Inherited form of behavior. The influence of genetic inheritance on behavior. Form of behavior in society

People differ from each other in a number of psychological characteristics. These differences are caused by both different living conditions and dissimilar genotypes, since people's genotypes contain different forms of genes. Psychogenetics studies the relative contribution of heredity and environment to the diversity of people in terms of psychological properties and behavior. To assess the influence of heredity and environment on human behavior, scientists compare people with varying degrees of genetic commonality (identical and multiple twins, siblings and half-siblings, children and their biological and adoptive parents).

Many genes exist in multiple forms, just as there are different forms of the gene that determines eye color. Some genes have dozens of forms. The genotype of a particular person contains two copies of each gene, the forms of which may be different or may be the same. One is inherited from the father, the other from the mother. The combination of forms of all genes is unique for each human body. This uniqueness underlies genetically determined differences between people. The contribution of genetic differences to the diversity of people in psychological properties is reflected by an indicator called the “heritability coefficient”. For example, for intelligence the heritability rate is at least 50%. This does not mean that 50% of intelligence is given to a person by nature, and the remaining 50% must be added through training, then intelligence will be 100 points. The heritability coefficient is not related to a specific person. It is calculated to understand why people differ from each other: whether the differences arise because people have different genotypes, or because they were taught differently. If the heritability coefficient of intelligence were close to 0%, then we could conclude that only learning creates differences between people, and applying the same educational techniques to different children will always lead to the same results. High values ​​of the heritability coefficient mean that even with the same upbringing, children will differ from each other due to their hereditary characteristics. The end result, however, is not determined by genes. It is known that children adopted into prosperous families are close to their adoptive parents in terms of intellectual development and can significantly exceed their biological ones. What then is the influence of genes? Let us explain this using the example of a specific study.*

Scientists examined two groups of adopted children. Conditions in the adoptive families were equally good for all, and the children's biological mothers differed in their level of intelligence. The biological mothers of children from the first group had above average intelligence. Approximately half of the children from this group demonstrated above average intellectual abilities, the other half - average. The biological mothers of the children in the second group had slightly reduced (but within normal limits) intelligence. Of this group, 15% of children had the same low intelligence scores; the remaining children had an average level of intellectual development. Thus, under the same conditions of upbringing in foster families, the intelligence of children, to a certain extent, depended on the intelligence of their blood mothers.

The above example can serve to illustrate the significant differences between the concept of heritability of psychological qualities and the heritability of certain physical characteristics of a person, such as eye color, skin color, etc. Even with a high level of heritability of a psychological trait, the genotype does not predetermine its final value. How a child will develop under certain environmental conditions depends on the genotype. In some cases, the genotype sets the “limits” for the expression of a trait.

The influence of heredity on intelligence and character at different ages

Research shows that genes are responsible for 50-70% of the diversity of people in terms of intelligence and for 28-49% of the differences in the severity of the five “universal” most important personality traits:

  • anxiety,
  • friendliness,
  • consciousness,
  • intellectual flexibility.

This data is for adults. However, the degree of influence of heredity depends on age. The results of psychogenetic studies do not support the widespread belief that with age, genes influence human behavior less and less. Genetic differences tend to become more pronounced in adulthood, when character has already been formed. The values ​​of the heritability coefficient for most of the studied psychological properties are higher for adults than for children. The most accurate data have been obtained on the hereditary conditioning of intelligence. In infancy, the intrapair similarity of multiple twins is as high as for identical twins, but after three years of age it begins to decline, which can be explained by the greater influence of genetic differences. At the same time, the increase in differences does not occur linearly. There are stages during which differences between children are caused primarily by environmental influences. For intelligence this is the age of 3-4 years, and for personality formation it is pre-teen age 8-11 years.

In addition, different genetic factors operate at different ages. Thus, among the hereditary factors that cause differences in intelligence, there are both stable ones, i.e. acting at all ages (these are possibly genes associated with the so-called “general intelligence”), and specific to each period of development (probably genes that determine the development of particular abilities).

The influence of heredity on antisocial behavior

Since in all developed countries, crime and alcoholism of biological parents are common reasons for a child losing his birth family and being placed in foster care, we will take a closer look at the evidence from psychogenetics about the influence of heritability on these forms of behavior. Family and twin studies of criminal behavior have been conducted for more than 70 years. They give very different heritability estimates, most often falling in the 30-50% range. "Upper" heritability values ​​are obtained by studying twins. Some researchers believe that the twin method may overestimate heritability because it does not always separate genetic influences from the special environmental conditions in which identical twins grow up. By studying adopted children, heritability coefficient values ​​are approximately 2 times lower than when studying twins.

Danish Study of Adopted Children


Figure 1. Number of families analyzed,

(Danish study).

The most systematic studies of the heritability of criminal behavior by studying adopted children were carried out in the Scandinavian countries - Denmark and Sweden. Thanks to the cooperation of adoptive parents and a number of authorities, Danish scientists were able to trace the fate of more than 14,000 people adopted between 1924 and 1947. Figures 1 and 2 show the results of a study of criminal records among men raised in foster care. They refer only to property crimes because the number of violent crimes was low.


Figure 2. Proportion of sons with a criminal record in families
differing in the criminal record of the biological and adoptive father
(Danish study).

From Figure 2 it can be seen that the proportion of convicts among children whose biological fathers were criminals is slightly increased compared to those children whose biological parents did not break the law. In addition, it turned out that the more criminal records a biological father has, the higher the risk for the offspring of becoming a criminal. It was also shown that brothers adopted different families, tended to be concordant in criminal behavior, especially in cases where their biological father was a criminal. These data indicate a certain role of heredity in increasing the risk of criminal behavior. However, as from the above example with intelligence, from the data in Figure 2 it follows that unfavorable heredity does not predetermine the future of the child - of the boys whose biological fathers were criminals, 14% subsequently violated the law, the remaining 86% did not commit illegal actions.

In addition, it turned out that the adoptive family has a particularly strong influence on children with unfavorable heredity, which can be both positive and negative. Of the boys who grew up in foster care, 16% subsequently committed crimes (versus 9% in the control group). Among the biological fathers of these children, 31% had problems with the law (versus 11% in the control group). Those. Although the crime rate among adopted children was higher than the community average, it was almost half that of their biological fathers. According to a number of scientists, this indicates that a favorable environment in a foster family reduces the risk of criminal behavior in children with a family history.

But in some cases, the family environment can increase the risk of criminal behavior. As can be seen from Figure 2, children whose biological and adoptive fathers had criminal records committed crimes more often than others. (Fortunately, there were very few such families (Fig. 1)). This means that there are genotypes that have increased vulnerability to unfavorable aspects of the family environment (such phenomena in psychogenetics are called genotype-environment interaction).

Swedish study

In a study of adopted children in Sweden, scientists initially did not find even a weak connection between the criminal records of children raised by adoptive parents and the behavior of their biological fathers. Among Swedes, crime was mainly a consequence of alcohol abuse. When the scientists excluded this type of crime from the analysis, they found a weak positive relationship between the criminal records of offspring and their blood fathers (Figure 3). At the same time, the crimes in both generations turned out to be not serious. These were mainly thefts and fraud.


Figure 3. Percentage of criminal records among adopted persons
depending on family type
(Swedish study).

The sensitivity of children with hereditary burdens to the characteristics of the adoptive family was also confirmed. There was no increase in crime rates among adopted Swedes compared to the national average, despite the fact that the conviction rate was increased among their biological parents. Among the Swedish adoptive parents there were no persons with a criminal record. Those. the most favorable family environment “neutralized” the effect of the genetic load. On the other hand, the highest risk of breaking the law was observed in those children with unfavorable heredity whose adoptive family had a low socio-economic status (Fig. 3).

American study


Figure 4. Results of studying the reasons leading to the formation of an antisocial personality,
in an American study of adopted children
(arrows indicate a statistically significant relationship between parental characteristics and the development of antisocial tendencies in children).

Scandinavian studies included analyzes of the behavior of adopted children born in the first half of the 20th century. Similar results were obtained in modern work American scientists from Iowa. True, it was not a criminal record that was analyzed, but the presence in adopted children of a tendency towards antisocial behavior of a wider spectrum. The behavior that serves as the basis for the diagnosis was assessed. antisocial disorder personality" and includes frequent behavior that could lead to arrest, as well as such traits as deceit, impulsiveness, irritability, disregard for safety, irresponsibility and dishonesty. We also took into account a number of characteristics of the adoptive family that could influence the formation of such tendencies. Figure 4 lists these characteristics and shows the main findings of the study when adoptees reached adulthood (ages 18 to 40).Only male data were analyzed because the number of females with “antisocial behavior” was too small Of the 286 men studied, forty-four were diagnosed with antisocial personality disorder.The results indicated that three factors independently contribute to the development of this disorder:

  1. criminal record of the biological parent (genetic),
  2. drunkenness or antisocial behavior of one of the members of the foster family (environmental),
  3. placement of a child with unfavorable heredity in a family with low socio-economic status (genotype-environment interaction).

What is a hereditary predisposition to antisocial behavior?

Obviously, in humans, genes do not trigger specific behavior in the same way as they do with some instinctive actions of animals. The relationship between the risk of criminal behavior and genes is mediated by psychological characteristics. Moreover, it is known that the risk of criminal behavior can be influenced by various unfavorable combinations of psychological properties, and each of these properties is under the control of several or a large number of genes and various environmental factors.

The first candidate for the role of a biological “substrate” of antisocial tendencies was the Y chromosome (a chromosome that is found only in the genotype of men and determines male sex). In approximately one in 1,100 men, biological errors in complex process the creation of a germ cell in the genotype there are two or more Y chromosomes instead of one. These men are distinguished by low intelligence (at the lower limit of the norm) and tall stature. In the 60s of the 20th century, it was first shown that among criminals serving sentences with reduced intelligence, a disproportionate number (4%) of men with an extra Y chromosome. At first, the connection between this genetic defect and criminal tendencies seemed obvious: since men are more aggressive than women, commit crimes more often, and, unlike women, have a Y chromosome, the presence of two or more Y chromosomes should lead to the formation of an aggressive “super man.” But later it turned out that criminals with an extra Y chromosome are no more aggressive than other prisoners, and they end up in prison mainly for committing thefts. At the same time, in men with this genetic pathology, a connection was found between a decrease in intelligence and the likelihood of being convicted. It is possible, however, that reduced intelligence did not affect the risk of committing a crime, but the risk of being caught and imprisoned. For example, one of the men with an extra Y chromosome broke into houses several times while the owners were in the room.

Studies of men with an extra Y chromosome lead to at least two important conclusions. First, the connection between genes and crime cannot be explained by a genetically determined increase in aggressiveness or cruelty, as one might assume based on “common sense.” This conclusion is consistent with data from studies of adopted children, in which the influence of heredity was found only for crimes against property. Secondly, even among men with such an obvious hereditary anomaly as an extra Y chromosome, the majority do not become criminals; we are only talking about a slight increase in the risk of such behavior among them.

Since the mid-90s, scientists have been searching for specific genes that could influence the risk of criminal behavior. All data obtained to date still needs confirmation and clarification. However, a study conducted in New Zealand deserves mention. It showed that among boys exposed ill-treatment in a family, carriers of a form of the gene that ensures higher activity of the MAOA enzyme in the body were less prone to antisocial behavior than carriers of another form of the gene - low activity. Among children who grew up in prosperous families, there was no connection between antisocial tendencies and the MAOA gene. Those. Individuals with certain genetic characteristics have been shown to be less vulnerable to parental abuse. This study made scientists think about whether it is even legitimate to talk about a hereditary predisposition (tendency) to antisocial behavior. Perhaps a more accurate concept would be the genetically determined vulnerability (insecurity) of some children in relation to adverse, traumatic events.

The influence of heredity on alcohol abuse

It has long been noted that crime and alcohol abuse are closely related. Moreover, psychogenetic studies have suggested that there are “predisposition genes” common to these forms of behavior. Some similar patterns have also been identified in the influence of heredity and environment on crime and alcohol abuse. For example, for both forms of behavior, a significant influence of the general environment** is found in adolescence. The influence of the common environment is manifested, in particular, in the fact that brothers and sisters growing up in the same family (even if they are not related) are more similar to each other in antisocial manifestations and habits associated with drinking alcohol than to their parents. However, alcohol abuse is a rather complex phenomenon from a behavioral and genetic point of view, since it includes everyday drunkenness and alcoholism as a gradually developing mental illness (the main diagnostic sign of which is an irresistible psychological attraction to alcohol).

Obviously, the role of genes in these two cases is different, but it can be quite difficult to separate these two forms of alcohol abuse in a psychogenetic study. This may be why estimates of the heritability of alcoholism vary widely. The most likely range seems to be 20-60%. Among sons of patients with alcoholism, according to various sources, on average 20-40%, and among daughters - from 2 to 25% (on average about 5%). At the same time, it can be considered established that the age at which they began to drink alcohol and the intensity of its consumption in the first stages is completely determined by the influence of the environment. Note that drinking alcohol at an early age (usually before 15 years of age) is a risk factor for the development of alcoholism. The absence of genetic influences on this trait indicates the important role of parental behavior that inhibits adolescent alcohol use in preventing the development of alcohol dependence. At the same time, genetic effects and genotype-environmental interactions are clearly detected in the further escalation of alcohol consumption and the development of alcoholism.

Let us emphasize, however, once again that a person is not born an alcoholic and there is no one “alcoholism gene,” just as there is no “crime gene.” Alcoholism is the result of a long chain of events that accompany regular drinking. A large number of genes influence these events to some extent. Yes, from character young man depends on how often he drinks and whether he knows when to stop, and character, as already mentioned, depends on both upbringing and genotype. In addition, due to their genetic characteristics, people are sensitive to the toxic effects of alcohol to varying degrees. For example, some Japanese, Koreans and Chinese have found this form of a gene that affects the processing of alcohol in the liver, the possession of which leads to very severe alcohol poisoning. A person with this form of the gene, after drinking alcohol, feels nausea, flushing, dizziness and irritation. These unpleasant sensations keep a person from further drinking alcohol, so among carriers of this form of the gene there are almost no alcoholics. Finally, not all people who regularly drink alcohol develop an irresistible craving for it. There are genes (they are now being intensively searched) that determine whether the long-term effect of alcohol on the brain will lead to alcohol dependence. At the same time, genes do not trigger specific forms of behavior, they do not “force” a person to go and drink. If a person knows they are susceptible to alcoholism, they can avoid situations that encourage drinking and remain healthy.

Children of alcoholics are often called a multiple-risk group. About 1/5 of them have various problems that require special attention from parents, teachers, and sometimes doctors. Mostly these are restlessness and neurotic disorders (tics, fear of the dark, etc.). Difficulties in mastering the school curriculum are observed less frequently, and other more serious disorders, for example, convulsive conditions, are even less common. These disorders are not manifestations of any defects in the genetic apparatus and are caused by unfavorable conditions in which mothers carry their pregnancies and raise their babies. Studies of adopted children have shown that alcoholism of birth parents does not increase the likelihood that the child will develop any serious mental disorder in the future.

Summarizing the existing data on the influence of heritability on antisocial behavior and alcoholism, the following conclusions can be drawn.

