior of members of the herd. But unlike the communicative actions of a person, any communicative actions of monkeys do not serve as an instrument of thinking.

The study of the communicative abilities of monkeys, especially apes, has been carried out for a long time and in many countries. In the USA, scientist D. Premack tried for a long time to teach chimpanzees human language using various optical signals. Animals developed associations between individual objects, which were pieces of plastic, and food. In order to receive a treat, the monkey had to choose the desired one from various objects and show it to the experimenter. The experiments were based on the “sample selection” technique developed by Ladygina-Kots. Using these methods, reactions to categories of objects were developed and generalized visual images were formed. These were representations such as “more” and “smaller,” “same” and “different,” and comparisons of different types, which animals below anthropoids are most likely incapable of.

This and similar experiments clearly demonstrated the exceptional abilities of great apes for generalizations and symbolic actions, as well as their great ability to communicate with a person that arises under conditions of intensive training on his part. However, such experiments do not prove that anthropoids have a language with the same structure as that of humans. The chimpanzees literally “imposed” human language instead of trying to make contact in the language of this primate. In this sense, experiments of this kind are unpromising and cannot lead to an understanding of the essence of the language of an animal, since they give only a phenomenological picture of artificial communication behavior that outwardly resembles the operation of human language structures. The apes developed only a system of communication with humans, in addition to the many systems of human-animal communication that he created since the time of the domestication of wild animals.

According to K.E. Fabry, who has been studying the problem of the language of great apes for a long time, “despite the sometimes amazing ability of chimpanzees to use optical symbolic means when communicating with humans and, in particular, to use them as signals of their needs, it would be a mistake to interpret the results of such experiments as evidence of an allegedly fundamental identity "the language of monkeys and human language or to derive from them direct indications of the origin of human forms of communication. The illegality of such conclusions follows from an inadequate interpretation of the results of these experiments, in which conclusions about the patterns of their natural communication behavior are drawn from the behavior of monkeys artificially formed by the experimenter." [35]

As Fabry noted, “the question of the semantic function of animal language is still largely unclear, but there is no doubt that not a single animal, including apes, has conceptual thinking. As has already been emphasized, among the communicative means of animals there are many “symbolic” components (sounds, poses, body movements, etc.), but there are no abstract concepts, no words, articulate speech, no codes denoting the objective components of the environment, their qualities or the relationship between them outside a specific situation. Such a fundamentally different way of communication from animals could only appear during the transition from the biological to the social plane of development." [36]

The language of animals in the general sense is a very conditional concept, in the early stages of development it is characterized by a large generalization of the transmitted signals. Later, during the transition to a social way of life, it was the conditionality of signals that served as a biological prerequisite for the emergence of articulate speech in the course of their joint labor activity. At the same time, only emerging social and labor relations could fully develop this prerequisite. According to most scientists, the first language signals carried information about the subjects included in the joint labor activity. This is their fundamental difference from the language of animals, which informs primarily about the internal state of the individual. The main function of animal language is social cohesion, recognition of individuals, location signaling, danger signaling, etc. None of these functions goes beyond biological patterns. In addition, the language of animals is always a genetically fixed system, consisting of a limited number of signals, defined for each species. In contrast, human language is constantly enriched with new elements. By creating new combinations, a person is forced to constantly improve the language, study its code values, learn to understand and pronounce them.

The human language has come a long way of development in parallel with the development of human labor activity and the change in the structure of society. The initial sounds accompanying labor activity were not yet genuine words and could not denote individual objects, their qualities or actions performed with the help of these objects. Often these sounds were accompanied by gestures and were woven into practical activities. It was possible to understand them only by observing the specific situation during which these sounds were uttered. Gradually, of the two types of information transfer - non-verbal (using gestures) and voice - the latter became a priority. This marked the beginning of the development of an independent sound language.

However, innate sounds, gestures, facial expressions have retained their significance since primitive people and have come down to our days, but only as an addition to acoustic means. Nevertheless, for a long time the connection of these components remained so close that the same sound complex could designate, for example, the object pointed to by the hand, the hand itself, and the action performed with this object. A lot of time passed before the sounds of the language were quite far removed from practical actions and the first genuine words arose. Initially, obviously, these words denoted objects, and only much later did words denoting actions and qualities appear.

Subsequently, the language began to gradually move away from practical activities. The meanings of words became more and more abstract, and language increasingly acted not only as a means of communication, but also as a means of human thought. At their inception, speech and social and labor activity constituted a single complex, and its separation radically influenced the development of human consciousness. K. Fabry wrote: “The fact that thinking, speech and social and labor activity constitute a single complex in their origin and development, that human thinking could develop only in unity with social consciousness, constitutes the main qualitative difference between human thinking and the thinking of animals. The activity of animals, even in its highest forms, is entirely subordinate to natural connections and relationships between the objective components of the environment. Human activity, which grew out of the activity of animals, has undergone fundamental qualitative changes and is no longer subject to so much natural as social connections and relationships. This is social- labor content and reflect the words and concepts of human speech." [37]

Even in higher animals, the psyche is capable of reflecting only spatio-temporal connections and relations between the objective components of the environment, but not deep causal relationships. The human psyche is on a completely different level. It is able to directly or indirectly reflect social ties and relationships, the activities of other people, its results - this is what allowed a person to comprehend even cause-and-effect relationships that are inaccessible to observation. As a result, it became possible to reflect objective reality in the human brain outside the direct relationship of the subject to it, i.e., in the human mind, the image of reality no longer merges with the experience of the subject, but the objective, stable properties of this reality are reflected.

