2.3. Internal and external factors. Structure of instinctive behavior

Internal factors of instinctive behavior. For a long time, it was believed that learning was determined by external factors, and instinctive behavior was determined solely by internal factors, and the nature of these factors was unknown. Search, clarification of internal factors of instinctive behavior would allow to answer the question, what determines the motivation of behavior.

Internal factors undoubtedly influence the instinctive behavior of animals. In the middle of the XX century. American biologist P. Whit conducted experiments with spiders, during which he studied the weaving of cobwebs when various chemical substances were ingested into an animal. The desired substance was applied as a drop directly on the web or injected with a syringe into the victim. Each substance stimulated a spider to weave a web of a certain type, while the web weaving reaction itself is hereditary in a spider. So, caffeine made spiders weave a shapeless web of randomly matted threads, while the spider experienced a semblance of neurosis. When pervitin was introduced into the organism, the spider became very restless and did not spin the web in its entirety. Hydrochloride caused torpor in the spider, and he did not doze the web. And lysergic acid promoted increase of concentration of attention on weaving, and the spider wove a web very carefully and exactly, at the same time its quality exceeded natural.

The internal environment of the body is constant, various regulatory processes are aimed at maintaining the physico-chemical composition of the medium. It is constantly updated, but all its parameters are maintained at a certain level due to self-regulation, which ensures the flow of all biochemical reactions. The peculiarity of the internal processes of the animal organism is that they often occur in the form of rhythms. In the 1930s Soviet zoopsychologist V.M. Borovsky suggested that it is the deviations of these internal rhythms of the body from the norm that are the primary motivation of behavioral reactions. In certain conditions, the internal consistency of physiological rhythms is broken, and the previous balance in the new conditions does not ensure the normal functioning of the body. An inner impulse arises, aimed at restoring the internal equilibrium, that is, a need arises. Instinctive behavior in this case will be aimed at meeting this need.

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

For intrinsic motivation, rhythmic processes that occur in the central nervous system are of particular importance. The rhythmic activity of its stem in vertebrates and abdominal nerve structures in invertebrates provides orientation of behavior over time. It is known that animals have so-called “biological clocks” - autonomous oscillatory processes that regulate all the rhythms of the vital activity of the organism. "Biological clock" determines the fluctuations in the external activity of animal behavior, all actions that repeat with a certain cyclical nature. They, as it were, lay the foundation for the animal's instinctive behavior, and the environmental factors introduce their corrections into these rhythms. Changes may be associated with the action of a variety of external stimuli (auditory, visual, etc.), and may also depend on the general physiological state of the animal at the moment. Most often in the behavior of animals circadian or daily rhythms are noted, the period of which is equal to days.

It is interesting to note that the activity of the animal is subject to such rhythmic daily fluctuations even in conditions of complete isolation from all environmental factors. For example, an animal can be placed in the conditions of full round-the-clock illumination and nevertheless observe in it an alternation of periods of sleep and wakefulness, which is close to natural. In addition, short-term rhythms may be noted in the behavior of animals during the day. An example would be the observations of the German ethologist V. Schleidt for turkeys. He noted that the turkey’s clogging during the day is repeated with a certain rhythm, which persists even with complete isolation of the bird and deprivation of its hearing.

In addition to the orientation of the animal's behavior in time, the “biological clock” orients it in space. For example, migratory birds with orientation on the position of the sun should at every moment of time relate its position to the time of day. This happens when they relate information about the position of the sun to internal circadian rhythms.

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

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

German neurophysiologist E. Holst discovered several zones in the hen's brainstem. When exposed to these zones with a weak electric current, instinctive movements arise, corresponding to a particular zone. It was noted that if, for a long time, one and the same zone was affected, increasing the force of irritation, one could observe a whole series of instinctive actions that would be performed in the same manner as in natural conditions. For example, a hen performed movements that were made when a ground predator approached: it showed a slight anxiety, then rose, flapped its wings, screamed, and then took off. At the same time, the stimulus itself (predator) was not in sight. Thus, under the influence of exclusively internal factors, not only individual instinctive movements, but also whole instinctive actions can manifest themselves. However, it should not be forgotten that in natural conditions instinctive actions are “triggered” by external factors. Approaching a land predator that a chicken would see would lead to the initiation of the corresponding zone of the bird's brain, which was artificially stimulated in the experimental conditions.