  • There is a positive, although very weak, relationship between the criminality of blood fathers and their sons who grew up in foster families.
  • This pattern is found only for minor crimes, so there is no reason to believe that the risk of becoming a criminal in adopted children is explained by a genetically determined increase in aggressiveness or cruelty.
  • Data indicate that a favorable family environment can neutralize innate characteristics associated with an increased risk of criminal behavior, while an unfavorable family environment can enhance them.
  • The development of antisocial tendencies is not inevitable even among carriers of serious genetic abnormalities.
  • The age at which they started drinking alcohol and the intensity of its consumption in the early stages is completely determined by the action of various environmental factors. Genetic effects and genotype-environment interactions are detected only for the subsequent escalation of alcohol consumption and the development of alcoholism.

*Willerman L. Effects of families on intellectual development. Quote according to "Psychogenetics" I.V. Ravich-Scherbo et al.

** Environmental influences in psychogenetics are divided into general and individual environments. The common environment refers to all non-hereditary factors that make compared relatives from the same family similar to each other and not similar to members of other families (we can assume that for psychological properties these are parenting styles, the socio-economic status of the family, its income, etc.). The individual environment includes all non-hereditary factors that form differences between family members (for example, a unique circle of friends, classmates or teachers for each child, memorable gifts or actions of adults, forced isolation from peers as a result of some kind of trauma or other individual events).

Alfimova Margarita Valentinovna,
candidate of psychological sciences,
Leading Researcher, Laboratory of Clinical Genetics
Scientific Center for Mental Health of the Russian Academy of Medical Sciences

Commentary on the Project "Towards a New Family"

It is worth considering that at the time the research problem was formulated, very narrow boundary conditions were set, which did not take into account several serious factors:

  • motivation and degree of preparedness of adoptive parents for parents' roles,
  • anxiety level of future parents,
  • the age at which the child was brought into the family and the level of his deprivation in the birth family or the institution where he was raised,
  • the family’s ability to methodically, independently or with the help of specialists, work to compensate for the child’s somatic and psychological problems.

All these factors were not previously given significant importance.

When studying the cancellation of adoptions and emerging psychological problems in adoptive families, a very high relationship was identified between the success and motivation of adoptive parents, as well as their preparedness for the role of parents. Quite often, future parents were not sufficiently prepared to accept a child. For example, they wanted to resolve issues of the family’s status in society by accepting a child into the family, restore relationships between each other, find an heir, raise an ideal child or child prodigy, and were not ready to accept him with all his characteristics and problems. This led to the fact that they were unable to love him and create a friendly, but only mentoring, upbringing environment. Until the age of 6-12 years, the parenting style does not greatly influence the occurrence of serious child-parent conflicts and antisocial manifestations in the child’s behavior, however, a mentoring environment or the so-called “responsible parenting style” works to adolescence and sharply increases the likelihood of conflicts developing into forms of protest (often antisocial) behavior of the child.

The situation is aggravated by increased suspicion and anxiety about the child’s behavior, which often leads to educational errors expressed in extreme forms of educational influence - impulsive, unjustifiably harsh measures or connivance, justified by the “inevitability of fate” and attributing one’s educational incompetence to genes. Thus, the antisocial behavior of biological parents is not genetic, but a powerful psychological factor of pressure on adoptive parents, provoking the risks of inadequate educational influence on the child. The influence of anxiety will be discussed in detail in a separate article.

The second most influential factor on the occurrence of antisocial behavior is the initial level of damage to the child’s nervous system and the success of its compensation in the foster family. Such damage to the nervous system occurs due to:

  • prenatal intoxication of the fetus with alcohol, drugs,
  • oxygen starvation, lack of microelements for the normal development of the nervous system with poor nutrition of the expectant mother,
  • birth injuries,
  • maternal deprivation of the child in the first days and years of life, and when the child enters an institution, the lack of natural communication with him and proper care.

The seriousness of the influence of the institutional environment was noticed quite a long time ago and was described in the 30s of the 20th century (Emmy Pickler), but the influence of parental competence in compensation on the success of adoption was noticed only in the late 70s. A child with deprivation problems requires special corrective action, otherwise undercompensated medical and psychological problems begin to manifest themselves in the form of deviant behavior during a period when rapid hormonal changes occur in the child’s body, and parents no longer have full power over the child - in adolescence.

Discussion

We also adopted a boy at 1.5 years old. They gave him all their soul and strength. All the mothers admired... But, unfortunately, now it is clear that he does not want to make an effort in anything. Everything is interesting at the level of pampering, he doesn’t want to strain and study. It’s as if there is no will... he can’t try, it’s easier for him to refuse the most tempting prospects... Now the child is 10 years old. But now I don’t know what he will do... I don’t exaggerate my demands. I don’t know whether it’s heredity (he’s a foundling, nothing is known about his parents at all), or a birth trauma, but the fact remains a fact. We are seeing a neurologist... They recommended a psychotherapist, let's go... maybe he can give me some advice... It's funny for me to read all sorts of accusations... like they don't love enough... And we loved and we love... but for now we can find a use for him in this life they couldn’t... Many good people got involved, wanted to help... they also put a lot of soul into it, to no avail... I’m afraid that the plant will grow... Honestly, I read letters from other adoptive parents and understand that my fears are justified. The child, however, is happy with everything :)

07/29/2012 22:26:09, Polinaaa

Unfortunately, more and more people are prone to jumping to conclusions without thinking through all the possible influences and combinations. This happens because people need to quickly know the answer to pressing questions and problems, especially those problems that pose a threat to people (in this article, this is the increasing crime in the world, thereby receiving a deceptive illusion of information (“forewarned is forearmed”) , turning our science into a familiar religious system of faith and primitive dogma. Once upon a time, ancient people (and some modern ones) tried, due to their lack of knowledge about the causes of the problems that confronted them, to absolutize their hasty conclusions, turning them into a system of information -mental submission - faith, since these conclusions did not have the necessary formalization, they could not be realized in the world without a system of unquestioning submission that avoids changing these conclusions. This article makes the same attempt to “quickly” find the causes of the problem, based on insufficient data and make hasty conclusions without taking many factors into account.

05/13/2008 15:22:14, Argyrogespera El "Feya

The elders are right. But, unfortunately, not only sinful entities are transmitted, but also behavioral patterns, and even karmic predestination of the future development of the fate of children. (How a person’s energy matrix is ​​fixed on the physical level is a separate conversation.) But this is important to understand not only when adopting, but also when choosing a partner to start a family. That is, for example, it is stupid to marry someone whose family and parents are far from what you would like for your children. No matter what love she has, over time she will still begin to repeat the natural line of behavior and fate for her. It’s not for nothing that they say, if you want to know what your wife will be like, look at your mother-in-law, if you want to know what awaits you in the family, look at her family. Unfortunately, the law that if bad things are possible, then they inevitably happen is quite true for fairly long periods of life. Both the child and the spouse must be selected in the image and likeness of what you would like for yourself. It is necessary and correct to assess your responsibility to future generations of your descendants. And in general, it’s time to think not only about quantitative, but also about qualitative parameters of demographic trends.

05/11/2008 19:29:15, Boris

Religious people say: parental sins are transmitted through blood. Some elders do not advise adopting - it is very difficult. In any case, it depends on spiritual training (how to deal with “blood”?). The case that Nina talks about is not an isolated one.

05/02/2008 12:19:52, Olga

We adopted a boy at six months old. before the age of 7, I don’t have time to turn away, he runs to the trash heap. I didn’t listen as soon as I started talking. I didn’t want to study from the 1st grade. They helped me get a higher education, but I didn’t want to study. He hasn’t worked for 18 years. Now he’s 35 years old. He gave all his strength and health in vain. Nina

26.04.2008 19:56:56

There would be more articles like this. Adoptive parents have to go through trial and error.
It is desirable to have the experience of our specialists and, most importantly, the invaluable experience of experienced foster parents.
The head of a foster family raising 12 children, nine of them are adopted.

13.07.2006 20:10:40, Starostin Sergey

Haven’t the rest of us read it???

By the way, thanks to Irina Shamaeva, we contacted the authors of the Colorado Psychogenetics Research Project (on adopted children) and are now in the process of discussing what’s new and interesting and receiving articles.

2003 marks the 27th year of the Colorado Adoption Project (CAP), classifying it as one of the longest running studies of its kind. The purpose of the CAP is to study both nature and nurture, to determine the genetic predispositions as well as the environmental influences that contribute to traits such as intelligence, personality, and behavior. In order to do this a wide range of interviews are conducted with participating families. These include in-person and telephone interviews that measure cognition, social attitudes, and behavioral choices. The CAP is an ongoing research project of the Institute for Behavioral Genetics,

Finally, at least one intelligible article (and not just for specialists). When you know that almost everything is in your hands, you gain the strength to do even more.
As well as Alexey’s remark that it is not genes that dominate the child, but the parents’ fear of these genes.
Thank you.
R.S. In our particular case, the article helped us make a decision - not to look for biological parents. To nothing.

Previous classes were devoted to hereditarily fixed forms of behavior that allow the animal to carry out the most adequate forms of behavior to a slowly changing or constant environment. These hereditarily fixed or “instinctive” forms of behavior are leading at the lower stages of phylogenesis, and in invertebrates (especially insects) they can acquire extremely complex forms.

Let us recall the main characteristics of these species-specific forms of behavior. Firstly, those programs that determine the behavior of these animals are laid down and fixed hereditarily. These behavioral programs do not need training, each individual is born with these programs, and that is why these programs, despite their complexity, are not only individual forms of behavior (distinguishing one individual from another), but stable species-specific forms of behavior. They are often called instinctive behaviors.

As I said last time, these complex behavioral programs, very well adapted to the constant, little changing living conditions of a given species, are activated by certain signals to which the animal has an innate sensitivity. These signals automatically turn on innate behavioral programs, so that the trigger signal is often very simple, and the programs of innate reactions to this signal are very complex. Thus, the animal has a repertoire of complex reactions to elementary environmental influences. Some psychologists denote these signals with the symbol M, and the behavioral programs they evoke carry out the animal’s adaptation to the environment. In all these cases, the animal itself does not analyze the environment; it captures individual properties and includes ready-made innate forms of behavior. This behavior is characteristic of the stage of the sensory psyche, at which behavior is set in motion by the elementary perception of individual properties of things, characterizing complex programs congenital forms behaviors that are automatically triggered by these signals - this can be called "instinctive" behavior.

It is significant that innate behavioral programs are so well adapted for constant, little changing conditions that they can create the impression of intelligent forms of behavior; however, they turn out to be completely inadequate for rapidly changing living conditions. In this they differ from the rational forms of behavior of higher animals and humans, as well as from those individually variable forms of behavior that develop at the highest stages of development of the animal world and which gradually begin to occupy a leading place.

Thus, the characteristic of the behaviors we have encountered is that they are innate behaviors, evoked by relatively simple sensory signals, and are adapted to constant or slowly changing conditions. A whole large branch of the development of the animal world, the branch of invertebrates (especially insects) follows the path of developing precisely such complex forms of innate behavior, and adaptation to the environment, carried out with the help of species-specific, hereditarily fixed behavior programs, takes a leading place in these animals.

It should be noted that such forms of behavior are adequate only under constant or slowly changing conditions. If the conditions of existence change, this form of adaptation with the help of innate behavioral programs turns out to be very inefficient, and often even inadequate. The main disadvantage of these forms of behavior is the lack of plasticity of these programs. In fact, sometimes it is enough to make very minor changes in conditions for these programs to cease to ensure the adaptation of the animal to the environment. It is for this reason that the described type of adaptation to reality through hereditarily fixed behavioral programs very soon becomes inadequate, and increasingly complex living conditions begin to require new, much more plastic forms of behavior.

However, in the process of evolution, the very type of adaptation of animals to the environment changes radically: innate behavioral programs, which occupy a leading place in insects and lower vertebrates, recede into the background at higher stages of evolution, and in vertebrates, especially in mammals, behavior becomes completely different. another principle.

At this stage of development, living conditions become so complex that it is necessary to develop new forms of adaptation of individuals to changing forms of the environment.

The birth of a large number of young, which sharply distinguishes higher animals and especially mammals from lower vertebrates, suggests a sharp change in the principles of existence with a shift in the plasticity of species-specific forms of adaptation of animals to the changing forms of behavior of individual individuals. It is necessary for such forms of behavior to develop in which an individual would change its behavior depending on changing conditions.

How are these individually variable forms of behavior possible, by what mechanisms are they carried out and by what laws do they proceed? I dedicate today's lecture to the analysis of these more complex individually variable forms of behavior.

Let me dwell on this in more detail. It is often thought that at the lower stages of development there are only innate and unchangeable forms of species behavior, and at the higher stages of development plastic, individually changeable forms of behavior are formed. In general, this is true, but this situation should also be treated with caution. It is known that individual variability of behavior is characteristic of all stages of development, and that individual forms of adaptation of animals to the environment can be observed even in the simplest. Let us remember only those experiments that I presented several lectures ago.

If a shoe is placed in a narrow tube, it goes in the direction in which positive tropism pushes it. But if the sign changes, it begins to go in the opposite direction. To do this, you need to turn over in the tube, which takes a certain amount of time. I told you that if you repeat this experiment many times, then this single-celled organism begins to turn over many tens of times faster; if at first it took him 1.5 - 2 minutes to turn over and go through the tube, then after a series of training it only takes a few seconds. This means that an individual plastic form of behavior develops here.

Another experiment, which we cited above, involves the development of a path that a single cell makes under the influence of conditions that complicate its path to a positively acting agent. If the tube into which the unicellular descends is curved at the end, and the end faces away from the light towards which the unicellular is moving, it is forced to make an arc, not immediately heading towards the path determined by positive phototropism. If, however, after this experiment, the single-celled animal descends into the tube without such a knee, it makes the same loop that it learned, but gradually the loop disappears, and the animal returns to the original direct movement directed towards the agent acting on it.

This means that single-celled animals already have the possibility of individual adaptation, there is individual variability in behavior. An essential feature of this individual, however, is that it is based on a change in the immediate physico-chemical properties of protoplasm: this “skill” does not last long, it disappears very quickly in animals, and it returns to elementary innate forms of behavior.

The transition to multicellularity and the development of a network-like nervous system contributes only relatively little to individual forms of adaptation. Here the speed of passage of impulses in the nervous system only accelerates and the sphere of excitation covers a larger area. However, the predominance of innate - and mostly poorly dissected diffuse forms of adaptation remains.

With the formation of complex types of recipes and the ganglion nervous system, many things change.

Animals with a ganglion nervous system (primarily insects) can functionally perceive a whole complex of stimuli, but practically react only to individual signaling properties that excite innate behavioral programs in them. At this stage of development, innate forms of adaptation continue to be leading.

It would be wrong to think that these animals do not have individual forms of behavior, and such animals cannot be taught anything.

Insects also have individually variable forms of behavior, and they can be trained and retrained, and this is used with great success in agriculture.

