Lecture
When studying unconditioned reflexes and instincts, it became necessary to create a classification of the main forms of animal behavior. For the first time attempts of such a classification were made in the pre-Darwinian period, but they reached their greatest development at the beginning of the 20th century. So, I.P. Pavlov divided the innate elements of behavior into indicative, defensive, nutritional, sexual, parental and childish. When new data on conditioned-reflex activity of animals appeared, it became possible to create more fractional classifications. For example, orienting reflexes began to be subdivided into indicative and research proper, an indicative reflex aimed at searching for food, called approximate-alimentary, etc.
Another classification of forms of behavior was proposed by A.D. Slonim in 1949 in the article "On the relationship between unconditioned and conditioned reflexes in mammals in phylogenesis." In his scheme, there were three main groups of reflexes:
1) reflexes aimed at preserving the internal environment of the body and the constancy of the substance. This group includes food behavior that ensures the constancy of the substance, and homeostatic reflexes that ensure the constancy of the internal environment;
2) reflexes aimed at changing the external environment of the body. These include defensive behavior and environmental, or situational, reflexes;
3) reflexes associated with the preservation of the form. These include sexual and parental behavior.
Later, Pavlov’s school scientists developed other classifications of unconditioned reflexes and conditioned reflexes formed on their basis. For example, the classification of D.A. Biryukova, created in 1948, N.A. Rozhansky (1957). These classifications were quite complex, they included both the proper reflexes of behavior and the reflexes of regulation of individual physiological processes, and therefore did not find wide application.
R. Hind gave several classifications of types of behavior based on certain criteria. The scientist believed that there are a lot of such criteria to choose from and in practice the most often selected criteria are suitable for the particular problem that is being considered. He mentioned three main types of criteria for classification.
1. Classification for immediate reasons. According to this classification, types of activity that are determined by the same causal factors are combined into one group. For example, all types of activity are combined, the intensity of which depends on the male sex hormone (male sexual behavior), types of activity associated with male-rival irritants (agonistic behavior), etc. This type of classification is necessary to study the behavior of an animal, it is convenient to use in practice.
2. Functional classification is based on the evolutionary classification of activity types. Here the categories are smaller, for example, such types of behavior as courtship, migration, hunting, and threat are distinguished. Such a classification is justified, while categories are used to study functions, but it is rather controversial, since identical elements of behavior in different species may have different functions.
3. Classification by origin. In this group, classification according to general ancestral forms, based on a comparative study of closely related species, and classification by acquisition method, which is based on the nature of a change in a behavioral act in the process of evolution, stand out. As examples of categories in these classifications, we can distinguish behavior acquired as a result of learning and ritualized behavior.
Hind emphasized that any classification systems based on different types of criteria should be considered as independent.
For a long time, a classification based on the classification of Pavlov's reflexes has been popular among ethological scholars. Its formulation was given by G. Tembrock (1964), who divided all forms of behavior into the following groups:
1) the behavior determined by the metabolism (eating and eating, urination and defecation, food storage, rest and sleep, sipping);
2) comfortable behavior;
3) defensive behavior;
4) behavior associated with reproduction (territorial behavior, copulation and mating, care for offspring);
5) social (group) behavior;
6) construction of nests, holes and shelters.
Let us dwell on some forms of behavior.
The behavior determined by the metabolism. Nutritional behavior. Nutritional behavior is common to all members of the animal world. Its forms are very diverse and species-specific. The basis of eating behavior is the interaction of the central mechanisms of excitation and inhibition. The constituent elements of these processes are responsible both for the reaction to various food stimuli, and for the nature of movements when eating. A certain role in the formation of eating behavior is played by the individual experience of the animal, in particular, the experience that determines the rhythms of behavior.