Most major psychologists tend to think that the development of human thinking to its present level would be impossible without language. Any abstract thinking is linguistic, verbal thinking. Human knowledge presupposes the continuity of acquired knowledge, the ways of their fixation, carried out with the help of words. Animals are deprived of the possibility of both verbal communication and verbal fixation of acquired knowledge and their transmission to offspring with the help of language. This, firstly, determines the limit of the thinking and communication capabilities of animals, and secondly, characterizes the biological, purely adaptive role of their communication. Animals do not need words to communicate; they can do just fine without them, living in a narrow circle limited by biological needs and motivations. Communication of a person without words, which are the highest, ideal objects of thinking abstracted from things, is impossible.

Thus, there is a clearly defined line between the intellect of an animal and the consciousness of a person, and thus this line also passes between an animal and a person in general. The transition through it became possible only as a result of an active, radically different impact on nature in the course of labor activity. This activity, performed with the help of tools, mediated the relationship of its performer to nature, which served as the most important prerequisite for the transformation of the preconscious psyche into consciousness.

The first elements of a mediated relationship to nature can be traced back to the manipulative actions of monkeys, especially during compensatory manipulation and in tool actions, as well as in demonstrative manipulation. But, as was discussed above, in modern monkeys even the highest manipulative actions serve other reasons and are not capable of further developing into complex labor activity. Genuine tool actions that took place among the ancestors of modern man are situationally determined, therefore their cognitive value is extremely limited by the specific, purely adaptive meaning of these actions. These instrumental actions received their development only after the merging of compensatory manipulation with instrumental actions, when attention is switched to the object being processed (future tool), which occurs during labor activity. It was this indirect attitude to nature that allowed man to reveal the essential internal interdependencies and laws of nature that were inaccessible to direct observation.

The next important stage in the development of human consciousness was the formation of social labor activity. At the same time, it became necessary to communicate with each other, which led to the consistency of joint labor operations. Thus, articulate speech was formed simultaneously with consciousness in the process of labor activity.

Despite the fact that the historical development of mankind is fundamentally different from the general laws of biological evolution, which psychologists have repeatedly emphasized in their works, it was the biological evolution of animals that created the biological basis and prerequisites for an unprecedented transition in the history of the organic world to a completely new level of development. This can be seen by carefully examining all the stages in the development of the mental activity of animals. Without the development of the simplest instincts, their long-term improvement as a result of evolution, without the lower stages of the development of the psyche, the emergence of human consciousness would also be impossible.

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.

Literature

1. Anokhin P.K. Key questions of the theory of functional system. M., 1980.
2. Vagner V.A. Biological foundations of comparative psychology. T. 1, 2. St. Petersburg; M., 1910-1913.
3. Vagner V.A. Animal psychology. 2nd ed. M., 1902.
4. Voitonis N.Yu. Background of the intellect. M.; L., 1949.
5. Darwin C. Expression of emotions in humans and animals. T. 5. M., 1953.
6. Dembovsky Ya. Psychology of monkeys. M., 1963.
7. Dewsbury D. Animal behavior. Comparative aspects. M., 1981.
8. Krushinsky L.V. Biological foundations of rational activity: evolutionary and physiological aspects of behavior. M., 1986.
9. Ladygina-Kots N.N. Study of the cognitive abilities of chimpanzees. M., 1923.
10. Ladygina-Kots N.N. Constructive and tool activity of higher apes (chimpanzees). M., 1959.
11. Ladygina-Kots N.N. Background of human thinking. M., 1965.
12. Ladygina-Kots N.N. The development of the psyche in the process of evolution of organisms. M., 1959.
13. Leontiev A.N. Problems of the development of the psyche. 3rd ed. M., 1972.
14. Lorenz K. Ring of King Solomon. M., 1970.
15. Lorenz K. A man finds a friend. M., 1971.
16. McFarland D. Animal behavior. Psychobiology, ethology and evolution. M., 1988.
17. Nesturkh M.F. Primatology and anthropogenesis. M., 1960.
18. Orbeli L.A. Questions of higher nervous activity. M.; L., 1949.
19. Panov E.N. Signs, symbols, languages. 1983.
20. Ponugaeva A.G. Imprinting (imprinting). L., 1973.
21. Promptov A.N. Essays on the problem of biological adaptation of the behavior of passerine birds. M.; L., 1956.
22. Rubinshtein S.L. Fundamentals of General Psychology: In 2 vols. M., 1989.
23. Ruler K.F. Selected biological works. M., 1954.
24. Severtsov A.N. Evolution and the psyche. M., 1922.
25. Slonim. HELL. Instinct. L., 1967.
26. Tinbergen N. The world of the herring gull. M., 1975.
27. Tinbergen N. Animal behavior. M., 1969.
28. Tikh N.A. Background of society. L., 1970.
29. Fabre J.-A. Insect life. M., 1963.
30. Fabry C.E. Animal play. M., 1985.
31. Fabry C.E. Weapon actions of animals. M., 1980.
32. Fabry C.E. Fundamentals of zoopsychology. M., 1992.
33. Firsov L.A. The behavior of anthropoids in natural conditions. L., 1977.
34. Frisch K. From the life of bees. M., 1966.
35. Hind R. Animal behavior. M., 1975.
36. Heinrot O. From the life of birds. M., 1947.
37. Chauvin R. Animal behavior. M., 1973.

