Back at the beginning of the XX century. American researcher W. Craig ("Attractions and antipathies as components of instinct", 1918) showed that any instinctive action consists of separate phases. Craig identified two phases that were named: search engine (preparatory) phase, or appetizing behavior, and final phase (final act).
Craig showed that in natural conditions, animals look for those key stimuli or their combinations (triggering situations) that are necessary for the implementation of a certain instinctive response. For example, animals look for food, individuals of the opposite sex during the breeding season, places for nesting, etc. Craig called these search forms of behavior appetizing, and the state of the animal at this moment - appetite. Intermediate stimuli perceived at the search phase of behavior are not a goal for the animal; they are only necessary to lead to the perception of key stimuli of the final behavior. The final phase of instinctive behavior is the very consumption of the elements of the environment by the animal, it is this phase that is directly instinctive behavior.
The final phase is hereditarily determined, species-typical, it contains the biological meaning of the entire 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 animal's body. During this phase, only minor individual behavioral variations are possible, which are genetically determined. The acquired components of behavior practically do not play a role in the final act, and more often than not they are completely absent. K. Lorenz named 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 for the species. It intertwines innate and acquired forms of behavior, the individual experience of the animal. The exploratory behavior is characterized by the orientation-research activity of the animal. It is through changes in appetite behavior that instinctive responses can be plastic. The preparatory phase is always divided into several stages. Its end occurs when the animal reaches a situation in which the next link in this chain of reactions can be realized. For example, the choice of a nesting territory by a male sometimes requires only returning to the old, last year's territory, and sometimes it may require lengthy searches and even struggle with other males. According to K. Lorentz, the search phase of a behavioral act should be attributed to purposeful behavior. At this stage, various actions are performed, but they are all subordinated to a specific goal. The search phase is very important and is as vital for the animal as the consumption in the final phase. It is appetite behavior that is a means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes the 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 external world.
Craig's concept of two phases of instinctive behavior was based on data obtained from studying the feeding behavior of animals. A hungry predator starts looking for prey. However, at first he does not have information about its location and therefore his search activity is not yet directed. Soon, the predator sees a potential prey, from which the first key stimulus emanates, for example, size and color details, and its searching behavior moves to the next stage, which already has a certain direction. The predator begins to clarify the location, the speed of movement of the prey, focusing on other key stimuli. Then the predator chases the prey or sneaks up on it imperceptibly, after which it grabs and kills it. If necessary, the victim is dragged to another place, where it is cut into pieces. Only after this does the animal's behavior enter the final phase, which includes direct eating of prey. All actions of the animal associated with the search, catching and killing of the victim refer to appetizing behavior. All of them have an instinctive basis, but in to a large extent depend on the process of individual learning, the experience of the animal and the situation.
Each stage of the search behavior has its own preparatory and final phases. The end of one stage is a signal for the beginning of the next, etc. Successive stages often have several degrees of subordination, so a complex structure of animal behavior is formed. For example, search behavior can 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. First, the territory for the nest is selected. When she is found, the next stage of search behavior begins - building a nest, then the next - courting a female, etc.
Animal behavior is largely composed of cycles, which, in turn, are composed of a series of repeated simple acts. For example, a bird engaged in building a nest does it according to a certain pattern. First, she goes in search of building material, then, having found it, assesses the suitability. If the bird suits the material, it carries it to the nest, otherwise it throws it and looks for a new one. Having arrived in the nest, the bird weaves the brought materials into its structure with certain movements, forms the shape of the nest, and then flies away again in search. This cycle begins spontaneously and continues as long as the bird needs to complete the nest. Switching to each next stage of the behavioral reaction is carried out upon the perception of a certain external stimulus. N. Tinbergen gives an example of female hymenoptera insects - philanthropists (bee wolves), which feed their larvae with honey bees. The wasp, to make supplies, flies to places of accumulation of bees, where it flies randomly until it meets a suitable prey. Noticing a flying insect, the wasp flies up to it from the leeward side and stops at about 70 cm. If a bee is deodorized with ether, the wasp will not grab it. The next stage of the wasp's behavior will be the paralysis of the victim with a sting blow. The stimulus associated with touching the sacrifice is needed to start this stage. If you present a bee model to a wasp, which does not feel like it to the touch, but has the same smell, the wasp will not sting such a model. Thus, when an animal passes through various stages of a behavioral reaction, a change in stimuli occurs, which are key for it at the moment.
condition appetite 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 intensified 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 theory. The basis for this theory was the data of the German physiologist E. Holst.