It is known, for example, that when it is necessary to retrain a bee to the new kind plants, this turns out to be possible. To do this, this new type of plant is lubricated with a liquid that has the smell of the type of plant to which the bee has an innate system of reactions. After the bee begins to fly to a new plant with the smell of the previous one, this smell is eliminated, and reflexively the bee begins to fly to a new plant that does not have this smell.

This means that bees can be retrained; however, this is a very slow process and does not have the required stability.

What is the limit of this learning and acquisition of an individually variable form of behavior in these animals that have a ganglionic nervous system?

First of all, this retraining can only take place within the limits of instinctive behavior programs.

An illustration can be the experiments of the German researcher Frisch cited last time, in which it was discovered that a bee easily learns to respond to complex figures similar to flowers, but only with great difficulty distinguishes simple geometric figures that have no natural analogues.

This means that it is possible to teach a bee to distinguish a certain form only within the limits of only those innate implementing mechanisms that are hereditarily fixed and that correspond to its ecology. Exactly the same thing happened when Frisch tried to teach bees to distinguish between mixed colors and pure colors. It was very difficult to train a bee to distinguish between pure colors, for example, pure blue from green, black and white, but it was easy for her to develop skills for mixed colors, for example, the difference between yellow-green and green-blue because the flowers she sits, are characterized by mixed and impure flowers.

This means that here, too, individually variable forms of adaptation can arise only within hereditarily fixed species forms of animal behavior.

Similar facts can be observed in birds. For example, it is easy to induce inhibitory lying in wait reactions in a bird of prey, but it is absolutely impossible to induce such inhibitory lying in wait reactions in a chicken, whose ecology does not include the act of lying in wait. Travelers note with surprise that in Antarctica it is absolutely impossible to teach penguins to be afraid of people because in their life repertoire there is no alertness reflex, no fear reflex; on land they do not encounter predators that are dangerous to them, and their repertoire does not include such behavior; therefore, they cannot be taught complex forms of defensive reactions on land.

This means that in all cases, lower animals or insects have the opportunity to change innate forms of behavior, but these changes are limited by the limits of instinctive reactions, which occur very slowly and are very little mobile.

Good examples The relatively low plasticity of behavior can be seen in the experiment conducted by A. N. Leontiev’s employees with the American catfish.

The aquarium was partitioned with a screen of gauze, and fish trying to reach food bumped into this screen. Gradually they learned to swim around this screen, and, consequently, they developed an individually variable form of behavior. But when the screen was removed, the fish continued for a long time to make the arc that they made when going around the screen, although it had now become unnecessary.

This means that even developed changing individual forms of behavior are so inert that they remain relatively unsuitable as forms of adaptation of individual behavior. Thus, at the stage of the sensory psyche, that is, at the stage when animals respond to only one signal that triggers innate programs of instinctive behavior, there is also individual variability, but it is possible only within this innate repertoire of behavior and its limits are very limited. Therefore, the thesis that the sensory psyche and the corresponding instinctive forms of behavior are good under constant environmental conditions remains valid.

Therefore, it is clear that for animals living in more complex conditions of a rapidly changing environment, a leap to another level of behavior is necessary. The most significant thing for this leap is the need to change the form of reflection of reality that the animal has.

In order for an animal to take into account changing environmental conditions and react with a rapid change in behavior and the corresponding environment, it is not enough for it to reflect only individual signals that set into motion the innate repertoire of behavior. For more complex forms of plastic individual behavior, analysis and synthesis of environmental conditions becomes necessary. To do this, it is necessary that the animal reflects not individual properties, but entire objects, entire objects, entire situations, a whole complex of properties; in order to analyze these changing environmental conditions, the animal is able to navigate the reality around it and develop those forms of behavior that would be corresponding to a specific subject environment.

Naturally, this requires the emergence of apparatuses that would not only make it possible to isolate individual properties, but would also make it possible to analyze environmental conditions; it is necessary that, on the basis of the analysis, new, no longer innate, but individual conditioned reflexes and programs that correspond to changing environmental conditions can be created. Finally, it is necessary that animals that develop new, changeable behavior programs can compare the results of actions with environmental conditions, detect erroneous actions in time, change these erroneous actions into correct ones in time, and thereby provide the necessary forms of plastic, changeable behavior.

All this requires a leap to completely new forms of nervous mechanisms and, above all, to the apparatus of the cerebrum and its cortex.

This new apparatus is built on top of the elementary ancient brain and, at the stage of evolution that interests us, becomes the main apparatus of individual forms of changeable behavior.

In the figure below () I give a schematic diagram of the three main stages in the development of the nervous apparatus and behavior of vertebrates. At the top is a frog's brain, in the middle is a lizard's brain, and at the bottom is a schematic diagram of a mammal's brain. Let's take a closer look at how this scheme is built. The eye receives a certain signal, this signal is transmitted to fibers that go to the midbrain; in the midbrain, the excitation reaching these fibers immediately switches to another cycle of fibers conducting motor impulses; the elementary reflex closes. A frog that sees the fluttering or flickering of a fly or piece of paper immediately makes a jump and captures the prey. This is an elementary reflex that occurs as an innate implementing mechanism. Reflexes catch the flicker, the caused excitation reaches the midbrain, immediately switches to motor neurons, and the innate program of jumping and grabbing is activated. Note that a frog that has only these mechanisms is very difficult to retrain, it is very difficult to develop inhibition of this innate reflex, and, perhaps, it is impossible to develop new forms of behavior under these conditions.

We see a completely different structure when we move on to the next large stage of reptiles (snakes, lizards). Here, to the floor that only exists in the frog, at least two new floors are added. Visual stimulation goes here along the visual fibers, which bifurcate; part of the fibers goes to the midbrain, in the quadrigemina, and here and in the lizard (as well as in the frog) there is a rapid switching of this excitation to the motor apparatus, but the other - and at that the main - part of the fibers goes to the formations of the diencephalon (subcortical visual centers) - this path is not a dead end; from it, the visual fibers go further and are sent to the cerebral cortex, which is a powerful apparatus that provides the most complex processing of the information received. Here they communicate with fibers carrying information from individual receptors and enter into the most complex neural apparatus, which provides the most complex forms of analysis and synthesis of these stimuli.

This mechanism appears even more clearly in higher vertebrates, in mammals. Only single branches of optic fibers go to the quadrigemina and therefore elementary visual reflexes no longer play a decisive role here. Only the simplest visual functions - the simplest components of the orientation reflex, the pupil constriction reaction - are carried out by this mechanism; the main pathway of the visual system is different here. the fibers of this visual pathway go to the external geniculate bodies and further - as part of the visual radiance - to the cerebral cortex, where the vast majority of visual fibers pass.

What happens in the apparatus of the visual cortex of the brain?

It is known that a number of fibers come to the occipital region of the cerebral cortex (Brodmann area 17), which end in the 4th layer. These are afferent fibers. The excitation brought by these fibers is only partially transmitted to the afferent pathways, thereby forming relatively simple visual reflexes. Most of these fibers, through the so-called interneurons (or stellate cells), transmit excitation to other cortical cells, forming constantly circulating circles of excitation, which form the basis for the organization of visual processes at a higher cortical level.

This circulation mechanism in the cerebral cortex allows for the implementation of complex functions of analysis and synthesis, that is, that form of behavior that is necessary for the animal to develop complex individually variable behavior programs that would correspond to the perceived object of the external world and the whole changing situation, in which the animal turns out to be.

Unlike the elementary levels of the nervous system, where neurons form a nucleus, usually consisting of elements belonging to the same modality and arranged in a random order, the cerebral cortex is built from at least three types of completely different neurons; afferent neurons that bring excitation from the periphery; switching or stellate, which take over excitation from afferent ones, enable excitation to circulate for a long time throughout the cerebral cortex and transmit this excitation to the third type of neuron - efferent neurons. On the other hand, these neurons, the number of which is enormous (you know that in the human cerebral cortex there are about 12 - 14 billion of them), unlike ganglia, are not located randomly, but in an organized manner along the plane. The cerebral cortex is a screen that has a six-layer structure and which allows excitation to circulate spatially, to a certain extent reflecting the structure of those stimuli that come from the outside world. In some areas, which are called primary projection areas of the cortex, elements of the fourth afferent layer occupy a predominant place. In other sections, the afferent layer of cells disappears and occupies a very insignificant place; but a huge amount of space begins to be occupied by the second and third layers of cells, which in their function are associative cells.

It is interesting that in different animals these layers are not equally developed, and the higher an animal is on the evolutionary ladder, the more developed its upper associative layers of the cortex become.

The third distinctive feature of the cerebral cortex is very significant. The structure of individual sections of the cerebral cortex is not the same.

The cortex is divided into a number of completely differentiated fields, which you have already studied quite clearly in the course of anatomy and evolutionary morphology.

At the lower stages of development, those areas of the cortex where visual, tactile, and auditory stimuli come are located very close to one another, actually overlapping each other. This means that the cerebral cortex of lower animals, say, a hedgehog, represents the possibility of direct synthesis of auditory, visual, and tactile stimulation. In more organized animals, the apparatus, where the fibers from the visual, tactile and auditory receptors reach, are moved apart and special secondary, tertiary fields are inserted between them; These fields consist mainly of neurons or zones into which fibers come, coming from those nuclei of the thalamus opticum, which themselves are not directly connected to the periphery and which themselves receive already processed impulses.

It is interesting that the higher an animal stands on the evolutionary ladder, the more space the secondary and tertiary fields occupy in its cortex and the more the primary projection fields are relegated to the background.

Let us turn to the last feature in the structure of the cerebral cortex, which is of significant importance for us.

We have already said above that different sections of the cerebral cortex are related to different receptors (visual, auditory, tactile). However, the place occupied by these zones varies among animals with different ecology. The greater the role played by one or another receptor, one or another analyzer in the behavior of an animal, the more it is represented in the cerebral cortex. In other words, the representation of individual receptors (visual, auditory, tactile) in the cerebral cortex is built not on a geometric, but on a functional principle. Therefore, in the hedgehog, in whose behavior the sense of smell plays a leading role, the olfactory areas of the cortex are represented by a particularly large area, occupying approximately two-thirds of the hemispheres. In the monkey - which is mainly oriented to visual stimuli - the visual sections of the cortex occupy a particularly large place; finally, in humans, in whom, as we will see later, the leading place is occupied by complex forms of reflection of reality mediated by speech, the overwhelming part of the cortex belongs to the secondary and tertiary zones, which carry out the most complex forms of cortical activity.

To illustrate the point that the leading analyzer is especially strongly represented in the cortex, one example can serve.

It is known that in a pig, the organ with which it perceives the outside world is the snout, and in sheep it is the lips, with which they get acquainted with food before putting it into the mouth. The pig's snout and the sheep's lips play the same role as the hand does in humans. This corresponds to very interesting features of the cerebral cortex.

When the famous English physiologist Adrian tried to trace which parts of the brain of different animals react to irritation of different parts of the body, he found that the largest area in a pig is represented by its “patch”, and in a sheep its lips.

This is an excellent illustration of the fact that these organs of the body are represented in the cerebral cortex in a manner inconsistent with the role they play in the behavior of the animal.

Everything we have said shows that the cerebral cortex of vertebrates and especially higher vertebrates is specially adapted not only for receiving signals, but also for analyzing and synthesizing complex signals.

The cerebral cortex allows you to reflect a complex of signals, highlight the essential, inhibit the unimportant, reflect entire objects and entire situations, and program complex variable forms of conduction. This is the most important function of the cerebral cortex. And that is precisely why the cerebral cortex, which greatly increases in the animal series and plays an increasingly important role, should be regarded as a mechanism that allows an animal to move from receiving individual signals that set in motion innate forms of behavior - to an apparatus that allows one to analyze and synthesize complex irritation and providing a transition from the sensory psyche to the perceptual psyche, reflecting entire images.

In the process of evolution, the proportion of this organ, which makes it possible to move on to the analysis and synthesis of external stimuli and to implement individual forms of behavior, increases more and more. In this regard, more and more functions begin to depend on the cortex. Researchers call this rule the movement of functions to the cortex or the law of “progressive corticalization of function.”

The fact that at higher stages of evolution more and more functions depend on the cerebral cortex can be proven by simple facts. If you cut out a bird's still poorly developed cerebral cortex, it continues to see, can avoid obstacles, and sit on platforms. If you cut out the cerebral cortex of a rat, it continues to distinguish light, although it loses the ability to distinguish shapes; a monkey whose bark is cut out turns out to be blind. A bird with its bark cut out can fly as smoothly as before; If you destroy a cat’s cerebral cortex, then after a few hours fairly smooth movements will return. This means that smooth movements in a cat do not yet depend on the cortex, but are carried out by subcortical apparatuses. A dog whose bark has been removed can stand after 24 hours, but cannot move independently; movements do not return to a monkey with the cortex cut out; In humans, damage to small areas of the motor cortex causes complete paralysis.

Consequently, indeed, in the process of evolution, the regulation of functions begins to depend more and more on the apparatus of the cerebral hemispheres.

Let us now move on to the main thing, for which we recalled the given anatomical data. At a certain stage of vertebrate development, the elementary cortical ganglia are supplemented with new level: cortex. This marks a decisive shift in animal behavior. if the subcortical ganglia is an organ of inherited behavior that allows you to perceive signals and turn on inherited behavioral programs, then the cerebral cortex is an organ that allows you to analyze and synthesize external stimuli, reflect objects and reflect entire situations, make new connections and build behavior no longer according to innate ones , embedded hereditary programs, and according to these conditions of the objective environment that the animal perceives. Therefore, with the development of the cerebral cortex, fundamentally new possibilities for individually variable forms of behavior arise, which are based on the analysis and synthesis of signals coming from the external world, on the reflection of entire images of the objective external world.

It is in this regard that the stage of the psyche that I will now talk about, in contrast to the sensory psyche, can be called perceptual, that is, the psyche, which is based on the reflection of not individual properties, but entire objective situations, which makes possible the formation subjective image of the objective world and allows one to adapt behavior to these incomparably more complex forms of reflection.

It is at this stage of the sensory psyche that the leading role shifts from innate behavioral programs to the analysis of rapidly changing forms of the environment and to the development of individual variable forms of behavior.

The more developed the cerebral cortex, the more the animal begins to analyze the external environment and respond accordingly to the analysis of this environment.

The physiological mechanisms of these individual forms of behavior are well known from the physiology of higher nervous activity; they come down to the processes of analysis and synthesis and the formation of temporary connections, well studied in the school of I. P. Pavlov. Therefore, I will not dwell on them and will briefly mention only those forms of behavior that arise at this stage.

What experiments can be used to show that animals with a well-developed cerebral cortex not only pick up signals, but also analyze entire situations and retain a certain image of external activity? Experiments showing these facts were carried out by many researchers and were called experiments with “delayed reactions”. In our country, among others, they were carried out for a long time by an excellent Soviet psychologist, the late Professor N. Yu. Voitonis.

If you place two boxes in front of the animal at a distance of several meters and put a piece of meat in one of the boxes (hermetically sealed) in front of the animal’s eyes, close the box and then hold the animal for 2 - 3 minutes and only then release it, you can observe an interesting picture. If the animal retains the image of the object, then it will definitely run to the box where the bait is placed. If the image of the object is not preserved, then it may accidentally run to either one or another box.