External factors of instinctive behavior. If the task of the internal factors of instinctive behavior consists mainly in preparing the body to perform a certain behavioral act, then external factors more often play the role of peculiar activators of this instinctive action.

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

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

The key stimuli can be simple physical or chemical signs, their spatial relationships (for example, size correlation) or vectors.

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

Symbolic stimuli are extremely diverse and by their nature: they can be visual, acoustic, chemical, etc. For example, in the sexual behavior of many insects, amphibians and some mammals, chemical stimuli (sexual attractants, pheromones) serve as key stimuli. Sound stimuli include a variety of screams, songs specific to a particular type of animal. Visual key stimuli are called “releasers”. These include various morphological features (features of body color, ridges, crest in birds, growths). For example, for female mallards, the releasers are “mirrors” on flight feathers of drakes. There are also special species-specific complexes of movements that can act as symbolic stimuli (postures of submission, threatening poses, greeting rituals, marriage rituals).

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

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

The existence of key stimuli and their role in the development of instinctive reactions have been proven by a multitude of observations and experiments. N. Tinbergen studied the food reaction of chickens of silvery gulls and thrushes with the appearance of the parent individual by the method of mock-ups.

The natural reaction of a hungry gull chick to a parent is that it bites a red spot on the beak of a (yellow) adult bird. Tinbergen in his experiments used several layouts. Only one layout exactly repeated the appearance of the head of an adult herring gull. On the other layouts, individual parts were excluded, and gradually the layout became less and less like the head of a seagull. The last layout was a flat red object with an oblong protrusion. However, the reaction of chicks to this subject not only was not weaker than the reaction to the first layout, but even surpassed it. The chick’s reaction to the mock-up in the form of a thin white stick with transverse dark red stripes became even more intense. From this we can conclude that the key stimuli for the appearance of the “begging” reaction in chicks of the silvery gull are the red color and oblong shape.

In experiments with ten-day thrush chicks, flat discs were used as mock-ups. If the thrush chicks were offered a circle, they reached for its upper part, where the head of the parent bird was assumed. If a small circle was attached to a small circle, the chicks began to reach for it, and when two small circles of different sizes were attached, the relative size of the circles became decisive. With a large size of the “trunk,” the chicks reached for an extra large circle, and for a small one, to a smaller one. Thus, the key stimuli in this case are the relative position and relative magnitude of the details of the layout.

Experiments on the study of key incentives in birds were carried out by Russian ornithologists G.L. Skrebitsky and T.I. Bibikova. In the course of the experiments, the relation of the gull to its eggs was investigated. Researchers transferred eggs from one nest to another, replaced them with eggs of other species of birds, other objects of different shapes, sizes, colors. Gulls readily began to “hatch” alien eggs, as well as eggs of other birds, variously colored dummies made of different materials (glass, clay, etc.), foreign objects (balls, potatoes, stones). Birds did not refuse to roll even heavy stone balls into the nest, i.e., it was not the egg weight that determined this reaction. G.L. Skrebitsky wrote: “... the seagulls sitting on such objects represented a very original picture, but the spectacle became especially unusual when the bird that had been driven from the nest came back to it and, before sitting down, carefully adjusted the colored balls, pebbles or potatoes with its beak. ". ^[eight]

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

If the seagull was offered two eggs of different size, it began to roll into a larger nest. The researchers even observed such a situation when a seagull tried to incubate a wooden mock-up of an egg of such gigantic size that it hardly climbed onto it. In this situation, there is a super-optimal reaction. The animal is confronted with overstimulus, which possesses superoptimal signs of a key stimulus, and begins to show a reaction to it more strongly than normal. Thus, key stimuli are subject to the law of summation: with an increase in the parameters of the stimulus, the instinctive reaction increases proportionally. This phenomenon may serve as an explanation for the increased reaction of the gull's chicks to the stick that is cross-streaked with red stripes.