Holst in his experiments focused on the rhythmic activity of the central nervous system. He noted that rhythmic discharges of impulses can be observed in the isolated abdominal nerve cord of the earthworm, which exactly correspond to the contraction of the segments of the worm. In further research, Holst studied the swimming mechanism of the eel. He fixed the middle segments of his body and did not allow them to contract. 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, they actually start moving 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 disturbed. Thus, the movements of the eel's body are performed not according to the mechanism of a reflex arc (depending on external stimuli), but obeying the rhythmic discharges of impulses in the central nervous system... Experiments by other scientists have confirmed this. For example, it was noted that decerebrated (with deleted large hemispheres) in cats, the antagonist muscles of the leg, completely devoid of sensory nerves, can rhythmically contract. Tadpoles and fish with one intact sensory nerve retain the ability to swim and normal coordination of movements. This means that endogenous automatism is inherent in the central nervous system, which does not depend on external stimuli. In this case, a minimum level of afferent impulses is required to maintain excitement, ("Specific energy of action") in the corresponding nerve centers at a certain level.
The works of E. Holst and his colleagues confirmed that the level of arousal in the corresponding nerve centers influences the nature of the course of instinctive reactions. The experiments were carried out on chickens, which were stimulated by the current of the brain stem structures. Depending on the localization of the irritated structure, the researchers noted elementary behavioral reactions (turning the head, pecking) or complex acts of behavior (courtship). And if simple reactions always proceeded approximately the same, regardless of the parameters of stimulation and environmental conditions, then complex behavioral reactions depended on these factors. So, with a weak current strength, the rooster pounced on a stuffed ferret, and with an increase in the current, even on the researcher's hand (non-specific stimulus).
The history of the study of instinctive behavior goes back several centuries, but there is still no clear, unambiguous and universally accepted definition of instinct.
Ever since the works of Charles Darwin, instinctive behavior has been understood as that part of animal behavior that is species-typical and hereditary. 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 identified five points on which "instinctive" behavior differs from "rational".
Instinctive behavior:
Hereditarily fixed;
No additional training is required for its development;
It is the same in all individuals of a given species, that is, species-typical;
Optimally matches the organization of the animal, its physiology;
Optimally adapted to the natural living conditions of animals of this species, it is often associated even 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.
Famous Russian physiologist I.P. Pavlov(1849-1936, "Conditioned reflexes: a study of the physiological activity of the cerebral hemispheres of the brain", 1925), one of the founders of the reflex theory, proposed to consider concepts reflex and instinctidentical. In this case, an unconditioned reflex corresponds to instinctive behavior. This concept 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 neural mechanism that responds to certain proposed and permissive impulses (external and internal) with fully coordinated, vital and species-specific movements."
Tinbergen created a hierarchical theory of instinct, which we will explore later.
Soviet physiologist HELL. Slonimgives the following definition: “Instinct is a set of motor acts and complex behaviors, characteristic of an animal of this species, arising from irritations from the external and internal environment of the body and proceeding 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. "
Slonim drew attention to the fact that instincts can appear and disappear during the life of the organism. For example, the instinctive behavior associated with the sucking reflex in young mammals disappears over time, but instincts for reproduction and nest-building activities appear. HELL. Slonim points to constancy as the main property of instinctive behavior. In his opinion, minor individual differences cannot refute this property, but are only fluctuations in its manifestation.
The plasticity of instinctive behavior.This question is one of the key in zoopsychology. In order to understand the behavioral responses of an animal, it is important to find out whether the innate behavior is permanent or it can be subject to modification. Currently, scientists have come to the conclusion that individual instinctive actions are not inherited, only the framework within which instinctive reactions develop 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 "The 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, so it can not be invariable. It is a plastic and labile system that develops under the influence of natural selection. Only species-typical frameworks that determine the amplitude of instinct variability are stable.
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 for a given species. They are formed by a set of innate species-typical instinctive reactions and conditioned reflexes acquired on their basis in ontogeny. 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 animal behavior are very important for a correct understanding of the problem of instinctive behavior. However, according to these concepts, the instinctive actions themselves are not subject to variations, their variability is provided only by acquired components.
It is now believed that instinctive behavior is subject to changes within the hereditarily fixed rate of response. These frames are species-typical; outside of them, instinctive behavior under normal conditions cannot change. At the same time, in extreme conditions that go beyond the normal response, the individual experience of the animal plays a large role. It allows instinctive behavior to change quite a bit. In addition, in addition to highly conservative innate mechanisms, there is a variable component of behavior.
