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.

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

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