N. Tinbergen drew attention to the quantitative side of the action of symbolic stimuli when studying the response of the female's pursuit in male marigolds during the breeding season. Observations showed that the male takes off not only when approaching individuals of his own species, but also at the sight of other flying insects, as well as small birds and even leaves falling from the trees. The scientist concluded that in this situation, some visual key stimuli are of paramount importance. In this case, chemical stimuli cannot be significant, because the direction of flight of the males is in no way connected with the direction of the wind, which means that they are not guided by smells. Tinbergen and his assistants made mock-ups of paper butterflies and fixed them on a thin fishing line tied to a long fishing rod. Each series of models left only one characteristic external feature: color, size, specific shape. When the rod was twitched, the mock-up butterfly began to move, which caused a mating response in the marigold. The intensity of the reaction was recorded by observation.

The results of the experiment showed that the pursuit reaction caused mockups of all colors, but the males followed the black “butterflies” most actively - the reaction to them was even more pronounced than with brown-colored mockups that corresponded to the natural color of the female velvet. In this case, you should talk about enhancing the visual stimulus - a dark color.

A similar picture was obtained when comparing the intensity of the reaction to the size of the layout. Males most actively pursued larger layouts than the natural size of a female. Such an incentive, as the shape of the object's body, turned out to be not so important for velvet women. Males reacted to models of all forms, with the least effective were long rectangular models. However, observations have shown that this is due to the violation of the nature of the movement of such “butterflies”: it became less “dancing”.

Tinbergen drew attention to another peculiarity of the action of key stimuli, which he called the stimulation adder. The scientist wrote: “... a weakly attractive white model will cause the same percentage of reactions as the black one, if it is shown at a smaller distance than the black one. The effectiveness of a white small model is also noticeably enhanced if you force it to “dance.” Таким образом, недостаточную эффективность одного параметра можно скомпенсировать усиленной привлекательностью совершенно иного параметра... стимулы складываются в некоем „сумматоре стимулирования“, который и заставляет бархатницу реагировать соответствующим образом». ^[9]

Кроме того, Тинберген отметил, что состояние самца определяет, какие стимулы в данный момент входят в этот сумматор. Например, в обычных условиях самцы реагировали только на тон окраски макета (темный или светлый), т. е. сами цвета в сумматор не входили. При кормлении же на макетах, окрашенных в яркий цвет, у самцов отмечалась реакция исключительно на голубые и желтые модели, т. е. знаковым стимулом становился цвет.

Реакция на ключевой стимул не всегда бывает адекватна ситуации и может не привести к желаемому результату. Так, Тинберген описывает явление, получившее название «осечки». Осечка в поведении животного происходит при столкновении со «сверхстимулом». Примером такого «сбоя» может служить выкармливание певчими птицами птенца кукушки. Ключевыми стимулами, заставляющими птицу-родителя кормить птенца, являются большой клюв и ярко окрашенный зев птенца. Оба эти признака у кукушонка имеют «сверхнормальное» выражение. Тинберген пишет: «Вполне возможно, что многие певчие птицы не только кормят птенца кукушонка, но и получают удовольствие из-за его огромного и привлекательного рта». ^[ten]

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

In conclusion, it should be noted that the instinctive behavior of animals is most often caused not by individual factors, but by their complex. At the same time, a combination of external and internal factors is necessary. For example, pigeons feed their chicks, belching "goitre", rich in proteins. The very process of formation in the goiter "milk" is stimulated by the release of the hormone prolactin (internal stimulus). However, the burping reaction is not caused by filling the goiter, but by external stimulation from the side of the chick, which by its weight puts pressure on the goiter of the parent. In winter, even with the goiter filled with food, the pigeon does not have this reaction, because there is no external stimulation.

Структура инстинктивного поведения. Еще в начале XX в. американский исследователь У. Крэг («Влечения и антипатии как составляющие инстинкта», 1918) показал, что любое инстинктивное действие состоит из отдельных фаз. Крэг выделил две фазы, которые получили названия: поисковая (подготовительная) фаза, или аппетентное поведение, и завершающая фаза (завершающий акт).