The initial phase of eating behavior is a search behavior caused by arousal. Search behavior is determined by the deprivation of animal food and is the result of increased reactivity to external stimuli. The ultimate goal of search behavior is to find food. In this phase, the animal is particularly sensitive to stimuli that indirectly indicate the presence of food. Types of stimuli depend on the availability and taste of different types of food. Signs that serve as irritants are common to different types of food or characterize its specific type, which is most often observed in invertebrates. For example, for bees, coloring of the flower rims can be such an irritant, and for termites - the smell of rotting wood. All these stimuli cause different types of activity. Depending on the circumstances and the type of animal, this may be the setting of prey, its preliminary preparation and absorption. For example, wolves have a certain way of hunting for different types of ungulates, while the lynx hunts for all kinds of prey equally (jumping from ambush to the nape of the victim). In predatory mammals, there are certain "rituals" when eating prey. Weasel eats mouse-like rodents from the head, and when there is a lot of prey, it is content only with the brain of the victim. Large predators also prefer to eat the prey, starting with the muscles of the neck and viscera.
When the animal begins to saturate, the feedbacks caused by irritation of the receptors of the mouth, throat and stomach, shift the equilibrium in the direction of inhibition. This also contributes to the change in the composition of the blood. Usually the processes of inhibition are ahead of the compensatory abilities of the tissues and proceed at different speeds. In some animals, the processes of inhibition only affect the final act of eating behavior and do not concern the behavior of the exploratory. Therefore, many well-fed mammals continue to hunt, which is typical, for example, of a kunim, to some large cats.
There are many different factors that determine the attractiveness of different types of food, as well as the amount of food consumed. These factors are best studied on the example of rats. In these rodents, differing in complex behavior, the novelty of food can serve as a factor contributing to both an increase in food eaten and a decrease in its quantity. Monkeys more often eat new food for themselves in small doses, but if a monkey notices that its congeners eat this food, the amount of food eaten increases markedly. In most mammals, young animals are the first to try new foods. In some schooling mammals and birds, individual individuals more often try unfamiliar food, being surrounded by their relatives, and are very cautious about it, being isolated. The amount of food consumed can also depend on the amount of food available. For example, in the autumn period, bears eat pears in orchards in noticeably more quantities than from detached trees.
For food can be attributed to such a widespread behavior as storing food. To provide food for insect larvae, it is reduced to the activity of laying eggs on living objects (gadflies), the manifestation of parasitism, and the activity of grave diggers. Feed storage is widespread among mammals. For example, food is stored by many species of predators, and the forms of storing them are extremely diverse. A domestic dog can simply bury a piece of meat left over from dinner, and an ermine, a marten, arrange entire warehouses consisting of corpses of small rodents. Many species of rodents store food, some of them (hamsters, saccular rats) have special cheek bags in which they carry the food. For most rodents, food storage times are strictly limited, in most cases they are confined to autumn, when seeds, nuts and acorns ripen.
Indirectly, urinary secretion and defecation can be correlated with food behavior, or rather, with behavior determined by metabolism. In most animals, urination and defecation are associated with specific postures. The mode of these acts and the characteristic postures are observed both in animals and in humans. The latter was proved by numerous experiments carried out during the wintering in the Arctic.
States of rest and sleep, according to Timbrock, relate to the behavior determined by the metabolism, but many scientists associate them with comfortable behavior. It was found that the postures of rest and postures taken by animals during sleep are species-specific, as are certain types of movement.
Comfortable behavior. These are diverse behavioral acts aimed at caring for the body of an animal, as well as various movements that do not have a specific spatial direction and location. Comfortable behavior, namely, that part of it that is related to the care of an animal for its body, can be considered as one of the options for manipulation (for more details on this see 5.1, 6.3), and in this case the body of the animal acts as an object of manipulation.
Comfortable behavior is widespread among various representatives of the animal world, from the most underdeveloped (insects that clean wings with limbs) to fairly well-organized ones, from which it sometimes acquires a group character (grooming, or mutual searching in apes). Sometimes, to perform comfortable actions, an animal has special organs, for example, some animals use the toilet claw for special hair care.