As research has shown, these experiments gave a clear result; the animal, as a rule, reacted to the place where the object was hidden. This means that it is clear that the animal retains in memory where the object was hidden and walked towards it. If the object moved, then the animal went to another place corresponding to this object.

Is this similar to experiments with insects? Not at all. These experiments show that the animal easily changes its reactions depending on the movement of the reinforced place, without revealing any inertia. Moreover, it turned out that the animal retains not only place cues, but also a clearer image of the object that was placed there.

The experiments that confirmed this were carried out as follows; in front of the animal's eyes, one bait was placed in the box, and then, unnoticed by the animal, it was replaced with another bait. Sometimes the animal was released, it ran to the box, opened it and saw a bait that was equivalent in taste to the first one, but different in appearance. This is where an interesting feature emerged; the animal turned away and ran to look for the bait it had seen before. This means that the animal retained the image of the bait, the image of the stimulus, and this image was selective in nature. These experiments led to very interesting conclusions. Consequently, at this stage of development, the external environment, which is perceived by the animal, causes it to focus on a certain image of the expected object. If the image of the found object coincided with the expected one, the action was stopped, and if the image of the found object did not coincide with the image of the expected one, the search continued. In the latter case, they said that the real image was inconsistent with the expected one, but the first time it was consistent with it. The behavior of the animal was determined by this agreement or mismatch between the image that the animal expected and the image of the object that it received. Thus, the behavior of the animal at this stage was characterized by a completely different structure; the behavior program began to be regulated by this image of objective reality, and the effect of behavior was a comparison of the received image with the expected image.

Thus, the entire behavior of the animal at this stage acquires an incomparably more complex and qualitatively different character than it had at the stage of the sensory psyche and instinctive behavior. The signal here begins to evoke not innate forms of behavior that do not change from environmental conditions, but causes an analysis of the situation, creates a certain subjective image of the objective world; this image causes the corresponding program of action; if this program of action leads to the expected action, the action resumes and continues.

Thus, the action of the animal here is based on the reflection of complex objective reality and adapts to it. Therefore, this form of reflection is called perceptual. This action changes depending on the signals that the animal receives from the environment. Therefore, it is not called instinctive, but individually changeable behavior.

The higher an animal is on the evolutionary ladder, the more complex images it can perceive and the longer the influence of these images lasts.

As studies of delayed reactions have shown, the effect of such an image in a rat lasts 10 - 22 seconds, in a dog - about 10 minutes; the monkey can maintain the image for 16 to 46 hours. The higher the animal, the more stable is the image that the animal has formed, and the more durable is its regulating influence.

However, the stage of the perceptual psyche differs not only in that the animal reacts to a complex set of stimuli and reflects images of the objective world. It also differs in that the animal programs its behavior in a completely different way than the behavior was programmed at the instinctive stage.

New forms of behavior arise, which many researchers called skills or habits or learned forms of behavior. An example of such a skill would be behavior that was carefully studied by one of the Soviet psychologists, Professor Protopopov. He calls this behavior under “stimulus-obstacle” conditions.

In these experiments, the animal is given a known signal, but it is given in difficult conditions - usually placed behind an obstacle. The animal must reflect the conditions in which it is placed and adapt its behavior to them. Facts have shown that if such an experiment is carried out with an animal that is at a lower stage of development, in which the cerebral cortex is still very underdeveloped, then the type of food triggers in the animal a direct hereditary program of behavior that does not take into account the conditions in which this stimulus is presented. Therefore, the chicken, which perceives the grain through the mesh, begins to beat against this mesh, and its behavior is entirely determined by the direct instinctive reaction program. But a dog or monkey in a stimulus-obstacle situation acts differently; she first tries to directly get the bait, and then runs around the obstacle and gets the bait from the other end. This means that if a chicken’s behavior in stimulus-obstacle conditions is limited to the inclusion of instinctive behavior programs, then in higher animals, in a dog, the action is not determined by a hereditary program, but is programmed according to the analysis of the situation.

Consequently, here we have individually variable behavior, and behavior changes depending on changes in the perceived external situation.

The higher the vertebrate, the more place is occupied by these complex individual forms of perceptual behavior based on an analysis of the situation, and the less place is occupied by forms of instinctive behavior. Physiological mechanism You study these individual forms of adaptation in the course of higher nervous activity, and we will turn to some behavioral mechanisms of skill formation next time.


The main material obtained in the initial period of studying higher nervous activity was based on the analysis of the autonomic conditioned reflex - salivation, caused by a variety of external signals that were not previously associated with this type of activity. Today it is called a classical, or conditioned reflex of the first type, which is a copy of the unconditioned one. Based on innate motor activity, conditioned reflexes of the second type (instrumental) are formed. They can correspond to the unconditioned ones (example: moving away from the flame of a fire before it burns, i.e. in response to the action of light and other conditioned stimuli) or differ significantly from them (example: standing on the hind legs, reinforced by giving food). Usually, a complex of instrumental and classical conditioned reflexes is formed simultaneously, i.e. the same signals cause a motor response and a variety of reactions from the respiratory, cardiovascular and digestive systems. The formation and extinction of these conditioned reflexes often occurs at different times.

A complex of unconditioned and conditioned, classical and instrumental reflexes fixed over a long period of time forms a system called a dynamic stereotype. It is produced with difficulty and has a certain inertness. Automatic skills associated with this stereotype can be gradually extinguished when living conditions change. Based on where the signal comes from to the nerve centers - from the external or internal environments, exteroceptive and interoceptive conditioned reflexes are distinguished. In the first case, visual, auditory, gustatory, olfactory, temperature, tactile conditioned reflexes are distinguished, in the second - mechanical, chemical, osmotic, temperature. In general, interoceptive conditioned reflexes (associated with irritation of internal organs) are developed more slowly than exteroceptive ones. However, this pattern is relative. The formation of conditioned reflexes is determined by the correspondence of this act to the natural relationships of the body and the external environment. For example, an innate change in the choice of certain foods, which occurs when the chemoreceptors of the stomach and intestines are irritated, very quickly becomes a conditioned reflex and takes place already upon stimulation of the mechano-receptors of these organs.

Here we should dwell on the category of natural conditioned reflexes, which have much in common with imprinting. They are associated with the action of environmentally adequate stimuli. It has been shown, in particular, that the smell of meat does not cause an innate reaction of salivation in a puppy, but after a single feeding of raw meat, a conditioned reflex secretion of saliva to its smell is produced, which does not disappear throughout life. Unlike natural ones, artificial conditioned reflexes, which are based on a connection between stimuli and innate reactions that are not connected by natural relationships, are formed slowly.

Conditioned reflexes can arise when combined with reinforcement of simple signals, their complexes (sound + light), and sequential stimuli. When the signal and the unconditioned stimulus coincide in time, a conditioned reflex is developed. If you regularly, at certain intervals, give reinforcement, then later, after their expiration, an unconditioned reaction (conditioned reflex for time) appears without any additional influences.

The central nervous system is constantly bombarded with signals from the external and internal environments. The cerebral cortex simultaneously evaluates these signals in a given life situation. Consequently, the reactions caused by conditioned signals are determined by this assessment, and if the signals did not cause the expected effect, then one integral state was replaced by another (for example, there was a transition from food to defensive or sexual behavior). Typically, such a change depends on many stimuli (pain, noise, smell, etc.). Their appearance during dog training can seriously complicate the development of the necessary skill. External (unconditioned) inhibition in the higher parts of the brain plays a significant role in this. Thanks to it, the current activity stops, indicative-exploratory behavior manifests itself and the transition to another type of activity. External inhibition is thus the most important regulator of the relationship between the organism and the environment, since it “turns the needle” towards the most important form of behavior in a given situation.

In the case of the action of super-strong stimuli, extreme inhibition develops in the cells of the cerebral cortex, which in extreme cases manifests itself in complete immobility - stupor. This condition can interrupt the dog's performance of an important task. It is possible to increase the performance of cortical cells by introducing pharmacological substances, thereby preventing the development of extreme inhibition. These include caffeine, phenamine, sydnocarb, combined with tranquilizers. Individual selection of these drugs and their dosage is required. In some cases, their administration can lead to disruption of higher nervous activity. It is also difficult to predict what long-term consequences are possible with one or another pharmacological effect.

Internal (conditioned) inhibition plays an important role in the regulation of acquired adaptive activity. If a dog with a well-developed food salivary conditioned reflex is given a conditioned signal without reinforcing it with feeding, then the secretion of saliva will gradually decrease. The internal inhibition underlying this phenomenon is called extinction. It manifests itself in waves - the inhibited reflex can appear again.

Another type of internal inhibition, differentiation, eliminates reactions to non-reinforced stimuli that are close to the reinforced one, which makes it possible to more accurately tune behavior to a biologically important stimulus. In puppies, this type of inhibition matures gradually, so they often exhibit a generalized (generalized, nonspecific) reaction. Adult dogs differentiate between significant and insignificant signals well (manifestations of this phenomenon in dogs are colorfully described by K. Lorenz).

Internal inhibition also includes conditioned inhibition - the absence of a reaction to a positive conditioned signal combined with an indifferent stimulus. Thus, if the sound signaling food is combined with the application of a cold plate to the skin, conditioned reflex salivation is not caused.

Currently, the mechanisms of the conditioned reflex are well studied using electrophysiological and neurochemical methods. The unity of the mechanisms of external and internal inhibition has been established, and it has been proven that inhibition is the same active process as excitation. The elucidation of the mechanisms underlying conditioned reflex activity has intensified research into such an important component as memory.

3.5. Mechanisms of memory

The behavior of higher animals would be impossible without the use of past experience, i.e. without storing information about it in the nervous system. Sensory memory manifests itself in the retention of a trace of receptor stimulation in a neuron for a very short time - up to 0.5 s. Erasing a trace takes 0.15 s. Longer retention of information about current events is determined by short-term memory. It is studied mainly using delayed reactions. The animal develops a conditioned reflex, after which a pause (delay) is introduced between the signal and response associated with it.

Long-term retention of information is associated with long-term memory. The transition from short-term to long-term memory, i.e. consolidation of a memory trace in the central nervous system is carried out using intermediate memory. The mechanisms underlying these phenomena are still far from clear; it has been shown, however, that they are determined by electrical factors (circulation of nerve impulses) and structural and chemical changes in the central nervous system.

It is customary to distinguish different forms of long-term memory: figurative (preservation in the brain of signs of objects that the animal has encountered), emotional (dog breeders know well how long dogs retain a negative attitude towards people who have caused them harm) and conditioned reflex, responsible for the reproduction of certain motor and secretory acts.

A large amount of experimental material has been obtained characterizing the mechanisms of action of biologically active substances produced by the body on various forms of memory. One might think that by pharmacologically increasing or decreasing the balance of these substances and their effect on neurons, we can improve memory, and therefore increase the level of the dog’s performance. This is partly true, but we should not forget that the same pharmacological effect has different effects on different aspects of learning. In addition, it is difficult to predict the long-term consequences of introducing chemicals into the body.

Data obtained in laboratories on the influence of chemical compounds that have a pharmacological effect on memory do not yet provide a sufficiently complete answer to the question of the neurochemical mechanisms of memory. Psychostimulants and tranquilizers, which are often recommended to improve learning, can sometimes, on the contrary, cause retrograde amnesia (forgetting previously stored information). It is known, however, that memory is closely related to the functioning of such biologically active substances contained in the brain as acetylcholine, norepinephrine, dopamine, serotonin, gamma-aminobutyric acid (GABA), and peptide hormones. It is incorrect to associate any one neuronal function with each of them.

One should also take into account the uniqueness of the neurochemical characteristics of different parts of the brain and their role in organizing certain forms of learning. In addition, we evaluate memory by the duration of the manifestation of various skills, but we can only indirectly judge the preservation of traces of certain influences in the nerve centers. At the same time, it is known that memory processes are associated with the central metabolism of biologically active substances, changes in which improve or worsen the development of a skill and its retention. Particular attention in the study of these processes is paid to brain monoamines - dopamine, norepinephrine, adrenaline and serotonin. It is shown that their balance largely determines the speed of learning and stabilization of the memory trace. Often, an increase in the content of one of these compounds in the brain has different effects on different forms of learning. Thus, with the development and strengthening of defensive reactions, the exchange of norepinephrine in the brain increases, and that of food - decreases. The administration of drugs that increase the content of serotonin in the brain significantly improves the learning process with food reinforcement, but worsens it with pain. Serotonin accelerates learning and promotes the preservation of conditioned reflexes developed under an emotionally positive background, acting in the opposite way in the formation of reactions (mainly defensive) accompanied by negative emotions.

Acetylcholine, GABA, and glutamic acid also play a significant role in memorization. The use of pharmaceutical drugs that affect the metabolism of these substances can affect learning and memory. However, let us once again recall the difficulties associated with the individual characteristics of the animal, the selection of appropriate doses (overdose often leads to the opposite effects), and the lack of sufficient knowledge about the neurochemical basis of memory.

The introduction of any pharmacological agent into the body causes a complex chain reaction. Data on the participation of neuropeptides in the regulation of memory processes are very interesting. The latter are quickly destroyed, and it can be assumed that their action is mediated through a cascade of biochemical reactions that determine a wide range of changes in the state of the central nervous system. Neuropeptides have been identified that have a significant effect on learning and memory. These include vasopressin, cholecystokinin, neurotensin, angiotensin and many others. Particular importance is attached to opioid peptides - endorphins and enkephalins. Some of them improve, others worsen memory. It also happens that the same opioid peptide stimulates the development and maintenance of a conditioned reflex in poorly learning animals and inhibits it in well-learning animals.

Thus, at the current stage of development of the science of memory, it is impossible to unconditionally recommend the introduction of drugs into the dog’s body that improve learning and memory, for several reasons. Firstly, their mechanism of action is largely unclear. Secondly, while they may have beneficial effects on some types of learning, these influences may hinder the formation and retention of other equally important skills. Thirdly, an artificial imbalance of biologically active substances in the body can disrupt normal functioning and lead to long-term changes in the behavior and activity of internal organs. Fourthly, many of the substances mentioned are still very expensive and are used in limited quantities even for experimental purposes.

3.6. SYSTEMIC BRAIN ACTIVITY

The functional state of the central nervous system continuously changes in accordance with changes in life activities. It is determined by posture, position of the limbs, and many signals from the external and internal environments. At any given moment, a special system of processes of excitation and inhibition develops in the brain and its individual structures, which determines the behavior of the organism under given conditions.

To understand the mechanisms by which the central nervous system controls the internal economy and activity aimed at interacting with the outside world, one should consider some of the main aspects of its systemic activity associated with the formation of complex forms of behavior.

3.6.1. The principle of dominance

A. A. Ukhtomsky created the doctrine of the dominant (from the Latin dorninantus = dominance) - the temporary predominance of excitation in nerve centers that are functionally united to carry out a certain activity. The focus of excitation, as it were, attracts streams of nerve impulses from extero- and interoreceptors and thereby predetermines the body's reaction aimed at a certain result. A “functional center” arises in the central nervous system, uniting departments that are distant from each other. Several foci of increased arousal may occur at the same time, making it possible to quickly switch from one activity to another. But at any given moment, the most persistent focus of excitation determines the direction (vector) of a certain activity of the body. Thus, stretching of the bladder creates a dominant focus of excitation associated with urination. Against this background, impulses coming into the central nervous system from different receptors support this focus and increase the level of its excitability. It drops sharply after the corresponding act - emptying the bladder.