Крэг показал, что в естественных условиях животные ищут те ключевые стимулы или их комбинации (пусковые ситуации), которые необходимы для осуществления определенной инстинктивной реакции. Например, животные ищут пищу, особей другого пола в период размножения, места для гнездования и т. д. Крэг назвал эти поисковые формы поведения аппетентными, а состояние животного в этот момент – аппетенцией. Воспринимаемые на поисковой фазе поведения промежуточные раздражители не являются для животного целью, они необходимы лишь для того, чтобы привести к восприятию ключевых стимулов завершающего поведения. Завершающая фаза инстинктивного поведения представляет собой само потребление животным необходимых ему элементов среды, именно эта фаза и является непосредственно инстинктивным поведением.

The final phase is hereditarily determined, species-specific, in it is the biological meaning of the whole instinctive action. This phase of behavior consists of a small number of movements, always performed in a clear sequence. It is stereotypical, determined by the structure of the body of the animal. In this phase, only minor individual variations of behavior are possible, which are genetically determined. Acquired components of behavior play virtually no role in the final act, and most often are completely absent. K. Lorenz called the final acts of behavioral reactions endogenous movements, they are type-specific, hereditary and do not require special training.

In contrast to the final act, the search phase is more changeable and adaptive in relation to the conditions, although it is also typical of the species. It is intertwined with innate and acquired forms of behavior, the individual experience of the animal. Orientational research activity of an animal is typical for search behavior. It is due to changes in the appetizing behavior instinctive reactions can be plastic. The preparatory phase is always divided into several stages. It ends when the animal reaches a situation in which the next link in this chain of reactions can be realized. For example, the selection of a breeding territory by a male sometimes requires only a return to the old, last year’s territory, and sometimes it may require a long search and even struggle with other males. According to K. Lorenz,the search phase of the behavioral act should be attributed to purposeful behavior. Various actions are performed at this stage, but they are all subordinate to a specific goal. The search phase is very important and is for the animal the same vital necessity of life, as well as consumption in the final phase. It is the appetizing behavior that is the means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the external world.The search phase is very important and is for the animal the same vital necessity of life, as well as consumption in the final phase. It is the appetizing behavior that is the means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the external world.The search phase is very important and is for the animal the same vital necessity of life, as well as consumption in the final phase. It is the appetizing behavior that is the means of individual adaptation of animals to a changing environment. This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the external world.This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the external world.This phase of the behavioral act includes manifestations of the elementary rational activity of animals. To achieve a certain ultimate goal, the animal chooses a path, while it operates with concepts and laws that connect objects and phenomena of the external world.

Концепцию о двух фазах инстинктивного поведения Крэг построил на данных, полученных в результате изучения пищевого поведения животных. Хищник, испытывающий ощущение голода, начинает искать добычу. Однако вначале он не имеет сведений о ее местонахождении и поэтому его поисковая деятельность пока ненаправленна. Вскоре хищник видит потенциальную жертву, от которой исходит первый ключевой стимул, например размер и детали окраски, и его поисковое поведение переходит на следующую стадию, которая уже имеет определенное направление. Хищник начинает уточнять местоположение, скорость передвижения добычи, ориентируясь при этом и на другие ключевые стимулы. Затем хищник преследует добычу или незаметно подкрадывается к ней, после чего схватывает и умерщвляет. Если в этом есть необходимость, жертва перетаскивается на другое место, где разделывается на куски. Только после этого поведение животного вступает в завершающую фазу, которая включает в себя непосредственное поедание добычи. Все действия животного, связанные с поиском, ловлей и умерщвлением жертвы, относятся к аппетентному поведению. Все они имеют под собой инстинктивную основу, но в большой степени зависят от процесса индивидуального научения, опыта животного и ситуации.