A.N. Severtsov in his writings analyzed the variability of the instinctive and acquired component of behavior. Severtsov showed that in mammals, adaptation to changes in the external environment is carried out in two ways: through a change in the organization, i.e., the structure and functions of the body, and through a change in behavior. Organizational changes can only adapt to slow changes in the environment, because they take a long time. Changes in behavior do not require a restructuring of the structure and functioning of the animal's body, therefore, they occur at a sufficiently high speed. Such changes arise from acquired, individual forms of behavior and allow the animal to adapt to abrupt changes in the environment. At the same time, 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 vertebrate brain that is taking place in evolution.
According to Severtsov, instinctive behavior is not variable enough, therefore its significance in evolution is approximately comparable to changes in the structure of the animal's body. Changes in innate behavior can also allow an animal to adapt to slow environmental changes. However, the role of such changes should in no way be underestimated.
According to Severtsov, “instincts are the essence of species adaptations, useful for the species to the same extent as those or other morphological signs, and just as constant. "
The ability to learn, according to Severtsov, depends on the hereditary height of the mental organization. Actions in this case are not instinctively fixed. And in instinctive behavior, both the action and the level of mental organization are hereditarily fixed. In other words, instinctive behavior is an innate program of action 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 implemented in the course of acquiring individual experience. Instinctive behavior should be sufficiently unchanged and stereotyped, because it concerns the functions vital for the animal. If instinct depended on the conditions in which each member of a species develops, individuals would not be able to benefit from species experience. 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 acquired components of behavior, learning processes. These processes make it possible to adapt a rather 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 fulfillment of vital functions.
The structure of instinctive behavior.Back at the beginning of the XX century. American researcher W. Craig ("Desires and antipathies as components of instinct", 1918) showed that any instinctive action consists of separate phases. Craig identified two phases, which were named: search engine(preparatory) phase, or appetizing behavior,and finalphase (final act).
Craig showed that in natural conditions, animals look for those key stimuli or their combinations (triggering situations) that are necessary for the implementation of a certain instinctive response. For example, animals look for food, individuals of the opposite sex during the breeding season, places for nesting, etc. Craig called these search forms of behavior appetizing,and the state of the animal at this moment - appetite.Intermediate stimuli perceived in the search phase of behavior are not a goal for the animal; they are only necessary to lead to the perception of key stimuli of the final behavior. The final phase of instinctive behavior is the very consumption of the elements of the environment by the animal, it is this phase that is directly instinctive behavior.
The final phase is hereditarily determined, species-typical, it contains the biological meaning of the entire 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 animal's body. During this phase, only minor individual behavioral variations are possible, which are determined genetically. The acquired components of behavior practically do not play a role in the final act, and more often than not they are completely absent. K. Lorenz named 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 for the species. It intertwines innate and acquired forms of behavior, the individual experience of the animal. The exploratory behavior is characterized by the orientation-research activity of the animal. It is through changes in appetite behavior that instinctive responses can be plastic. The preparatory phase is always divided into several stages. Its end occurs when the animal reaches a situation in which the next link in this chain of reactions can be realized. For example, the choice of a nesting territory by a male sometimes requires only returning to the old, last year's territory, and sometimes it may require lengthy searches and even struggle with other males. According to K. Lorentz, the search phase of a behavioral act should be attributed to purposeful behavior. At this stage, various actions are performed, but they are all subordinated to a specific goal. The search phase is very important and is as vital for the animal as the consumption in the final phase. It is appetite behavior that is a means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes the 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 external world.
Craig's concept of two phases of instinctive behavior was based on data obtained from studying the feeding behavior of animals. A hungry predator starts looking for prey. However, at first he does not have information about her location and therefore his search activity is not yet directed. Soon, the predator sees a potential prey, from which the first key stimulus emanates, for example, size and color details, and its searching behavior moves to the next stage, which already has a certain direction. The predator begins to clarify the location, the speed of movement of the prey, focusing on other key stimuli. Then the predator chases the prey or sneaks up on it imperceptibly, after which it grabs and kills it. If necessary, the victim is dragged to another place, where it is cut into pieces. Only after this does the animal's behavior enter the final phase, which includes direct eating of prey. All actions of the animal associated with the search, catching and killing of the victim refer to appetizing 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 the search behavior has its own preparatory and final phases. The end of one stage is a signal for the beginning of the next, etc. Successive stages often have several degrees of subordination, therefore a complex structure of animal behavior is formed. For example, search behavior can 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. First, the territory for the nest is selected. When she is found, the next stage of search behavior begins - building a nest, then the next - courting a female, etc.