Dominant is an important principle of the central nervous system. When training, it is necessary to remember this, since the development of a skill can be seriously affected if during training the dog develops a dominant focus associated with another type of activity. Switching of dominants can, for example, take place when an individual of the same or opposite sex appears (inhibition of the worker and manifestation of a new dominant aimed at sexual, play or defensive activity), with a sharp shout or with painful irritation.

Changes in the functional state of the body (estrus, pregnancy, lactation) are associated with the formation of long-term, persistently persistent dominant foci of excitation. A similar phenomenon is observed with insufficient nutrition, selective deprivation of any food product necessary for the body, and fatigue. Therefore, against such a background, developing skills that are not related to the current dominant state of the central nervous system is ineffective. Training is successful when the reinforcement corresponds to the character of the dominant (for example, when developing salivary or food-procuring conditioned reflexes in an unfed dog). In this case, a conditioned reflex can be developed even to painful stimulation. So, in the laboratory of I.P. Pavlova M.N. Erofeeva gradually converted the defensive reaction in a hungry dog ​​caused by the action of electric current into a food reaction. She reinforced the electric shocks with feeding, and eventually they stopped causing the dog to have a negative attitude towards the experimental room and the desire to chew the electrical cord, break free and run away. After several experiments under the influence of hunger, the defensive reaction began to gradually slow down. Further, when the dog lost significant weight and the dominant hunger sharply manifested itself, the electric current began to cause salivation and the entire repertoire of food-procuring behavior with signs of a positive emotional state. This example illustrates the plasticity of the central nervous system and the ability, against the background of a certain dominant, to transform defensive conditioned reflexes into food ones (we will see later that reverse transformations are also possible).

3.6.2. Conditioned reflex switching

The same conditioned stimulus, depending on the situation against which it operates, can be a signal for the manifestation of different types of activity. In fact, the above experiments with a change in the signal value of the electric current can serve as one of the examples of switching. The fact is that in these experiments the food reaction was caused by painful irritation of a certain area of ​​the skin in a certain environment. Outside the experimental room, the electric shocks caused the dog to react normally defensively.

To develop conditioned reflex switching, some kind of light or sound stimulus is usually used, the signal value of which changes depending on additional circumstances. For example, one experimenter develops a salivary conditioned reflex in response to the knocking of a metronome of a certain frequency during food reinforcement, while another, in response to the same signal, develops a defensive reflex, reinforced by electrocutaneous stimulation. A few days later, in experiments conducted by the first experimenter, the dog responds to the beats of the metronome by salivating, and in experiments conducted by the second, by withdrawing its paw. Time can also be a “switch” of the nature of the conditioned response: different conditioned reflexes can be developed to the same stimulus in the morning and evening hours. Conditioned reflex switching can occur within the same activity. If you reinforce the metronome beats with food in the morning and do not reinforce it in the evening, then this stimulus causes salivation in the morning and inhibits it in the evening.

The young cocker Don Quixote was home trained by his young owner to show stand. Don Quixote was immediately exhibited at a prestigious exhibition abroad, and behaved disgustingly. This is how the owner’s unforeseen conditioning of the learned exhibition pose by the home environment manifested itself. Experienced dog breeders usually practice posing in the ring at ordinary exhibitions, during which they do not so much show the dog as train it to posing in real exhibition conditions.

Observing the behavior of dogs, we are faced with the phenomenon of switching at every step. Yes, dog, completed the course general training, reacts positively to food offered by the owner (salivation, food-procuring movements, tail wagging), but refuses to take it from strangers, showing signs of a negative emotional state; to the same command “forward!” The dog reacts differently in front of a boom, a barrier and a ladder.

Of great interest is the so-called dissociated behavior, which can also be considered as switching. This form of training has been studied in the development of conditioned reflexes against the background of the introduction of pharmacological substances that change the functional state of the central nervous system. It is possible, using the same stimulus (light, sound, etc.), to develop, for example, against the background of caffeine administration, a food-procuring conditioned reflex, and against the background of small doses of strychnine, a defensive one. In the future, such an irritant will cause salivation or withdrawal of the paw, depending on the background created by the preliminary administration of one of the pharmacological drugs.

The given examples of complex forms of learning are well explained within the framework of the doctrine of the dominant. The state of the central nervous system, determined by the environment or internal factors, is the basis on which goal-directed behavior is built.

3.7. FACTORS THAT PROMOTE AND OBSTACLE LEARNING

3.7.1. Indicative-exploratory behavior

Above, attention was already drawn to the difficulties that arise when trying to draw a clear boundary between innate and acquired forms of behavior during life. These difficulties are associated with the fact that the former undergo significant changes in the process of life. The plasticity of unconditioned reflex reactions is determined by various factors and, above all, by the influence of developmental conditions in the prenatal period and in the first stages of life after birth on the deployment of the genetic program. The characteristics of the formation of many components of the puppy’s body, including the development of its central nervous system, and, consequently, many aspects of behavior depend on the nutrition received by the puppy, on the state of its endocrine organs, and on the external signals falling on it. The environmental conditions surrounding it after birth also play an important role. Thus, heritable reactions may manifest themselves differently depending on the influences on the organism in the early stages of life.

Unconditioned reflex reactions can also change significantly in subsequent age periods. Learning can influence all aspects of innate activity, for example, perception (conditioned reflex reduction in receptor sensitivity thresholds). Immediately after birth, inherited reactions begin to “overgrow” with conditioned ones. This is why it is so difficult to separate inherited and developed forms of behavior. The combination of these forms is very clearly manifested in orientation-exploratory behavior, one of the most important manifestations of higher nervous activity.

The orienting reflex occurs when exposed to new stimuli. Outwardly, it manifests itself in a reorientation of attention to them and the suspension of current activities. I.P. Pavlov called this mechanism the “what is it?” reflex. and proposed to distinguish the reflex of “biological caution” and the actual exploratory reflex - the movement of the animal towards a new source of irritation in order to better understand and evaluate it. Yu. Konorsky designated the orientation reflex as a “targeting” reflex, since it helps to configure the senses for optimal perception of a certain stimulus.

The indicative reaction is an integral part of conditioned reflex behavior. It increases the excitability of the cortical and subcortical structures of the brain, which is necessary to “close” the temporary connection between an indifferent stimulus and unconditional reinforcement when they coincide in time. One of the properties of this reaction is the ability to be extinguished upon repeated presentation of the stimulus. When this reaction is extinguished, the development of a conditioned reflex becomes significantly more difficult.

In puppies, the indicative reaction appears immediately after birth. From the first hours of life, they respond positively or negatively to odor stimuli. From the 10-15th day, the generalized form of the indicative reaction is replaced by local movements - turning the head towards the stimulus and subsequent elements of its examination. Following the olfactory, gustatory and tactile analyzers, other sense organs gradually mature - hearing and vision. An approximate reaction to sound stimuli appears on the 4-6th day of life, when the ear canals are still closed; it becomes more definite on the 15th day. In relation to light stimuli, a clear indicative reaction occurs at the age of 15-18 days. It gradually becomes more complex and by the age of one month reaches the level characteristic of an adult dog.

Usually we pay attention to the motor manifestations of this reaction. In fact, it also includes somatic and autonomic components. At the same time, such of them as a decrease and then an increase in respiratory movements, an increase in the number of heart contractions reflect the body’s preparation for the best implementation of a motor reaction. Such a complex allows the animal to quickly assess changes in the situation and react correctly to them. This determines the most important role of the orienting reaction as the basis for ensuring active adaptation to the constantly changing external environment.

In the process of development, the orienting reaction becomes more complex. From the 1st to the 15-25th day of a puppy’s life, it is associated mainly with eating behavior. For example, any touch to a puppy causes him to suck, smack and move towards the source of irritation. This age period is characterized by a state of prolonged excitation of the food center, supported and reinforced by any irritations in accordance with the principle of dominance. Food dominance, expressed in the appearance of a food search reaction in response to any irritation, has great biological significance - it facilitates the mother’s finding, the act of sucking and ultimately ensures survival and normal development puppy.

As already noted, as a result of the newborn puppy’s encounter with new stimuli, the restructuring of innate behavior begins already in the first hours of life. A functional complex is formed in which congenital and acquired components closely interact. In this case, the orienting-exploratory reaction is organically included in the orienting-exploratory behavior.

3.7.2. Passive defensive behavior

In the process of evolution, different forms of behavior have arisen, each of which has important adaptive significance. One of them is passive-defensive behavior or, as it was called in Pavlov’s laboratories, the “biological caution reflex,” a self-protective mechanism that is replaced by exploratory behavior as one becomes familiar with the external environment. It is essential for adaptation to the action of unfamiliar stimuli and, with moderate manifestations, cannot be attributed to pathology. However, it is bordered by a high level of inhibition and the development of widespread sleep inhibition. In these cases, it is difficult to use the animal for work that requires a good orientation reaction and weakly expressed passive-defensive behavior. These qualities should be inherent in dogs of many breeds (shepherds, Airedales, Dobermans, pointers, St. Bernards, etc.). At the same time, their manifestation is determined not only by the genotype, but also by the conditions of upbringing. Optimal care for your puppy in the early stages of life deserves special attention. Negative effects may not immediately affect the behavior of the animal, but appear after a long time. Thus, in puppies exposed to stressful influences at one month of age, at six months of age passive-defensive behavior sharply manifests itself, slowing down the development of food-procuring skills. With the help of pharmacological interventions, this behavior can be corrected.

In particular, in the above case, such a correction can be made by introducing metamizil, a substance that reduces the content of acetylcholine in the brain and thereby reduces fear. In this way, it is possible to reduce the level of passive defensive reactions and significantly speed up the learning process. The same effect can be achieved by using other drugs that have a sedative (tranquilizing) effect.

It is necessary to take into account the characteristics of animals’ responses to certain stimuli at different periods of development. Weak defensive reactions to sharp stimuli are already observed in newborn puppies (having smelled ammonia or acetic acid, the puppy becomes restless, turns its head away, and whines). Such reactions were probably observed by many breeders when they lubricated the wounds of puppies with iodine or alcohol. Puppies aged 15-25 days require special attention when the color of the indicative reaction changes - it is joined by elements of passive defensive behavior, accompanied by defecation and urination. This type of response to unexpected sounds, light, smells, vestibular and tactile stimulation occurs in almost all puppies before the 40-45th day of life.

The biological significance of including passive-defensive elements in orientation-exploratory behavior is enormous. By the end of the first month of a puppy’s life, its range of vital activity expands. We rarely think about how many dangers await a puppy in the most ordinary environment if his actions were completely random. If previously the puppy was under the protection of its mother, now it is faced with new stimuli, the correct and quick response to which is still difficult due to the imperfection of the sensory organs and motor system, as well as the lack of sufficient life experience. Thanks to the presence of passive-defensive behavior, the puppy avoids many dangers. So, during walks, when there is a sudden horn of a car, headlights, the cry of a bird, or a sharp clap, the puppy usually stops moving and presses against the ground or the owner’s leg. This behavior is normal and does not require pharmacological correction. Puppy shyness should not be confused with cowardice. During this period of development, you need to be patient with the puppy, do not frighten him with a shout, do not pull the leash, and encourage the puppy with affection.

The further development of passive-defensive behavior largely depends on the conditions in which the puppy is raised. Its level decreases significantly in an “enriched” external environment (communication with peers, familiarity with new objects, phenomena, etc.). In the absence of sufficient stimuli (in isolation), the passive defensive reflex is strengthened and can persist throughout life.

From the 40-45th day to 3-4 months (a critical period called the period of “socialization” by the American scientist J. Scott), exploratory behavior reaches its maximum. With proper upbringing, elements of passive-defensive behavior are rarely manifested at this time. However, if the load is excessive, if necessary, decide complex tasks a breakdown may occur - the puppy refuses to work, whines, barks, falls asleep during training.

Passive-defensive behavior in puppies aged 15-45 days is more primitive than in puppies 3-4 months old. In the latter, it manifests itself against the background of complex analytical and synthetic processes in the central nervous system associated with rational activity. These animals are very vulnerable, they solve complex problems well, but at the same time they easily become neurotic. At this age, the typological characteristics of the nervous system are formed, so passive-defensive behavior begins to have a pronounced individual character. At some stage, the puppy notices that his threats by barking, growling, and finally attacking are good defenses in conflict situations. Thus, the passive-defensive reaction is gradually replaced by an active-defensive one, characteristic of many breeds of dogs, especially service dogs.

It is interesting to see the difference in the way the active-defensive reaction develops in dogs of different breeds. Thus, timidity makes an East European Shepherd puppy afraid of everything and everyone. He clings to the protector-owner and is ready to bark all over White light. As he gets older and bigger, he can really scare strangers. A conditioned reaction is developed that associates his attacks with safety. A brave, attacking defender of the owner grows up.

The development of anger in a Rottweiler or black terrier is completely different. Puppies of these breeds are less timid and early acquire a sense of security in the world around them. Often quite a lot of time passes and significant provocation is required for the owner to become convinced that this apparent good nature is the self-confidence of a strong defender dangerous to enemies.

For the correct development of defensive behavior in a puppy at the age of 3-4 months, it is necessary to monitor compliance with the regime, protect it from overload, and, if necessary, use bromides, valerian, Corvalol, Devican and other sedatives.

3.8. MOTIVATIONS

Complex forms of behavior are aimed at achieving certain results, which are associated with the satisfaction of various needs. The desire to satisfy these needs is referred to as drives, drives, or motivations. The motivations of hunger, thirst, fear, aggression, sexual, caring for offspring, interactions with other individuals and many others are described.

Motivation is a state of the central nervous system that underlies goal-directed behavioral acts. The presence of a need does not immediately affect the vector of behavior. This requires its transformation into motivation, i.e. the appearance of a corresponding focus of excitation in the central nervous system. The dominant principle contributes to the formation of motivation based on need - the dog can be very hungry, but the presence of a threat from the enemy or the presence of a female in heat will slow down the movement towards the food object.

It is convenient to further consider the mechanism of motivation using the example of eating behavior. Despite the apparent simplicity of satisfying the body's need for nutrients, the physiological processes underlying the motivation for hunger are extremely complex. Many parts of the central nervous system, located at its different levels, are involved in the regulation of food consumption. This is a functional association of I.P. Pavlov called it a “food center”.

Several subcenters responsible for food motivation are located in the hypothalamus (hypothalamus). In fed animals, electrical stimulation of the lateral sections of this area disinhibits the act of eating; Once they are damaged, the urge to eat completely disappears. Stimulation of the median nuclei of the hypothalamus inhibits food consumption, and their destruction leads to gluttony and obesity. Similar phenomena are observed during infectious processes or tumors affecting these areas of the brain.