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

We recommend interesting articles Section Lecture notes, cheat sheets:

▪ Logistics. Lecture notes

▪ Pedagogical psychology. Lecture notes

▪ History and theory of religions. Crib

See other articles Section Lecture notes, cheat sheets.

Read and write useful comments on this article.

<< Back

Latest news of science and technology, new electronics:

Artificial leather for touch emulation 15.04.2024

In a modern technology world where distance is becoming increasingly commonplace, maintaining connection and a sense of closeness is important. Recent developments in artificial skin by German scientists from Saarland University represent a new era in virtual interactions. German researchers from Saarland University have developed ultra-thin films that can transmit the sensation of touch over a distance. This cutting-edge technology provides new opportunities for virtual communication, especially for those who find themselves far from their loved ones. The ultra-thin films developed by the researchers, just 50 micrometers thick, can be integrated into textiles and worn like a second skin. These films act as sensors that recognize tactile signals from mom or dad, and as actuators that transmit these movements to the baby. Parents' touch to the fabric activates sensors that react to pressure and deform the ultra-thin film. This ... >>

Petgugu Global cat litter 15.04.2024

Taking care of pets can often be a challenge, especially when it comes to keeping your home clean. A new interesting solution from the Petgugu Global startup has been presented, which will make life easier for cat owners and help them keep their home perfectly clean and tidy. Startup Petgugu Global has unveiled a unique cat toilet that can automatically flush feces, keeping your home clean and fresh. This innovative device is equipped with various smart sensors that monitor your pet's toilet activity and activate to automatically clean after use. The device connects to the sewer system and ensures efficient waste removal without the need for intervention from the owner. Additionally, the toilet has a large flushable storage capacity, making it ideal for multi-cat households. The Petgugu cat litter bowl is designed for use with water-soluble litters and offers a range of additional ... >>

The attractiveness of caring men 14.04.2024

The stereotype that women prefer "bad boys" has long been widespread. However, recent research conducted by British scientists from Monash University offers a new perspective on this issue. They looked at how women responded to men's emotional responsibility and willingness to help others. The study's findings could change our understanding of what makes men attractive to women. A study conducted by scientists from Monash University leads to new findings about men's attractiveness to women. In the experiment, women were shown photographs of men with brief stories about their behavior in various situations, including their reaction to an encounter with a homeless person. Some of the men ignored the homeless man, while others helped him, such as buying him food. A study found that men who showed empathy and kindness were more attractive to women compared to men who showed empathy and kindness. ... >>

Random news from the Archive Chromebooks updated and discounted 01.01.2012 In the "cloud" Chrome OS has a number of innovations. In particular, the login screen has changed, the browser has acquired new tabs, music applications, games and a file manager have been added. A little earlier, the Chrome Web Store was redesigned, the main page of which is now an "endless wall" with the most interesting applications. At the same time, Google agreed with the manufacturers of new prices. In 2012, "chromobooks" will fall in price by $50: for example, prices for models from Acer and Samsung start at $299. At the same time, Samsung introduced a new version of the Chromebook Series 5 - completely black.

Other interesting news:

▪ Salty soil dries up planets

▪ Samsung Gear VR Innovator Edition - virtual reality glasses for smartphones

▪ Release of memristor chips delayed

▪ Electronics work inside the body

▪ A new company in the LCD backlight market

News feed of science and technology, new electronics

Interesting materials of the Free Technical Library:

▪ section of the site Electrician in the house. Article selection

▪ article With the seal of power on the forehead. Popular expression

▪ article Do any birds hibernate? Detailed answer

▪ article cleaner. Job description

▪ article MIDI keyboard on PIC16F84. Encyclopedia of radio electronics and electrical engineering

▪ article Electrolysis and electroplating installations. Copper electrolysis plants. Encyclopedia of radio electronics and electrical engineering

Leave your comment on this article:

All languages ​​of this page

Arabic	Hebrew	Polish
Bengali	Hindi	Portuguese
Chinese	Italian	Spanish
English	Japanese	Turkish
French	Korean	Ukrainian
German	Malay	Vietnamese

Home page | Library | Articles | Website map | Site Reviews

www.diagram.com.ua
2000-2024