На каждом этапе поискового поведения есть свои подготовительные и завершающие фазы. Окончание одного этапа является сигналом к началу следующего и т. д. Последовательные этапы часто имеют несколько степеней соподчинения, поэтому складывается сложная структура поведения животного. Например, поисковое поведение может привести не к завершающей фазе поведенческого акта, а к сочетанию раздражителей, стимулирующему следующую фазу поискового поведения. Примером может послужить поисковое поведение птиц в период размножения. Вначале происходит выбор территории для гнезда. Когда она найдена, начинается следующая стадия поискового поведения – строительство гнезда, затем следующая – ухаживание за самкой и т. д.

Поведение животного в значительной степени состоит из циклов, которые, в свою очередь, складываются из серии повторяющихся простых актов. Например, птица, занятая постройкой гнезда, совершает это по определенной схеме. Вначале она отправляется на поиски строительного материала, затем, отыскав его, оценивает пригодность. Если материал устраивает птицу, она несет его к гнезду, в противном случае бросает и ищет новый. Прилетев в гнездо, птица определенными движениями вплетает принесенные материалы в его структуру, формирует форму гнезда, а затем вновь улетает на поиски. Этот цикл начинается спонтанно и продолжается до тех пор, пока у птицы есть потребность в достраивании гнезда. Переключение на каждую следующую стадию поведенческой реакции осуществляется при восприятии определенного внешнего стимула. Н. Тинберген приводит пример с самками перепончатокрылых насекомых – филантов (пчелиных волков), которые выкармливают своих личинок медоносными пчелами. Оса, чтобы сделать запасы, летит в места скопления пчел, где беспорядочно летает до тех пор, пока не встретит подходящую жертву. Заметив летящее насекомое, оса подлетает к нему с подветренной стороны и останавливается примерно в 70 см. Если после этого оса уловит запах пчелы, который и будет ключевым раздражителем для перехода на следующую стадию поведенческой реакции, она схватит пчелу. Если пчелу лишить запаха с помощью эфира, оса не станет хватать ее. Следующей стадией поведения осы будет парализация жертвы ударом жала. Для начала этой стадии необходим стимул, связанный с прикосновением кжертве. Если предъявить осе макет пчелы, который на ощупь не похож на нее, но имеет тот же запах, оса не станет жалить такой макет. Таким образом, при прохождении животным различных стадий поведенческой реакции происходит смена стимулов, которые являются для него ключевыми в данный момент.

Состояние аппетенции происходит в условиях очень высокой возбудимости нервных центров, координирующих определенные физиологические реакции. К. Лоренц ввел понятие «специфический потенциал (энергия) действия». Этот потенциал накапливается под действием ряда внешних (температура, освещенность) и внутренних факторов (гормоны) в нервных центрах. Превысив определенный уровень, накопленная энергия высвобождается, после чего начинается поисковая фаза поведенческого акта. При усиленном накоплении «специфической энергии действия» завершающий акт может осуществиться спонтанно, т. е. в отсутствие соответствующих раздражителей, это так называемая «реакция вхолостую».

To explain the neurophysiological mechanism of these phenomena, Lorenz proposed his theory. The basis for this theory was the data of the German physiologist E. Holst.

Holst in his experiments focused on the rhythmic activity of the central nervous system. He noted that rhythmic pulses of impulses can be observed in the isolated ventral nerve chain of the earthworm, which exactly correspond to the contraction of the worm segments. In further research, Holst studied the mechanism of swimming eel. He fixed the middle segments of his body and did not allow them to contract. According to the reflex theory in this case, the back segments of the body will not get irritation, and therefore, also will not be able to contract. However, in reality they start moving after a certain period of time. If you cut the dorsal roots of the spinal cord of an eel, thereby disrupting the transmission of sensory information, the acne will retain the ability to swim movements, and their coordination will not be disturbed. Thus, the movements of the body of an eel are not performed by the mechanism of the reflex arc (depending on external stimuli), but by obeying the rhythmic pulses of impulses in the central nervous system. Experiments of other scientists have confirmed this. For example, it was noted that out-trimmed (with large hemispheres removed) cats can rhythmically contract antagonistic leg muscles, completely devoid of sensory nerves. Tadpoles and fish with one intact sensory nerve retain the ability to swim, normal coordination of movements. This means that the central nervous system is characterized by endogenous automatism, which is independent of external stimuli. At the same time, a minimum level of afferent impulsation is necessary to maintain the excitation, (“specific action energy”) in the corresponding nerve centers at a certain level.