The hypothalamus has extensive nerve connections with many parts of the brain. The process of excitation from the motivational centers of the hypothalamus spreads to these departments, due to which an indicative-exploratory reaction first arises, and then purposeful behavior. An ascending wave of excitation from the hypothalamic parts of the food center changes the functional state of the system that controls all stages of eating behavior: search, examination and absorption of food. Descending activating nerve impulses prepare the internal organs to receive and process it.

Domesticated dogs do not need to get food, like their wild ancestors - wolves. In the latter, food motivation usually alternates with aggression motivation, which can dominate during prolonged pursuit and killing of the victim. Therefore, in wild predators, the direction of behavior associated with obtaining food is determined by the involvement of centers of aggression. In dogs, elements of aggression often appear in isolation, not being associated with food activity.

The role of conditioned reflexes in the formation of food motivation is extremely large, but their action is manifested only under a certain state of the body. Thus, a well-fed dog will not respond to a conditioned signal by salivating and it is difficult for him to develop skills with food reinforcement.

It is known that under normal conditions, appetite reflects the body's needs for energy and plastic (construction) materials. How does the food center become aware of these needs? What determines the formation of states of hunger and satiety?

The presence in the brain, and primarily in the hypothalamus, of receptors that perceive the level of nutrients in the internal environment of the body has been established. States of hunger and satiety are characterized by changes in blood composition (the content of glucose, amino acids, fatty acids), which are captured by central receptors. In accordance with this, the flow of nerve impulses to many brain centers that control food motivation changes. Signaling from stomach receptors plays a special role in its enhancement or suppression. The stomach, which is not filled with food, begins to contract at certain intervals. The period of physical activity is replaced by a period of rest. Against the background of stomach contractions, the hunger urge increases sharply, and the animals’ search for food becomes more active. After eating, the mechanoreceptors of the stretched walls of the stomach send signals that inhibit food motivation.

The food center and its receptors are influenced by many biologically active substances, including various hormones - insulin, glucagon, pituitary hormones, peptide hormones of the duodenum, sex hormones, etc.

Thus, food motivation is under the control of many humoral and nervous stimuli. Its strengthening or inhibition is determined not only by the body’s need for nutrients, but also by a number of external and internal conditions.

A large amount of material has been obtained on pharmacological effects on appetite. It has been established that the administration of opioids, norepinephrine, GABA, insulin, somatotropin (growth hormone), pancreatic polypeptide, male sex hormones (androgens) stimulates appetite and accordingly increases food consumption. Adrenaline, serotonin, cholecystokinin, bombesin, thyroliberin, calcitonin, corticoliberin, somatostatin, neurotensin, and female sex hormones (estrogens) have the opposite effect.

The mechanism of action of many of these substances on the formation of food motivation has not been sufficiently studied. Correcting appetite disorders should begin with identifying the cause of the disease. Often this reason can be completely eliminated by ordinary measures - improving the diet (restoring its balance, enriching it with vitamins), the correct regimen, and treatment of concomitant diseases. The selection of medications to correct appetite should be carried out by a doctor, and the use of pharmacological drugs to change normal appetite in order to influence the animal’s physique is extremely dangerous in terms of consequences.

Consideration of the peculiarities of the formation of food motivation would be incomplete without drawing the attention of dog handlers and dog breeders to the problem of specialized appetites.

Theories explaining the mechanisms of regulation of hunger and satiety are based on the idea of ​​the role of maintaining energy balance in the regulation of food motivation. Although it is largely correct, one cannot help but admit that very often a specialized reaction aimed at searching for and consuming certain nutrients dominates, and the vector of behavior depends on the body’s predominant need for them under given conditions. In accordance with this, it is customary to distinguish protein, carbohydrate, fat and other appetites. Particular attention is paid to salt appetite. Selective food preferences associated with the need for certain vitamins are also described. In some cases, the choice and consumption of any substances are not related to either the energy or plastic needs of the body. So, a dog suffering from worms begins to eat Chernobyl. This is an example of an instinctive defensive reaction that has become entrenched in the process of evolution of the species.

The mechanisms for regulating specialized appetites are extremely complex; congenital and acquired factors are closely intertwined. It is still unclear which preferences are inherited and which are developed during life. Obviously, the given example of dogs eating Chernobyl illustrates the first type of preferences, but with regard to other reactions of food choice, such clarity is lacking. Previously, it was believed that salt (sodium) appetite was innate. This position is currently being questioned. In any case, preference for salty foods largely depends on training. Apparently, even in cases where specialized appetite is determined by the type of nutrition inherent in a given biological species, it can undergo significant changes in accordance with the state of the body and environmental conditions.

Let's take the following example. If a dog is fed from a bowl in which it previously received bland food, it will begin to eat very salty food offered in this bowl, which is rejected when offered in another bowl. Undoubtedly, an innate reaction is to reduce the consumption of salty foods when chemoreceptors in the stomach and intestines are irritated. However, such a reaction can also manifest itself as a conditioned reflex in response to stretching of the walls of the stomach.

Specialized food choices are largely determined by established nutritional stereotypes. Imitation of parents and preference for food that animals received during the transition from milk feeding to independent feeding are of great importance. It should be noted that food preferences are not always completely adequate to the body's needs for nutrients - they may reflect other interests of the body (protection from poisoning, emotional reward, research, transportation of supplies, etc.). Food motivation may be based on drug or drug addiction, which is often used when training dogs to detect drugs.

The eating behavior of animals is characterized by a reaction that prevents the adverse effects of food on the body - neophobia. It manifests itself in the caution with which the animal treats unfamiliar food objects, even if they have an attractive smell and taste. At first, this food is consumed in small quantities, and if it does not cause a negative effect, neophobia is gradually inhibited.

The severity of neophobia is unevenly distributed among different animals in the population. In nature, as a rule, most of the population are “conservatives”, characterized by pronounced neophobia, and a smaller part are “scouts” with weakened neophobia. For domestic animals, human care allows them to largely avoid harm from insufficient conservation. Therefore, for example, quite a few young dogs can eat completely unexpected “products” upon first meeting them. Boxer Prince in his youth ate (always at the first contact with an object) a telephone directory, a pack of cigarettes, 400 rubles, a set of pastel paints, a hexachlorane pencil, a wonderful cameo and much more!

Of particular interest are reactions called conditioned reflex rejection (aversion). They manifest themselves in the refusal of food, the consumption of which caused a painful condition. Very often, aversions occur in response to completely benign food, the consumption of which coincidentally coincided with some disease. The resulting aversion to this food can persist for a very long time.

The following features are characteristic of conditioned reflex taste aversions. They are produced when the consumption of a certain food is combined with a painful condition (often a digestive disorder). Unlike conditioned reflexes produced to sound, light and other stimuli, taste aversions are formed even in cases where several hours pass between the action of the conditioned signal (new taste) and the unconditioned reinforcement (painful condition). It is important to note that in males the duration of aversion is significantly longer than in females.

Taste aversions begin to develop from the first hours of a puppy’s life. At the same time, at certain stages of its development, the nature of the development and extinction of aversion varies significantly, which is determined by many factors and, in particular, the degree of maturity of certain parts of the brain.

The following circumstance is interesting. It would seem that when developing and maintaining an aversion, only the relationship between the taste of a given food and the painful state that coincides with it matters. In reality, aversion also depends on the environment. It fades away much faster in rooms where the dog is kept constantly than in an unfamiliar environment. Apparently, in the first case, the food dominant has an inhibitory effect on the defensive one. Therefore, if a dog refuses a certain food, this reaction can be reduced by feeding it in the animal’s usual environment (and on a trip you need to take food that the dog likes).

The problem of taste aversions is important not only for the development of food rations that ensure optimal manifestation of higher nervous activity and, accordingly, the performance qualities of the dog. The possible formation of aversion to a certain type of food or anorexia - decreased appetite - must be remembered when introducing an animal to a new type of food and introducing various pharmacological drugs into the body. Many medications, along with the healing effect, lead to discomfort, a painful condition that is associated with eating. The subsequent refusal to eat can be mistaken for a metabolic disorder and appetite regulation. In reality, a typical conditioned reflex aversion is manifested here, after the extinction of which eating behavior is completely restored. This is probably the nature of many perversions of the appetite of lap dogs - during the period of introducing the puppy to different foods, he is intensively “stuffed” with various medications, without coordinating the feeding regimen, and then they are surprised that the grown dog does not eat meat, porridge, soup, etc. In order to avoid perversions of appetite, it is necessary to take into account that the aversion is weaker the more time has passed between the action of the conditioned signal (taste) and the negative reinforcement (drug intoxication). It is advisable to maximize the intervals between the last feeding and the administration of the medicine.

Using the example of the formation of food motivation, we examined some aspects of satisfying the biological needs of the body, but did not touch upon one of the main problems of the physiology of higher nervous activity - elucidating the mechanisms of reinforcement. Therefore, it is necessary to move on to a description of emotions, without which it is impossible to carry out purposeful behavior.

3.9. EMOTIONS

Biological needs that transform into motivation can be satisfied only when the latter is accompanied by the experience of pleasure or displeasure. Motivation and emotions are so closely related that it is customary to talk about motivational-emotional reactions. It would seem that they cannot be separated, but it has been experimentally established that they can be studied in isolation, since they are associated with different morphofunctional systems. This possibility has arisen only recently due to advances in neurophysiology.

In the past, the motivational-emotional sphere of an animal was judged by its behavior. In everyday life, people have long been accustomed to evaluate. the animal’s experiences through facial expressions, posture, and certain motor and vocal reactions. Involuntarily, such manifestations of behavior were compared with one’s own experiences. Charles Darwin left aside the analysis of the subjective aspect of emotions. Having shown that facial and gestural reactions are components of aggressive, defensive and other forms of behavior, he assessed them as adaptive mechanisms formed in the process of natural selection.

The discovery of emotional centers marked the beginning new era in studying inner world animals. It turned out that by irritating individual brain structures with electrodes immersed in them, it is possible to cause positive or negative reactions. If the tip of the electrode is in the “reward zone,” the animal develops an instrumental conditioned reflex: it begins to independently press the pedal, which turns on the irritating current. When the tip of the electrode enters the “punishment zone,” the animal bounces off the pedal when the current is turned on and does not approach it in the future. It turned out that the urge to self-irritate the “reward zone” is stronger than hunger, and animals can stimulate it for hours, forgetting about food. In the process of self-stimulation, they can choose the intensity of the current and the rhythm of pedal presses that provide the highest level of positive reinforcement.

The state of the “reward zones” associated with the formation of a positive emotional state depends on the balance of biologically active substances in the central nervous system (dopamine, norepinephrine, serotonin, opioids, etc.). The introduction of drugs into the body that change this balance can significantly affect the emotional sphere, which should be taken into account if you want to pharmacologically activate the dog’s behavior.

Above was information about the role of biologically active substances in preserving and retrieving a memorable trace. Apparently, the processes underlying memory largely depend on the emotions against which learning occurs, and therefore, many pharmacological drugs used to improve learning act on memory indirectly, changing the state of the emotiogenic systems of the brain. This is precisely what can explain the multidirectional influence of catecholamines and serotonin on learning during emotionally positive and emotionally negative reinforcement.

It is very important that, as experiments have shown, low and high levels of emotional stress have a negative impact on learning and retention of a skill. Its average level is most favorable, providing moderate brain activation. Against this background, emotions have a beneficial effect on the state of internal organs and sensory processes (vision, hearing, smell, etc.). When determining the desired level of stress on an animal, its age and individual characteristics must be taken into account. A certain hormonal background and the content of active substances in the internal environment of the body have a significant impact on the emotional state.

3.10. DISORDERS OF HIGHER NERVOUS ACTIVITY

The combination of neurosurgical interventions in certain parts of the brain and the conditioned reflex method has led to an understanding of many aspects of higher nervous activity. The analysis of its disorders expanded the understanding of the functioning of the brain in normal and pathological conditions. Although these interventions cause irreversible changes, the presence of compensatory processes in some cases mitigates the manifestations of behavioral pathology. Psychopharmacology allows you to make corrections in the behavior of an animal. Within the framework of this book, it makes sense to consider only temporary disorders of higher nervous activity (even if they are long-lasting) and the possibilities of eliminating them.

Under the influence of stress, an animal can develop long-term deviations from the norm both in behavior and in the vegetative sphere. They belong to the category of neuroses and are expressed in a violation of the orienting-exploratory reflex, in inadequate reactions to signals from the external environment, in memory defects, in poor learning, unstable manifestation of skills, insufficient spatial orientation, emotional shifts, as well as in pathological changes in the cardiovascular system. -vascular, digestive, endocrine and other systems of the body.

The concept of “experimental neurosis” was introduced by I.P. Pavlov based on an analysis of the behavior of dogs during floods. Subsequently, it was shown that a neurotic state in dogs can be caused by depriving them of sleep, by frequent changes in the stereotypes of conditioned reflexes, by the use of super-strong stimuli, and by a collision of opposite motivations (for example, food and defensive). One of the main manifestations of neurosis is a violation of the law of force, i.e. discrepancy between the magnitude of the response and the intensity of stimulation.

In experiments with the differentiation of complex complexes of conditioned reflexes, with the alteration of stable stereotypes, with the collision of food and defensive motivations, it was shown that overstrain of the mobility of nervous processes plays an important role in the development of a neurotic state. Using extremely strong stimuli, requiring an animal to solve a difficult problem, one can cause in it inadequate reactions of fear (phobias), circulatory disruption of conditioned reflex activity, phenomena of explosiveness of the excitation process, pathological inertia of nervous processes with obsessive movements. Many deviations from normal behavior are determined by the environment in which the dog encountered extreme stimuli.

The type of nervous activity characteristic of a given individual is essential for the development of neurosis. I.P. Pavlov identified the following types. Weak - with excessive inhibition and low performance limits of cortical cells, a tendency to passive defensive reactions. Unbalanced - with a predominance of excitation processes, aggressiveness. Alive (mobile) - with great mobility, balance and sufficient strength of nervous processes. Calm (inert) - with low mobility of nervous processes with sufficient strength and balance. This scheme is conditional, since among animals there are many representatives of mixed types.

Typological features of higher nervous activity are determined not only by heredity, but also by environmental conditions. By systematically working with a dog, you can improve its typological qualities. In addition, the nature of the puppy’s environment is of great importance. In an enriched environment and if it is possible to lead a relatively free lifestyle upon leaving the “nest,” signs of strong types more often appear. When puppies are kept in cages, the properties of the weak type may predominate in them, although genetically these puppies belonged to the strong type. Education can restore the original typological characteristics of the animal, spoiled by improper rearing at the beginning of life.

Pathological development of higher nervous activity was discovered in puppies that were raised from the 1st to the 7th-10th month of life in relative isolation. Their orienting-exploratory behavior was disrupted, a diffuse reaction to new objects was observed, increased motor activity was combined with a high level of fear, and they learned poorly.

In adult dogs whose development took place under normal conditions, susceptibility to neuroticism is determined mainly by hereditary factors. In animals of the weak type it occurs easily, in animals of the strong type it is difficult. In the latter case, disruption of higher nervous activity is possible only in extremely difficult and long-term situations. Against the background of a general illness or with a weakened functional state of the nervous system, neurosis develops more easily. An obsessive manifestation of fear has been described after the administration of drugs that have an inhibitory effect on the central nervous system.