Animal behavior is largely composed of cycles, which, in turn, are composed of a series of repeated simple acts. For example, a bird engaged in building a nest does it according to a certain pattern. First, she goes in search of building material, then, having found it, assesses the suitability. If the bird suits the material, it carries it to the nest, otherwise it throws it and looks for a new one. Arriving at the nest, the bird weaves the materials brought into its structure with certain movements, forms the shape of the nest, and then flies away again in search. This cycle begins spontaneously and continues as long as the bird needs to complete the nest. Switching to each next stage of the behavioral reaction is carried out upon the perception of a certain external stimulus. N. Tinbergen gives an example of female hymenoptera insects - philanthropists (bee wolves), who feed their larvae with honey bees. The wasp, to make supplies, flies to places of accumulation of bees, where it flies randomly until it meets a suitable prey. Noticing a flying insect, the wasp flies up to it from the leeward side and stops at about 70 cm.If after that the wasp catches the smell 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 deodorized with ether, the wasp will not grab it. The next stage of the wasp's behavior will be the paralysis of the victim with a sting blow. The stimulus associated with touching the sacrifice is needed to start this stage. If you present a bee model to a wasp, which does not feel like it to the touch, but has the same smell, the wasp will not sting such a model. Thus, when an animal passes through various stages of a behavioral reaction, a change in stimuli occurs, which are key for it at the moment.
condition appetiteoccurs 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 intensified 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 theory. The basis for this theory was the data of the German physiologist E. Holst.
Holst in his experiments focused on the rhythmic activity of the central nervous system. He noted that rhythmic discharges of impulses can be observed in the isolated abdominal nerve cord of the earthworm, which exactly correspond to the contraction of the segments of the worm. In further research, Holst studied the swimming mechanism of the eel. He fixed the middle segments of his body and did not allow them to contract. 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, they actually start moving 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 disturbed. Thus, the movements of the eel's body are performed not according to the mechanism of a reflex arc (depending on external stimuli), but obeying the rhythmic discharges of impulses in the central nervous system. Experiments by other scientists have confirmed this. For example, it was noted that decerebrated (with the large hemispheres removed) cats can rhythmically contract the antagonist muscles of the leg, 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 endogenous automatism is inherent in the central nervous system, which does not depend on external stimuli. In this case, a minimum level of afferent impulses is required to maintain excitement, ("Specific energy of action")in the corresponding nerve centers at a certain level.
The works of E. Holst and his colleagues confirmed that the level of arousal in the corresponding nerve centers affects the nature of the course of instinctive reactions. The experiments were carried out on chickens, which were stimulated by the current of the brain stem structures. Depending on the localization of the irritated structure, the researchers noted elementary behavioral reactions (head turns, pecking) or complex behaviors (courtship). And if simple reactions always proceeded approximately the same, regardless of the parameters of stimulation and environmental conditions, then complex behavioral reactions depended on these factors. So, with a weak current strength, the rooster pounced on a stuffed ferret, and with an increase in the current, even on the researcher's hand (non-specific stimulus).
17. Instinctive behavior and communication in animals: ritualized and demonstrative behavior.
Communication - This is a physical (biological) and mental (information exchange) 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 in animals, there are necessarily special forms of behavior that perform the functions of transferring information between individuals. In this case, some actions of the animal acquire a signal value. Communication in this sense is absent in lower invertebrates, and in higher invertebrates it appears only in its rudimentary form. It is inherent in all representatives of vertebrates to one degree or another.
The German ethologist G. Tembrok studied the process of communication in animals and its evolution. According to Tembrok, one can speak about real animal communities in which individuals communicate with each other only when they begin a life 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.
Communication is based on communication (information exchange). For 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 with the help of which the transfer is carried out and the assimilation of information occurs, are hereditarily fixed, are instinctive.
Forms of communication.According to the mechanism of action, all forms of communication differ in information transmission channels. There are optical, acoustic, chemical, tactile and other forms.