The works of E. Holst and his colleagues confirmed that the level of arousal in the corresponding nerve centers influences the nature of instinctive reactions. The experiments were carried out on chickens, which were stimulated by the shock of brain stem structures. Depending on the location of the irritated structure, researchers have noted elementary behavioral reactions (turns of the head, pecking) or complex behavioral acts (courtship). And if simple reactions always proceeded in approximately the same way, regardless of the parameters of stimulation and environmental conditions, the complex behavioral reactions depended on these factors. So, with a weak current, the rooster attacked a ferret stuffed animal, and with a current increase, even into the hand of a researcher (non-specific stimulus).

Hydraulic model of K. Lorenz. Lorenz proposed a hypothetical model of instinctive reactions. The scientist borrowed the general principles of its work from hydraulics, so it was called the “hydraulic model”.

“Specific action energy” is represented in this model with water, which gradually fills the reservoir (energy accumulation takes place) through an open tap, denoting a continuous flow of potential energy formed in the process of the organism’s vital activity. Water (energy) enters the reservoir (organism) as long as the body feels the need for this form of behavior. Fluid pressure inside the tank is constantly increasing, creating a certain voltage in the system. The outflow of water from the reservoir, indicating the activity of the animal, occurs through the pipes, it is prevented by a valve (central decelerating mechanism). The valve can open in two situations: with a high pressure of water that has accumulated in the tank, or under the influence of gravity of the load suspended from the valve. Cargo refers to the influence of external stimuli specific to a given behavioral act. The increasing pressure of water (the accumulation of specific action energy) and the weight of the load (the strength of external stimuli) sum up their effect on the valve. The stronger the stimulus, the less energy is required. Conversely, the more energy is accumulated, the less external stimulus is needed for the implementation of an instinctive reaction. If the energy level is very high, the valve can open without an external stimulus, due to water pressure. This corresponds to the “idle reaction” (according to Lorentz, “the reaction in a vacuum”). So, Lorenz described the behavior of a starving starling who, in the absence of any stimulus from the external environment, such as an insect, fixes his gaze and "catches" in the air. Inclined tray with holes located at different levels, indicates different types of motor activity of the animal during a behavioral act. The lowest hole corresponds to the motor activity with the lowest threshold, the other holes - forms of activity with a higher threshold. If the valve is only slightly open, water will flow out in a small amount and will only fall into the lower hole area. If the valve opens more and the water flow rate increases, it will fall into the following holes. When the reservoir is emptied (the “specific energy of action” is exhausted), this behavioral act is terminated.

The Lorentz model explains well the situation when the threshold for committing an action decreases with long non-fulfillment (water accumulates in the tank to such a level that a small load is required to open the valve) and the possibility of reactions to nonspecific stimuli (accumulation of water in the tank to such a level when the load for opening the valve is not needed).

The hydraulic model has been repeatedly criticized because of the mechanical design and schematics. The concepts of “specific energy of action” and “key stimuli” in modern zoopsychology correspond to the concept of “specific urge”.

Hierarchical theory of instinct I. Tinbergen. It is noted that stereotyped motor reactions are in a certain relationship. Sometimes instinctive movements appear together, and raising the threshold of one of them causes an increase in the threshold of the second. From this we can conclude that both of them depend on one functional "center". There is some regularity in the sequence of manifestations of actions in complex instinctive reactions. An example is the aggressive collision of fish in the division of territory. In fishes of the cichlid family, direct collisions are preceded by a special demonstration of intimidation. In some species of cichlids, demonstrations are short-lived, and the fish almost immediately turn to attack. In other cichlids, collisions with injuries occur only with equal forces of males, and the demonstrations are very complex and lengthy. There are some cichlids whose fights are completely absent, and the ritualized ceremony of intimidating the enemy is performed until the exhaustion of one of the males, who retreats. Such rituals are sequential actions, starting with a demonstration of the sides of the body, then the dorsal fins rise vertically, followed by strikes from the tail. Opponents can estimate the strength of such a strike with the help of a sideline that perceives the fluctuations of the water. Then the opponents stand in front of each other, and in some species, reciprocal pushes begin with an open maw, while in others the opponents bite each other in open mouths. Rituals continue until one rival gets tired, his coloring turns pale in that case and he swims away. All motor reactions during the ritual demonstration are strictly stereotypical and clearly follow one after the other. Thus, tail strikes cannot begin until the dorsal fin is lifted, and mutual shocks occur only after strikes by the tail.