In a puppy's life there are periods of increased sensitivity to difficult situations. Thus, at 3-4 months, puppies successfully solve the problem of bilateral choice, but due to overstrain of the nervous system, they often exhibit neurotic reactions. Therefore, it is necessary to provide developing animals with gentle conditions, and especially carefully protect them from stressors.

At any age, it is important to monitor the condition of the animal, give it rest and introduce difficult elements during training gradually. It is known that when exposed to super-strong stimuli, when a dog is treated roughly, or through ill-considered coercion, the dog develops a negative attitude towards the trainer and the environment in which the training takes place. A state close to neurotic (“pre-neurosis”) may develop, and in some cases, pronounced neurosis with a predominance of lethargy or agitation, inadequate response to commands, etc.

At the beginning of the study of experimental neuroses, the main attention was paid to processes in the cerebral cortex. IN last decade Attention was also drawn to the role of subcortical structures in the development of pathology of higher nervous activity. It turned out that many manifestations of neurosis and pre-neurosis are determined by changes in the functioning of these structures. Since the hypothalamus is closely connected with the regulation of not only motivational processes, but also autonomic reactions, disruption of its activity is often accompanied by shortness of breath, palpitations, the appearance of trophic ulcers, and pathological changes in the functioning of the digestive system. In turn, these shifts can affect the state of the central nervous system and behavior. A significant role in this is played by a decrease in food motivation, leading to the manifestation of a number of somatic (bodily) disorders and difficulties in learning associated with food reinforcement.

The reactivity of the central nervous system depends on the state of the endocrine organs and, above all, on the phases of the reproductive cycle. Thus, against the background of a developing estrus in a bitch, conditioned reflex activity is first activated and then suppressed. In males, with strong sexual arousal, feeding and defensive behavior is inhibited, hypnotic phases may appear, and the development of conditioned reflexes is sharply disrupted. Such dominance of sexual activity cannot be attributed to manifestations of neuroticism, but against this background it can arise more easily than under normal conditions. Fluctuations in the excitability of brain neurons, instability of positive and disinhibition of inhibitory conditioned reflexes occur during pregnancy. In its second and partially in the third phases, stress resistance is increased, and previously manifested neurotic symptoms become less pronounced at this time.

There is a lot of data on changes in higher nervous activity with hypo- or hyperfunction of the reproductive, thyroid, parathyroid glands, pituitary gland and adrenal glands. Injections of hormones from these glands affect the development and maintenance of conditioned reflexes. Thus, long-term administration of the adrenocorticotropic hormone of the pituitary gland or the hormone of the adrenal cortex - cortisone - causes a neurotic state with a predominance of chaotic behavior. Such deviations, caused by a violation of internal inhibition, can persist for several months after the cessation of hormonal influence.

The nature of the influence of various hormones and other biologically active substances on behavior is determined by many factors, in particular, their dosage and the typological characteristics of the animal. Small doses can stimulate the development and implementation of conditioned reflexes, large doses can cause a breakdown of nervous activity with manifestations of motor excitation.

Treatment of neuroses and relief of pre-neurotic conditions are associated with serious difficulties. At the first unfavorable symptoms, it is necessary to reduce the load and stop training. Significant progress in the treatment of neuroses has been achieved thanks to the advances in psychopharmacology. In case of disruptions of higher nervous activity, a combination of bromine and caffeine drugs is traditionally used. Bromides enhance the inhibitory process and thereby normalize the balance of excitation and inhibition in the brain. There is evidence that in this case some structural changes occur in the nervous system, so it is extremely important to monitor the correct choice of dosages of this drug. Doses of bromide used in the treatment of neuroses in dogs with a weak type of nervous system should be minimal. Low doses should also be used in highly neurotic animals with a relative weakness of the inhibitory process and a predominance of excitation. In animals of the strong type with balanced nervous processes even large doses of bromides may not have an effect on conditioned reflex activity. Such doses are usually used for significant arousal, weak doses - for hypnotic phase states. Action various salts bromides are the same, but the ammonium salt is absorbed faster and is therefore more effective than potassium and sodium bromides. In case of overdose, irradiation of the inhibitory process and deterioration in learning are observed. Long-term administration of these drugs in large doses causes poisoning (bromism), which is more pronounced in weak dogs.

Caffeine, included in a complex of therapeutic measures for neuroses, increases excitation and thereby has a beneficial effect on inhibition processes. In dogs of the weak type and in excitable animals of the strong type, an overdose of caffeine can be accompanied by depletion of cortical cells, the development of extreme inhibition, and the manifestation of hypnotic phases. The use of caffeine, either alone or in combination with bromides, should be based on careful dosage selection.

Neurochemical changes characteristic of neurotic conditions were studied. It turned out that as a result of the collision, the content of acetylcholine in the blood decreases, and when conditioned reflex activity is restored, it returns to normal. The content of catecholamines increases with the development of neurosis, and in dogs with pronounced motor excitation the level of norepinephrine increases, and in dogs with signs of inhibition - adrenaline. In accordance with this, when treating neuroses, drugs should be selected whose administration restores the balance of the adrenergic and cholinergic systems of the brain.

Neuroleptics have a calming and normalizing effect on behavior. Good results in the treatment of neuroses were obtained using chlorpromazine, which reduces the overall level of activity of the central nervous system. It eliminates the phenomena of stagnant inhibition, fear of open space, cyclical behavior, reduces the level of passive-defensive reactions, stimulates appetite (if food is refused), and relieves autonomic disorders. If neurotic symptoms are less pronounced, you can use drugs of the same series that have a milder effect - propazine, triftazine, trioxazine and others.

The group of minor tranquilizers includes meprobamate, chlordiaze oxide, diazepam, phenazepam. They have a beneficial effect on the implementation of already developed conditioned reflexes in dogs placed in conflict situation(for example, a collision of food and defensive motivations). All tranquilizers reduce emotional stress such as fear and anxiety. Depressed animals become more active, more proactive, and their relationships with other dogs are restored. Elimination of fear can lead to an increase in aggressive reactions in dogs under the influence of sedative medications, which is important for watchdog and protective guard services. In this case, claims to a leading position in the group may arise. Overdoses of tranquilizers can make it difficult to develop new skills (memorization worsens), as well as cause a number of undesirable shifts in the formation and implementation of behavioral acts. The muscle relaxant effect of tranquilizers can reduce the strength and accuracy of a dog's movements. Daytime tranquilizers, such as gidazepam, are free from this deficiency.

In the treatment of neuroses, amizil and metamisil (anticholinergics) are used, which reduce the level of fear, relieve depression and improve the production of conditioned reflexes. An overdose of these drugs leads to serious intoxication.

The effectiveness of tranquilizers is most fully manifested in the stage of pre-neurosis. They correct emotional behavior, improve the development of skills, and eliminate autonomic disorders. However, in the stage of full-blown neurosis, their effect can be short-term and be replaced by deterioration in indicators of higher nervous activity. In laboratory conditions, a combination of tranquilizers with psychostimulants and antidepressants was used, which in some cases gave good results. We caution, however, dog owners against self-appointment these drugs. Their choice and dosage can only be determined by a specialist, who will be guided in his decisions by many criteria.

3.11. CONCLUSION

The solution to problems of higher nervous activity is successfully developing and, presumably, we will continue to witness new discoveries in this rather complex area. Undoubtedly, every new achievement in understanding the motivational-emotional sphere and behavior of animals, including, of course, dogs, is of paramount applied importance. Thanks to advances in physiology, biochemistry and pharmacology, it is possible to make significant corrections in the development of animals used by humans for a variety of service purposes. The search for adequate effects on the animal’s body at different age periods, put on a scientific basis, makes it possible to improve its phenotype. An example here is the use of an enriched environment in which the puppy is placed in a environment that promotes brain maturation and therefore optimizes behavior later in life. Advances in the science of nutrition and digestion are also noteworthy, thanks to which optimal diets have been developed.

Particular attention should be paid to the problem of pharmacological effects on the animal’s body in order to improve its appearance and performance qualities. In laboratory conditions, it has been shown that phenomenal results can be achieved with the help of hormones and psychopharmacological drugs. Here, however, many surprises await us. Stimulating a healthy body with psychostimulants (caffeine, sydnocarb, etc.) leads to short-term success, but begins an imbalance that is often difficult to correct over many months and even years.

Another thing is the correction of disorders of higher nervous activity. Thanks to the development of psychopharmacology, it has become possible to normalize disrupted behavior. With the help of aminazine, it is possible to tame the most ferocious animal; minor tranquilizers have become firmly established in the arsenal of means for combating neurotic reactions. But we must not forget about the purely individual sensitivity of dogs to the effects of these drugs. Their unwise use can do more harm than good. Diseases caused by an overdose of substances that affect the state of the central nervous system often turn out to be much more severe than the illnesses that the treatment was aimed at eliminating. Therefore, in all cases related to the need to correct higher nervous activity, it is necessary to consult a specialist with a deep understanding of the internal mechanisms that ensure behavior.

3.12. Editor's Notes on the Use of Psychotropic Drugs in Dog Breeding

The chapter on behavior management was written by one of the best specialists in the field of physiology of higher nervous activity. As the reader can see, the author’s position regarding the use of pharmacological agents to artificially regulate the behavior of dogs is not only strictly scientific, but also humane.

The author is cautious about the use of drugs to correct the behavior of animals, both due to the insufficiency of modern knowledge for a targeted influence on the individual psyche, and for reasons of a humane nature (fear of the animal becoming disabled). Professor V.G. Cassil believes that a deep knowledge of the physiology of behavior in most cases is quite enough to do without the still imperfect pharmacological agents. Indeed, as is clearly shown in the chapter, skillfully using physiological techniques and knowledge, you can very effectively, no worse than many drugs, influence the behavior of an animal. I don’t know, for example, of a medicine that would better increase a dog’s sensitivity, attention, orienting reaction than a slight feeling of hunger, or act in a calming manner better than satiety.

By by and large a physiologist always looks at the problem of regulating body functions more deeply and thoroughly than a practitioner and a doctor. The vulnerability of the position of a negative attitude towards the use of psychotropic drugs in dog breeding, apparently, is only in the fact that, despite almost the same arguments, medicine still uses them (psychotropic drugs today are among the most frequently used - the sales volume of psychotropic drugs in developed countries by 1990 approached $10 billion). In addition, the reality is that some dog breeders already use psychotropic drugs, and this creates inequality of opportunity and the need for knowledge of the pharmacology of psychotropic drugs to organize control over their use. It is also unlikely that the use of psychotropic treatment of dogs during their exploitation should be condemned. A working kennel, whose dogs must serve intensively, cannot refuse the opportunity to speed up training, relieve stress, increase the sensitivity of dogs, even if some of them break down faster. The only thing to be condemned is the illiterate use of the capabilities of modern psychopharmacology.

It should also be recognized that selection achievements in dog breeding have led a number of breeds (or lines within breeds) to actually consolidate psychopathology in the genotype of a significant group of animals. Indeed, pit bull terriers, to which the press has firmly assigned the epithet of “killer dogs,” differ from most breeds not so much in their exterior features as in their psyche (unbridled anger, insensitivity to pain, stubbornness, etc.). This may be considered a hereditary mental illness rather than a healthy natural behavior option. For a person who, for one reason or another, has become the owner of a pit bull, the use of psychotropic drugs may be necessary and the only humane method of controlling the behavior of his dog. The use of antidepressants can be a way out of a difficult situation for many owners of decorative indoor dogs with a vulnerable psyche. Apparently, it is no coincidence that the success of P. Neville’s book “Do Dogs Need Shrinks?”, 1992 (“Do Dogs Need a Psychiatrist?”), which has become a bestseller in recent years. This book and chapter written by V.G. Kassilem, different in many ways. In my opinion, the chapter is deeper and more accurate than Peter Neuville's book. Of course, this reflects not only the difference in the approaches of the authors, but also the difference in the problems facing domestic and foreign dog breeder readers. But the works of P. Neuville and V.G. Kassil is united by the humane approach and the priority of physiological methods of solving problems over pharmacological ones, which, however, are not excluded.

The temptation to use psychotropic drugs on a healthy animal as doping is quite great, and the standards of exhibitions and competitions are so vague and the chances of catching a violator are so small that one can only be surprised that the use of such methods has not yet become universal in our not very civilized conditions. I know for sure that dog breeders use painkillers, sedatives and stimulants as doping agents during dog shows. I have met completely illiterate trainers who specialize in dogs that are difficult to train (higher fees) and use a course of nootropics. A not very competent dog breeder sometimes takes a pharmacological reference book and selects from it psychotropic drugs with the desired effect, in the opinion of the dog breeder. In this case, preference is usually given to the most powerful drugs and completely forgotten about such rather mild and effective drugs as vitamins, adaptogens and other drugs, the use of which in most cases would be more justified due to the normalization of the functional state of the dog’s body and its nervous system. .

Brain cells are very sensitive to oxygen starvation, and therefore, in many cases, instead of saturating the internal environment of the body with foreign chemical compounds, it is better to provide the animal with exercise and good hematopoiesis. Only when physiological means of improving brain activity have been exhausted can one, for example, try antihypoxic drugs.

Conscious refusal to use psychotropic drugs on healthy animals is highly desirable. It requires a lot of educational work. Concealing information only leads to the fact that practitioners use the most crude techniques, the dramatic effect of which creates advertising for them and at the same time makes them the most dangerous in the wrong hands. The conviction of the need to refrain from the doping use of psychotropic drugs in dog breeding is all the more important since it is very difficult to establish appropriate effective control.

Control of the use of psychotropic drugs, in principle, can be based on recognizing the physiological signs of the action of these substances and on detecting traces and products of doping in the body. Biochemical test results are usually more reliable when the answer is positive than when it is negative. The latter can be obtained not only in the case of the doping purity of the animal, but also due to the limited capabilities of the detection technique (they were looking for the wrong thing, using the wrong method, not then, etc.). It is especially difficult to detect doping if it is indistinguishable from the natural components of the body’s biochemical composition. This applies to natural hormones and other regulators of nervous system functions, their precursors and rapidly metabolized “provocateurs” of a cascade of events leading to a doping effect by the time the original stimulus has already disappeared from the internal environment of the body.

Inadequately dilated pupils, surges in blood pressure, hyperemia of the mucous membranes, and other autonomic components of the effect of psychotropic drugs on a dog can draw the expert’s attention to the possibility of doping. In the absence of chemical control over the use of doping in dog breeding, in fundamental issues of breeding work, it is necessary to pay more attention to long-term observations of the animal, and repeated, unscheduled examinations of it.

Apparently, the problem of psychotropic doping is most acute in an industry that is unpleasant to most dog breeders, but actually exists - fighting and racing. Among other things, these types of competitions are usually associated with monetary stakes and the hard-heartedness of the owners (who love their dogs for victories and are offended by them for defeats). Here, equality of chances is possible either with permissiveness (everyone is equal, since everyone is allowed to do everything), or with the most stringent doping control. It may be necessary to introduce long-term holding of dogs before competitions in isolated kennels (short-term drugs will be eliminated), mandatory tests of urine, blood, etc. Due to the difficulty of detecting doping, it is probably necessary to legitimize the most severe punishments and ostracism of violators for using those methods that are prohibited. Severe punishment can enhance the deterrent effect of the possibility of being caught doping.