Among optical forms of communication, the most important place is occupied by expressive postures and body movements, which make up “ demo behavior”. This behavior consists of showing animals certain parts of their bodies, which, as a rule, carry specific signals. These can be brightly colored areas, additional structures such as ridges, decorating feathers, etc. At the same time, some parts of the animal's body can visually increase in volume, for example, due to ruffling of feathers or hair. The signaling function can also be performed by special movements of the body or its individual parts. By performing these movements, the animal can display colored areas of the body. Sometimes such demonstrations are performed 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 movement of the claw of a beckoning crab, which it performs when courting a female. Such movements are called "Allochthonous". Allochthonous movements are species-typical and stereotyped, their function is to transmit information. Their other name is ritualized movements. All ritualized movements are conditional. They are very rigidly and clearly fixed genetically, refer to typical instinctive movements. It is this kind of conservatism of movements that ensures the correct perception of signals by all individuals, regardless of living conditions.
Most often and in the greatest number, ritualized movements are observed in the sphere of reproduction (primarily mating games) and struggle. They transmit information to one individual about the internal state of another individual, about its physical and mental qualities.
18. Learning as an individually acquired form of animal regulation: general characteristics.
Learning is a modification of behavior that occurs as a result of the individual experience of an individual, and is not a consequence of growth, maturation, aging of the body, or a consequence of fatigue, sensory adaptation (for example, a scarecrow and crows) ..
As a result of learning, behavior can change as follows: 1) behavioral acts that are completely new for a given individual can arise; 2) the habitual behavioral response will be caused by a stimulus that was not previously associated with it; 3) it is possible to change the likelihood or form of the response, which usually appeared earlier to the presented stimulus in a given situation.
Learning may not be adaptive to the individual. (An example is "bad habits", that is, obsessive, compulsive behavior, the purpose of which is to find a more acceptable way of protection from the once experienced trauma.
The study of different taxonomic groups in animals has qualitative differences. However, behavioral zoopsychologists have formulated " general patterns learning ".
1) Thorndike's "law of effect". For a reaction that is followed by a reward or a state of satisfaction, the likelihood of repetition is increased, and for a reaction that causes a harmful or unpleasant consequence, the likelihood of repetition decreases.
2) Skinner's principle of least effort. Animals strive to receive rewards in the fastest and most convenient way.
3) The law of instinctive displacement of learned behavior (formulated by the Breland couple - American zoopsychologists). "Learned behavior shifts towards the instinctive whenever the strong innate instincts of the animal are similar to the conditioned response. The law of reinforcement is not sufficient to overcome the innate tendencies towards certain types of behavior."
4) Yerkes-Dodson law. Learning occurs most successfully with optimal motivation. If motivation exceeds the optimum, learning will be slower and more mistakes will be made.
In zoopsychology, there are two approaches to learning classification... The first approach to the classification of learning belongs to the non-behaviourist E. Tolman. The approach is based on the awareness of the fact that learning mechanisms are heterogeneous among representatives of different systematic groups. E. Tolman classified the ability of animals and humans to learn and highlighted the following ability groups:
1) the ability to expect the appearance of a stimulus and act in accordance with this prediction; 2) the ability to discriminate and manipulate; 3) the ability to retain experience; 4) the ability to foresee the result of one's actions in the case of an alternative choice of options for simple motor reactions (labyrinth); 5) the ability to form representations, allowing you to compare alternative ways of solving the problem; 6) "creative flexibility".
The second approach assumes that the whole variety of learned behavior can be reduced to several basic types. These are the classifications of Thorp, Godefroy, Fabry. The classification of J. Godefroy is based on the degree of participation of an individual in the learning process, and, with the active participation of the organism, the cognitive level is also highlighted.
Ethologists distinguish obligate and optional learning, according to the principle of the need for a particular learning reaction for survival under standard conditions. Obligatory (compulsory) learning is a set of skills necessary for the survival of an individual. Optional (optional) learning is skills that appear in some individuals, in response to specific conditions. Observing mammals in an anthropogenic environment, ethologists found out that in females the main role is played by obligate learning, which is formed mainly on the influence of natural factors. In the life of males, optional learning is more important, which appears in response to the action of anthropogenic factors.
The structure of instinctive behavior
Search and final phases of the behavioral act
It was said above that key stimuli act forcibly, that the animal is forced in its behavior to completely obey the triggering situation. But does this mean that animals have no opportunity to show their own initiative, to make some kind of independent choice? Not at all!
The proactive, selective attitude of the animal to the environment is manifested primarily in the active search for the necessary triggering situations and in the selection of the most effective opportunities for performing behavioral acts. It should be emphasized that we are talking about the search for stimuli emanating from biologically significant objects, and not these objects themselves. We now know that these are key stimuli with a guiding or triggering function.