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

As an illustration, Tinbergen provides a hierarchy of reproduction instinct centers for a male three-needle stickleback. The highest breeding center of the male stickleback is activated by increasing the length of the day, hormonal and other factors. Impulses from the higher center remove the block of appetizing behavior lying near the center. This center is discharged, and the male begins to search for suitable conditions for building a nest (appropriate temperature, territory, necessary soil, vegetation, and shallow water). After selecting such a territory, the braking is removed from the subordinate centers, they are discharged, and the nest building itself begins.

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

Contribution to the study of the hierarchical organization of the instinct made English zoopsychologist XX century. R. Hind. Using the example of stereotypical actions of a big tit, he showed that it is not always possible to arrange these actions in the form of a hierarchical scheme. Some actions may be characteristic of several types of instincts, and in some cases they will be final acts, and in others - only a means for creating conditions in which the final act may occur.

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

N. Tinbergen's scheme provides for the possibility of interaction between the “centers” of various types of behavior, for example, in a situation where one center suppresses another. So, if the male feels hunger while grooming females, he interrupts mating demonstrations and starts searching for food.

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

In a situation when one male stickleback invades the territory of another male, the owner of the territory attacks. He pursues a stranger, and he quickly swims away. When the floating male gets into his territory, they will switch roles, now the pursuer will start to flee. If males clash occurs on the border of their territories, both males will have both attack reaction elements and escape elements in their behavior. The closer the male is to the center of his territory, the more the elements of an attack will be expressed in his behavior. If you move away from the center of the territory, these elements will be suppressed, and the elements of flight will increase.

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

Sometimes in such conflict situations in animals there are so-called “replacement movements”: there is a sort of shift in the activity of the animal. For example, the starling, when meeting a rival, instead of attacking, begins to intensively sort out its plumage with its beak. Having met on neutral territory, the males of the silvery gull stand in a pose of threat, and then suddenly they start to clean the feathers. A similar reaction can be observed in other birds, for example, white geese in such a situation make movements, as when bathing, gray geese shake off, and roosters peck grass. Reactions associated with a shift in activity are innate.

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

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

The variability of the structure of an instinctive behavioral act. The structure of instinctive behavior is extremely complex. The search phase is not always the reaction of finding any environmental agents, it can be negative. In this case, the animal avoids certain irritants and avoids them. In addition, the individual stages of the search behavior may even fall out, then this phase is reduced. Sometimes the search phase does not manifest itself completely, because the final act occurs too quickly. The direction of the search behavior can get off, then the “alien” final act is possible. In some cases, the search phase takes the form of the final, while the present final phase is also preserved. Then the actions in both phases look the same, but have a qualitatively different motivation. In some cases, the final phase is not reached at all, then the instinctive act does not proceed to the end. In animals with a highly developed psyche, the goal of a behavioral act may be the search for stimuli itself, that is, the intermediate stages of search behavior (complex exploratory behavior).

Instinctive behavior and communication. Communication is a physical (biological) and mental (information exchange) interaction between individuals. Communication is necessarily expressed in the coordination of actions of animals, so it is closely connected with group behavior. When communicating, animals necessarily have special forms of behavior that perform the functions of information transfer between individuals. However, some of the actions of the animal acquire a signal value. Communication in this sense is absent in lower invertebrates, and in higher invertebrates, it appears only in rudimentary form. To all representatives of vertebrates, it is inherent in varying degrees.

German ethologist G. Tembrok studied the process of communication in animals and its evolution. According to Timbrock, the real animal communities in which individuals communicate with each other can only be talked about when they begin living together. When living together, several individuals remain independent, but together carry out homogeneous forms of behavior in different areas. Sometimes such joint activities involve the separation of functions between individuals.