The streamlined phrase in the current regulations on dog shows, which states that “a dog breeder should not use methods to hide the animal’s deficiencies,” should be specified taking into account the characteristics of psychotropic doping stimulation and modern control methods. Otherwise, this puts, for example, special training, handling and specialized feeding of dogs with muscular deficiencies on the same level with psychotropic treatment of animals with defects in higher nervous activity.

Notes:

When we talk about the coincidence in time of conditioned and unconditional signals, we mean the somewhat advanced action of the first (the advance should be at least 0.6 s). If the signal and reinforcement completely coincide, or if the command is delayed, the conditioned reflex is not developed. This is a common reason for failures among novice trainers - the signal and unconditioned reflex reinforcement are given at the same time and the skill is not developed. The experiment demonstrated the possibility of forming a conditioned reflex when the order of action of the stimuli is reversed to the usual, i.e. the indifferent stimulus precedes the conditioned one and “covers” it. In this case, however, the resulting conditioned reflex is unstable, quickly fades away and can turn into an inhibitory one.

During training, first-order conditioned reflexes, developed and strengthened using positive (food, stroking, etc.) and negative (punishment) reinforcement of distant conditioned stimuli, serve as the basis for the further formation of various skills. For example, a whistle signals encouragement, and a clap signals punishment. With the help of these signals, the trainer controls the behavior of the animal in free movement over a considerable distance. Traditional commands “Okay!” and “Ugh!” are such signals. Naturally, the signal rewards and punishments themselves require periodic reinforcement according to the laws of conservation of conditioned reflexes.

The further evolution of behavior is associated with the emergence of complex differentiated reception apparatuses, which make it possible to perceive highly specialized information coming from the external environment. It is also associated with the development of complex programs that allow the animal to adapt to complex, albeit constant, stable environmental conditions. All this becomes possible at further stages of the evolution of the ganglion nervous system and is especially clearly manifested in arthropods.

Complicated living conditions make it necessary to form a variety of sensitivity devices that make it possible to register various influences of the external environment. Let's look at this using evolution as an example. photoreceptors. At first, light-sensitive cells were simply concentrated on the front surface of the body. This gave the animal the opportunity to perceive the effects of light, but did not yet allow it to localize the light source in space. At the next stage of evolution, light-sensitive cells were concentrated in two light-sensitive plates located on both sides of the anterior end of the body. This made it possible to navigate in the spatial position of the light source and turn the body to the right or left, but did not yet make it possible to distinguish the properties of the object acting on the body. Only at the last stage of evolution did the supersensitive plates bend, taking the shape of a hollow ball. A small hole, which was then filled with a refractive medium (lens), allowed the incident ray to be refracted, and the effect of the light object was imprinted on the sensitive layer of this hollow ball. The apparatus of a complex light-sensitive receptor - the eye - emerged, which for the first time made it possible not only to react to the presence of light, but also to reflect the properties of the influencing object.

The structure of the eye, the most important light-receiving organ, varies from animal to animal. In insects it has the character of a “compound eye”, sometimes built from many thousands of independent cells. In vertebrates, it takes the form of a single eye, well known to us, which allows us to perceive the reflection of an object and change the clarity of the reflection using a self-regulating system of the refractive apparatus and muscles. However, in all cases, the emergence of a complex apparatus that allows one to navigate at a distance in influencing objects remains one of the most significant achievements of evolution.

Insects have a large number of highly differentiated receptors. Along with a complex photoreceptor (eye), they have:

Special tactile-chemical receptors (located in the antennae);


Taste buds (located in the mouth, on the legs), which detect subtle changes in taste;

Vibration receptors (located in the membranes of the legs), responding to the finest ultrasonic vibrations, sometimes up to 600 thousand vibrations per second.

It is possible that there are a number of other types of receptor apparatus unknown to us, the specialization of which was developed over millions of generations.

Excitations caused by influences falling on these receptor apparatuses spread along the nerve fibers and come to anterior ganglion, which is a prototype of the brain and an apparatus that combines (encodes) impulses reaching it and translates these impulses into highly complex systems innate behavioral programs that underlie the adaptive movements of the insect.

The anterior ganglion of higher insects, such as bees, has a very complex structure. It consists of a cluster of differentiated nerve cells that receive impulses from peripheral receptors. The anterior part of this ganglion contains predominantly visual cells, the middle part contains olfactory cells, and the posterior part contains sensitive cells of the oral cavity. It is characteristic that the arrangement of these cells is organized. In them one can already observe a planar “screen” structure, which allows the evoked excitations to spread through the neural structures of the anterior ganglion in an organized manner, thereby ensuring the reflection of known structurally organized influences.

It is characteristic, as has been established by recent studies, that already at this stage of evolution the anterior ganglion includes highly specialized neurons that respond to individual tiny signs of information reaching the body, decomposing it into a large number of constituent elements and allowing them to be subsequently combined into entire structures (the forms of operation of these neurons will be discussed below).

All this makes the anterior ganglion of higher insects a very complex central apparatus, allowing it to capture diverse environmental influences and encode them into entire systems.

Excitation codes that arise during certain stimulation in the anterior ganglion of insects are transmitted in the form of complex behavioral programs to the underlying thoracic ganglion, where impulses of complex adaptive movements of the insect that make up its behavior arise.

The most complex behavioral programs of insects are not only of great interest, but also require special detailed consideration.

The peculiarity of the most complex programs, which make up the vast majority of insect behavior, is that they are congenital And are passed on by inheritance take the well-known form instinctive behavior. These programs are developed over many millions of generations and are passed on hereditarily, just as the structural features of the body (the shape of the wings, the features of the proboscis, the structure of the receptor organs) are well adapted to the living conditions of insects.

Examples of innate behavior programs in insects are very numerous. Often they are so complex and expedient that some authors considered them an example of reasonable behavior.

It is known that the larva of the birch elephant cuts a birch leaf into an ideal geometric shape, which is close to the optimal, mathematically calculated structure, in order to then roll it into a tube and use it for pupation. The mosquito lays its eggs on the surface of the water and never lays them on land, where they will inevitably dry out. The sphex wasp lays eggs in the body of the caterpillar so that the emerging larvae do not lack food. To do this, she first pierces the thoracic ganglion of the caterpillar so that the caterpillar does not die, but is only immobilized, and does this with amazing accuracy. Should we talk about the innate behavioral programs of a spider, which weaves a web that is amazing in its design, or about the innate behavioral programs of a bee, which builds honeycombs of an ideal, from an economical point of view, shape, fills these honeycombs with honey and seals them with wax as soon as they are sufficiently filled .

The above examples of the most complex, purposeful behavior and many others are innate; the insect does not have to learn them; it is born with these forms of behavior, just as it is born with an ideal wing shape or with a structure of sense organs that is amazing in its purposefulness.

Only for Lately research by zoologists and, in particular, that branch of science called ethology(ethos - behavior), brought some clarity to the mysterious form of behavior and showed that behind this form of activity, striking in its complexity and apparent rationality, elementary mechanisms are hidden. These studies showed that the most complex programs of “instinctive” behavior are actually caused by elementary stimuli that set into motion innate cycles of adaptive acts.

Thus, the laying of mosquito eggs on the water surface is caused by shine water; therefore, it is enough to replace the water with a shiny mirror so that the mosquito begins to lay eggs on its surface. The complex innate activity of a spider that lunges at a fly caught in its web is actually caused by the vibration of the web, and if the web is touched by a vibrating tuning fork, the spider lunges at it in the same way as it lunges at the fly.

The described mechanisms allow us to take a significant step in improving our understanding of the processes underlying innate behavior and moving from simple description to his explanation, to show how instinctive behavior different from reasonable.

Let us give just one example showing how difficult such research proceeds and what interesting results it leads to.

It is known that some species of earthworms that store leaves for the winter pull them into their burrows by the end. This was considered a manifestation of the “rational activity” of worms, which was once spoken of by C. Darwin, and led to the assumption that the worm perceives the shape of the leaf and “calculates” which end is best to pull it into the hole.

This assumption changed significantly after a German researcher Ganeli performed the following experiment. He cut out a piece from a leaf, reproducing the shape of this leaf, but with the tip facing downwards. In this case, the worm attempted to pull the leaf into the hole not with its sharp end, but with its blunt end. The question of why he does this became the subject of research by another scientist - Chard. This researcher suggested that the worm's behavior was not dictated by shape perception, but by a much more basic chemical sense. To test this, he placed a row of identical sticks in front of the worm, but lubricated one end of these sticks with an extract from the tops of the leaf, and the other with an extract from the base of the leaf, or one end with an extract from the top of the leaf, and the other end with an extract from the cutting. As a control, experiments were carried out in which one end of the stick was lubricated with an extract from the top of a leaf or cutting, and the other end with neutral gelatin. The results of the experiment showed that in these cases the frequency with which the worm pulled the stick into the hole by one end or the other was not the same, and that the main factor deciding the matter was the difference in the chemical difference between the top of the leaf and its stem (Table 1.2).

Table 1.2

Results of Mangold's experiments

Hereditarily fixed reflexes underlie adaptive behavioral acts that manifest themselves without prior training. These reflexes are similar in all representatives of this species. At the same time, some innate reactions in representatives of different breeds of dogs can vary significantly (for example, in hunting and herding dogs).

The animal's body is well adapted to environmental conditions throughout its life, including during the prenatal period of development. It has been shown that in mammalian fetuses, in accordance with the unfolding of the genetic program, the central nervous system gradually matures and first generalized and then specialized reflex reactions to various stimuli arise. By the time of birth, complexes of central and peripheral nervous formations and associated apparatuses are formed, providing, during the neonatal period, first less complex and then increasingly complex specialized behavioral acts. They are aimed at interacting with the mother and surviving in the rather greenhouse conditions of the den.

I.P. Pavlov considered innate behavior as a set of complex unconditioned reflexes (instincts). He was primarily interested in the relationship between inherited and acquired reactions during life, studied under laboratory experimental conditions. Ethologists who study primarily the deployment of the genetic program in the natural habitat describe behavior from slightly different positions. Currently, the contradictions between these approaches are being smoothed out and attempts are being made to create a synthetic theory of behavior.

Can the manifestation of unconditioned reflex activity be considered unchanged in all individuals? Obviously this question should be answered in the negative. In accordance with the genetic program underlying development, all manifestations of an individual’s life activity are individual in nature. This indisputable position also applies to innate reactions to various stimuli. Thus, each animal has different thresholds for the perception of signals associated with the functioning of the sensory organs. Depending on the individual characteristics of the central and peripheral nervous systems, on the level of production of biologically active substances, innate reactions to certain stimuli manifest themselves differently. At the same time, the limits of variation in these reactions in representatives of a certain breed of dog are much narrower than in individuals of different breeds.

Environmental factors largely determine the characteristics of innate behavior. It can undergo significant changes as a result of influences on the animal’s body at different stages of life, but most of all during sensitive (critical) periods, which sometimes take only a few days (and sometimes hours). At this time, certain external stimuli can modify the nature of the animal's response at subsequent stages of life. For example, unfavorable living conditions for females at the end of pregnancy (diet insufficient in calories or unbalanced in composition, stressors) lead to disruption of the sexual behavior of male offspring, who, upon becoming adults, behave in many respects according to the “female type.” It has been shown that such loss in male puppies can be prevented by administering tyrosine (an amino acid used in the biosynthesis of catecholamines) or a beta-endorphin blocker - naltrexone. Good results have also been obtained with injections of the male sex hormone testosterone. We must not forget, however, that when correcting one important function may have an adverse impact on the development of others.

Nature has provided protective mechanisms that mitigate the effect of damaging factors on the developing organism. Thus, when there is a deficiency of essential amino acids in food, they are mainly supplied to the fruits. However, offspring are not protected from the consequences of stressors even with short-term isolation from the mother during the neonatal period. Such exposure leads to a slowdown in body weight growth and irreversible changes in the structure of the brain, which will manifest themselves in behavioral defects. This is why it is important not to rush into weaning puppies from their mother.

The diverse reactions of a mature organism to external and internal signals can change under the influence of excessively strong early influences on the body. They depend on many factors that are important for higher nervous activity. These factors include: the number of puppies in the litter, the mother’s attention, and the external environment enriched or depleted in various signals. Even before birth, the fetal body is exposed to chemical influences as a result of environmental and food pollution or if the puppy is treated with any pharmacological drugs. Being relatively harmless for an adult, they are capable of distorting the development of the genetic program and changing the rate of maturation of the brain and endocrine glands. Hypoxia, a condition that occurs when there is insufficient oxygen supply to the body, has serious consequences. It often occurs during abnormal births, as a result of which brain development is disrupted, and animals exhibit deviations from normal behavior throughout their lives. Improvements in metabolism nerve cells and normalization of brain function in these cases can be achieved by administering nootropic drugs to puppies, in particular piracetam (nootropil) and dimethylamine ethanol. Injections of ACTH-like peptides (ACTH 1-10, ACTH 4-10) also have a beneficial effect.

The above examples show that unconditioned reflexes are characterized by a certain variability associated with the details of the “biography” of the individual. They undergo changes in accordance with the state of the body and, above all, the control nerve centers. However, unconditioned reflexes as an innate form of behavior are relatively constant in comparison with acquired - conditioned reflexes, i.e. stereotypically manifest in response to irritation of certain nervous devices - receptors.

Many unconditioned reflexes have been described that are associated with various aspects of behavior and the regulation of vital systems of the body in accordance with the biological role of these reflexes, the type of stimuli that cause them, levels of control (connection with certain parts of the brain), and the order in which they occur in a specific adaptive act; several classifications have been proposed. I.P. Pavlov identified food, defensive, orientation, parental, and child reflexes, each of which can be divided into more specific ones. For example, food reflexes include reactions associated with search, extraction, inspection, grasping, tasting, absorption of food, secretion of digestive juices, movements of the stomach and intestines, etc.

When analyzing innate forms of behavior, the following reflexes are described: goals, collecting, caution, freedom, self-preservation (positive and negative), aggressive, watchdog, submission, sexual (male and female), play, parental, group (animal social), migration, saving strength , sleep regulation, restorative, imitation.

Unconditioned reflexes can be considered according to their level of complexity. The simplest include reactions of local significance, for example, blinking when a speck gets into the eye or withdrawing a burned paw. Coordination reflexes are more complex, as an example - a reflex that coordinates the contraction of flexor and extensor muscles. Integrative unconditioned reflexes include complexes of movements and accompanying changes in the body.

The mechanisms of nervous regulation at different levels are closely intertwined. The complexity of the organization of innate forms of behavior is clearly illustrated by the example of the salivary unconditioned reflex, which in the past was considered to be quite simple. In fact, it depends on the activity of many receptors, fibers of several cranial nerves, and many parts of the central nervous system. Salivation is associated with eating behavior, digestive processes, the work of the endocrine glands, blood circulation, breathing, and thermoregulation.

The relativity of any classification can be clearly seen in the example of one of the most important unconditioned reflexes - the indicative one. Due to its special role in behavior and connection with conditioned reflex activity, further attention will be paid to it.