More than half a century ago, the American researcher of animal behavior W. Craig showed that instinctive actions consist of separate phases. First of all, Craig identified two phases, which in the ethological literature are called "search" (or "preparatory") and "final". During the search phase, the animal searches (hence the name of the phase) those key stimuli, more precisely, their combinations (i.e. triggering situations) that will eventually lead it to the final phase in which it is embodied biological significance all instinctive action.
All intermediate stimuli do not constitute an end in themselves for the animal and are valuable only insofar as they lead to the perception of key stimuli of the final behavior. Only in the final phase does the animal actually consume the vital elements of the environment. But the very search for adequate stimuli is for animals the same primary vital necessity as the consumption of environmental elements.
The search phase is always subdivided into several stages; however, in the final phase, either such subdivisions are not found at all, or it consists only of a few strictly sequential movements.
Craig based his concept on data he obtained from studying the feeding behavior of animals. Here's an example of this area of \u200b\u200bbehavior. At first, a predator going hunting does not yet know where its possible prey is, so its first movements have the character of an undirected search. As a result, it sooner or later falls within the scope of the stimulus emanating from the prey animal. The first key stimulus was discovered, which includes the next stage - directed orientation to additional stimuli, clarification of the location of the prey animal. This is followed by sneaking (or pursuit), throwing (jumping) and taking possession of the prey, killing it, sometimes also dragging the carcass to another place, dismembering it into separate pieces and, finally, grabbing the pieces of meat with the teeth and swallowing them. In this chain of successively performed actions and movements, only the last two links (actually the act of eating) refer to the final phase of the described food-gathering behavior of the predator; all other stages in their totality make up search (or preparatory) behavior. True, within each such stage there are preparatory and final phases, with which each stage ends. At the same time, sometimes there are several degrees of subordination (like "nesting dolls"), so that on the whole, a very complex structure of activity develops.
The situation is similar in other seemingly much simpler spheres of behavior, such as rest and sleep. The animal first looks for a place to rest or sleep (trees, shelters, depressions in the soil, or just certain areas of open space), then it arranges (improves) the found place (digs, crushes vegetation), sometimes it is also cleaned and only after that it fits (moreover, species-typical way!). Only laying down constitutes the final phase, while the preceding stages are the search phase.
To these examples, many others could be added from any area of \u200b\u200bbehavior. However, already in the above, one can see the following deep differences between the two phases, which determine their essence.
Search behavior is the plastic phase of instinctive behavior. It is characterized by a pronounced orienting-exploratory activity of animals and an interweaving of congenital and acquired components of behavior based on individual experience. Everything that is connected with the plasticity of instincts, in particular with modifications of instinctive behavior, belongs to search behavior.
Finishing behavior, on the other hand, is a rigid phase. The movements performed in it are distinguished by a strict sequence, stereotype and are predetermined by the corresponding macro- and micromorphological structures. The purchased components play an insignificant role here or are even absent. Therefore, variability is limited to individual (genetically fixed) variability. This includes everything that has been said about the constancy, rigidity of instinctive behavior and the coercive effect of key stimuli. Here, almost everything is innate, genetically fixed. As for the movements performed in the final phase, these are actually instinctive movements, or "innate motor coordination", so called by the Austrian scientist K. Lorentz, one of the founders of modern ethology.