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

Forms of communication. According to the mechanism of action, all forms of communication differ in the channels of information transfer. Optical, acoustic, chemical, tactile and other forms are distinguished.

Among the optical forms of communication, the most important place is occupied by expressive postures and gestures, which constitute “demonstration behavior”. This behavior consists of the demonstration to animals of certain parts of their body, which, as a rule, carry specific signals. These can be brightly colored areas, additional structures such as combs, decorating feathers, etc. However, some parts of the animal’s body can visually increase in volume, for example, by ruffling feathers or hair. The signal function can also be performed by specific movements of the body or its individual parts. By performing these movements, the animal may exhibit colored areas of the body. Sometimes such demonstrations are made with exaggerated intensity.

В эволюции поведения появляются специальные двигательные акты, которые отделились от остальных форм поведения тем, что утратили первичную функцию и приобрели чисто сигнальное значение. Примером может служить движение клешни у манящего краба, которое он выполняет при ухаживании за самкой. Такие движения получили название «аллохтонные». Аллохтонные движения видотипичны и стереотипны, их функция – передача информации. Другое их название – ритуализованные движения. Все ритуализованные движения условны. Они очень жестко и четко закреплены генетически, относятся к типичным инстинктивным движениям. Именно такая консервативность движений обеспечивает правильное восприятие сигналов всеми особями независимо от условий жизни.

Чаще всего и в наибольшем количестве ритуализованные движения наблюдаются в сфере размножения (прежде всего это брачные игры) и борьбы. Они передают одной особи информацию о внутреннем состоянии другой особи, о ее физических и психических качествах.

Инстинктивные движения и таксисы. Таксисами называют врожденные, наследственно определяемые реакции на определенные компоненты окружающей среды.

По своей природе таксисы сходны с инстинктивными движениями, но имеют и отличие. Инстинктивные движения всегда возникают в ответ на ключевые раздражители, таксисы же проявляются при действии направляющих ключевых раздражителей. Эта особая группа стимулов сама по себе не способна вызвать начало или конец какого-либо инстинктивного движения. Направляющие ключевые раздражители стимулируют только смену направления протекания этой реакции. Таким образом, таксисы производят общую ориентацию инстинктивных движений. Таксисы тесно связаны с врожденными двигательными координациями и в совокупности с ними составляют инстинктивные реакции или их цепи.

Кроме таксисов, существуют кинезы. При кинезах не происходит ориентации тела животного относительно раздражителя. В данном случае раздражители либо вызывают изменение скорости передвижения животного, либо меняется частота поворотов тела. При этом происходит смена положения животного относительно раздражителя, но ориентация его тела остается прежней.

При таксисах тело животного принимает определенное направление. Таксис может сочетаться с движениями, в этом случае животное будет двигаться по направлению к раздражителю или от него. Если двигательная активность направлена в сторону благоприятных для животного условий среды, таксис будет положительным (активность животного направлена к раздражителю). Если, напротив, условия не ценны для животного или опасны, таксис будет отрицательным (активность животного направлена от раздражителя).

В зависимости от природы внешних стимулов таксисы делятся на фототаксисы (свет), хемотаксисы (химические раздражители), термотаксисы (температурные градиенты), геотаксисы (сила тяжести), реотаксисы (течение жидкости), анемотаксисы (поток воздуха), гидротаксисы (влажность среды) и др.

Различают несколько типов таксисов (по Г. С. Френкелю и Д.Л. Ганну; Fraenkel GS, Gunn DL «The Orientation of Animals», 1940).

1. Клинотаксис. При клинотаксисе для ориентации тела относительно стимула необязательна способность рецептора к определению направления источника раздражения. Животное сравнивает интенсивность раздражения с разных сторон простым поворотом органов, несущих рецепторы. Примером может служить установление траектории движения на свет личинки мухи. Фоторецепторы у личинки расположены на переднем конце тела, при ползании она отклоняет голову то в одну сторону, то в другую. Сравнение интенсивности