From the book Doping in Dog Breeding author Gourmet E G6.2.4. The structure of the coat An expert cynologist during the exhibition examination does not miss the opportunity not only to examine, but also to touch the dogs. For him, the texture of the wool of the evaluated animals, especially the leaders, is important. Most rock standards clearly specify gloss, hardness
From the book Fundamentals of Zoopsychology author Fabri Kurt ErnestovichInternal factors of instinctive behavior As already mentioned, the problem of instinct and learning is directly related to another equally important problem - the problem of internal and external factors, the motivation of behavior. For a long time it was believed that instinctive actions
From the book Fundamentals of Neurophysiology author Shulgovsky Valery ViktorovichExternal factors of instinctive behavior When talking about the autonomy of internal factors of behavior, about their independence from the external environment, it must be remembered that this independence is only relative. Already from the above experiments of Holst it is clear that
From the book Microbiology: lecture notes author Tkachenko Ksenia ViktorovnaAcquired components of the instinctive act Along with those indicated in the search phase of any instinctive act, there are always - in varying degrees and in different combinations - all elements of behavior that relate to learning, not excluding higher forms of behavior,
From the book General ecology author Chernova Nina MikhailovnaComplexity and Diversity of the Structure of Instinctive Behavior The two-phase structure of instinctive actions is presented here only in the form of a very complete, simplified general scheme. In reality, most often there are various complications and modifications. Primarily
From the book Forms of learning and methods of training dogs to search for explosives, explosive devices, weapons and ammunition author Gritsenko Vladimir VasilievichThe mental component of instinctive behavior The mental component of instinctive behavior, more precisely, mental reflection during the instinctive actions of animals, must be studied at the final phase of the behavioral act and, to some extent, directly
From the book Biology [ Complete reference to prepare for the exam] author Lerner Georgy IsaakovichChapter 4 PHYSIOLOGICAL MECHANISMS OF REGULATION OF VEGETATIVE FUNCTIONS AND INSTINCTIVE BEHAVIOR The vegetative functions include those functions that provide metabolism in our body (digestion, blood circulation, respiration, excretion, etc.). They also include providing
From the book Fundamentals of Psychophysiology author Alexandrov Yuri1. Structure of immunoglobulins Antibodies (immunoglobulins) are proteins that are synthesized under the influence of an antigen and specifically react with it. They consist of polypeptide chains. Four structures are distinguished in the immunoglobulin molecule: 1) the primary one is
From the book Stop, Who Leads? [The biology of human and other animal behavior] author Zhukov. Dmitry Anatolyevich1. Structure HIV belongs to the family of retroviruses. The virion has a spherical shape with a diameter of 100-150 nm. Cubic type of symmetry. The outer (supercapsid) envelope of the virus consists of a bimolecular lipid layer that originates from the cell membrane of the cell
From the book Human Genetics with the Basics of General Genetics [ Tutorial] author Kurchanov Nikolay Anatolievich7.2. The structure of the biocenosis The structure of any system is the regularities in the relationship and connections of its parts. The structure of the biocenosis is multifaceted, and when studying it, various
From the author's book1.2. Structure of Behavior The totality of possible behavior of an animal is called a behavioral repertoire. The unit of behavior is a behavioral act. A behavioral act is a set of elements of animal behavior from the moment of occurrence
From the author's book From the author's book7.3. The structure of the subjective world and the subject of behavior The specialization of neurons with respect to the elements of individual experience means that their activity does not reflect external world as such, but the relationship with it of the individual (see also paragraph 4). That is why the description
From the author's bookChapter 3 The structure of behavior The concept of behavior The objective observer judges mental phenomena not directly, but by measuring their manifestations in the activity of a living organism, primarily in behavior. Therefore, let us dwell on this concept in more detail.
From the author's bookSleep structure Night sleep consists of several cycles, each of which lasts about an hour and a half and includes two qualitatively different stages, called "slow sleep" and "REM sleep". Stages of REM and slow sleep are not only in humans, but also
From the author's book3.3. Chromosome structure Each chromatid contains one DNA molecule associated with histone and non-histone proteins. At present, a nucleosomal model of the organization of eukaryotic chromatin has been adopted (Kornberg R., 1974; Olins A., Olins D., 1974). According to this model, histone proteins (they
It was said above that key stimuli act forcibly, that the animal is forced in its behavior to completely obey the triggering situation. But does this mean
Search engineand final phasesbehavioral act
that animals have no opportunity to show their own initiative, to make some kind of independent choice? Not at all!
The proactive, selective attitude of the animal to the environment is manifested primarily in the active search for the necessary triggering situations and in the selection of the most effective opportunities for performing behavioral acts. It should be emphasized that we are talking about the search for stimuli emanating from biologically significant objects, and not these objects themselves. We now know that these are key stimuli with a 4-ounce guide or trigger.
More than half a century ago, the American researcher of animal behavior W. Craig showed that instinctive actions consist of separate phases. First of all, Craig identified two phases, which in the ethological literature are called "search" (or "preparatory") and "final". During the search phase, the animal searches (hence the name of the phase) those key stimuli, more precisely, their combinations (i.e. triggering situations) that will eventually lead it to the final phase, in which the biological significance of the entire instinctive action is embodied.
All intermediate stimuli do not constitute an end in themselves for the animal and are valuable only insofar as they lead to the perception of key stimuli of the final behavior. Only in the final phase does the animal actually consume the vital elements of the environment. But the very search for adequate stimuli is for animals the same primary vital necessity as the consumption of environmental elements.
The search phase is always subdivided into several stages; however, in the final phase, either such subdivisions are not found at all, or it consists only of a few strictly sequential movements.
Craig built his concept on data from his eating behavior studies
animals. Here's an example of this area of \u200b\u200bbehavior. At first, a predator going hunting does not yet know where its possible prey is, so its first movements have the character of an undirected search. As a result, it sooner or later falls within the scope of the stimulus emanating from the prey animal. The first key stimulus was discovered, which includes the next stage - directed orientation to additional stimuli, clarification of the location of the prey animal. This is followed by sneaking (or pursuit), throwing (jumping) and taking possession of the prey, killing it, sometimes also dragging the carcass to another place, dismembering it into separate pieces and, finally, grabbing the pieces of meat with the teeth and swallowing them. In this chain of successively performed actions and movements, only the last two links (actually the act of eating) refer to the final phase of the described food-gathering behavior of the predator; all other stages in their totality make up search (or preparatory) behavior. True, within each such stage there are preparatory and final phases, with which each stage ends. At the same time, sometimes there are several degrees of subordination (like "nesting dolls"), so that on the whole, a very complex structure of activity develops.
The situation is similar in other seemingly much simpler spheres of behavior, such as rest and sleep. The animal first looks for a place to rest or sleep (trees, shelters, depressions in the soil, or just certain areas of open space), then it arranges (improves) the found place (digs, crushes vegetation), sometimes it is also cleaned and only after that it fits (moreover, - in a typical way!). Only laying down constitutes the final phase, while the preceding stages are the search phase.
To these examples, many others could be added from any area of \u200b\u200bbehavior. However, already in
given above, one can see the following deep differences between the two phases, which determine their essence. Search behavior is the plastic phase of instinctive behavior. It is characterized by a pronounced orienting-exploratory activity of animals and an interweaving of congenital and acquired components of behavior based on individual experience. Everything that is connected with the plasticity of instincts, in particular with modifications of instinctive behavior, belongs to search behavior.
Finishing behavior, on the other hand, is a rigid phase. The movements performed in it are distinguished by a strict sequence, stereotype and are predetermined by the corresponding macro- and micromorphological structures. The purchased components play an insignificant role here or are even absent. Therefore, variability is limited to individual (genetically fixed) variability. This includes everything that has been said about the constancy, rigidity of instinctive behavior and the coercive effect of key stimuli. Here, almost everything is innate, genetically fixed. As for the movements performed in the final phase, these are actually instinctive movements, or "innate motor coordination", so named by the Austrian scientist KLorentz, one of the founders of modern ethology.
general characteristics instinctive movements have already been given above. It was also said that they are the "keepers" of the most valuable, vital that is accumulated by the species as a result natural selection, and that it is precisely this that determines their independence from random conditions environment... The fox, making instinctive movements of burying meat on the stone floor, behaves "senseless". But after all, the phylogenesis of foxes did not take place on a stone substrate, and it would be fatal for the further existence of the species, if due to chance
Instinctive movements and taxis
if an individual was temporarily staying in such conditions completely atypical for the foxes' habitat, a form of behavior so useful for these animals would disappear. So it is better to make burying movements under any circumstances, and the "rigid" innate program of behavior forces the animal to do this.
The general orientation of instinctive movements is carried out by taxis, which, according to Lorenz, are always intertwined with innate motor coordination and together with them form unified instinctive reactions (or chains of several such reactions).
Like instinctive movements, taxis are innate, genetically fixed responses to certain environmental agents. But if instinctive movements arise in response to triggering stimuli, then taxis respond to guiding key stimuli, which are incapable of determining the beginning (or end) of any instinctive reaction, but only change the vector of its course.
Thus, taxis provide a spatial orientation of the motor activity of animals towards favorable or vital environmental conditions (positive taxis) or, conversely, from biologically low or dangerous conditions (negative taxis). In plants, similar reactions are expressed in changes in the direction of growth (tropisms).
By the nature of orienting external stimuli, taxis are subdivided into photo-, chemo-, thermo-, geo-, rheo-, anemo-, hydrotaxis (reactions to light, chemical stimuli, temperature gradients, gravity, fluid flow, air flow, environmental humidity) etc. At different levels evolutionary development taxis have varying degrees of complexity and perform different functions, which will be discussed further in the review of the evolution of the psyche. It is now important to emphasize that taxis are permanent components of even complex forms of behavior, and the higher forms of taxis appear in close combination with the individual experience of the animal.