In comfort behavior, several forms can be distinguished: cleansing the hair and skin of the body, scratching a certain part of the body against the substrate, scratching the body with the limbs, rolling on the substrate, bathing in water, sand, shaking the hair, etc.
Comfortable behavior is typical, the sequence of actions to cleanse the body, the dependence of a particular method on the situation is innate and manifest in all individuals.
Close to comfortable behavior adjacent posture of peace and sleep, the whole complex of actions associated with these processes. These poses are also hereditarily fixed and species-specific. Studies on the study of resting and sleeping postures in bison and bison, conducted by Soviet biologist M.A. Derdyagina, allowed to isolate from these animals 107 types of typical postures and gestures belonging to eight different spheres of behavior. Of these, two-thirds of the movements fall into the category of comfort, rest and sleep. Scientists noted an interesting feature: the differences in behavior in these areas in young bison, bison and their hybrids are formed gradually, at a later age (two to three months).
Sexual behavior describes all the diverse behavioral acts associated with the process of reproduction. This form refers to the most important forms of behavior, as it is associated with the continuation of the genus.
According to most scientists, key stimuli (releasers) play an important role in sexual behavior, especially in lower animals . There are a great number of releasers, which, depending on the situation, can cause either rapprochement of sexual partners, or a fight. The action of the releaser directly depends on the equilibrium of the aggregate of its constituent stimuli. This was shown in the experiments of Tinbergen with a three-needle stickle stick, where the red coloring of the belly of the fish acted as an irritant. When using different models, it was found that the stickie males most aggressively react not to models that are completely painted red, but to objects that are closest to the natural color of the fish. Also aggressively, sticklebacks reacted to models of any other shape, the lower part of which was colored red, imitating the color of the abdomen. Thus, the reaction to the release agent depends on a combination of features, some of which can compensate for the lack of others.
When studying the releasers, Tinbergen used the comparison method, trying to figure out the origins of the marriage rituals. For example, in ducks, the courtship ritual comes from movements that serve to care for the plumage. Most of the releasers on display during the mating games are reminiscent of unfinished movements, which in everyday life are used for completely different purposes. Many birds in marriage dances can recognize the threat poses, for example, in the behavior of seagulls during mating games, there is a conflict between the desire to attack a partner and hide from him. Most often, the behavior is a series of individual elements that correspond to opposite trends. Sometimes the behavior can be seen manifestation of heterogeneous elements at the same time. In any case, in the process of evolution, any movements underwent strong changes, ritualized and turned into releasers. Most often, the changes went in the direction of enhancing the effect, which may consist in their repeated repetition, as well as an increase in the speed of their execution. According to Tinbergen, evolution was aimed at making the signal more visible and recognizable. The limits of expediency are reached when the hypertrophied signal begins to attract the attention of predators.
To synchronize sexual behavior, it is necessary that the male and female are ready to breed at the same time. Such synchronization is achieved with the help of hormones and depends on the season and the length of the daylight hours, but the final “fitting” occurs only when a male and female meet, as has been proven in a number of laboratory experiments. In many species of animals, the synchronization of sexual behavior is developed at a very high level, for example, in the trickles during the mating dance of the male, each movement of the male corresponds to a certain movement of the female.
In most animals, sexual behavior is distinguished by individual behavioral blocks, which are performed in a strictly defined sequence. The first of these blocks is most often the ritual of pacification. This ritual is evolutionary aimed at removing obstacles to the convergence of marriage partners. For example, in birds, females usually do not tolerate the touches of other individuals of their species, while males are prone to fights. During sexual behavior, the male is prevented from attacking the female by differences in the plumage. Often, the female takes the position of a chick requesting food. In some insects, pacification takes on peculiar forms, for example, in the case of cockroaches, the glands under the sheaths emit a kind of secret that attracts the female. The male lifts the wings and, while the female licks the secretions of the odorous glands, proceeds to mate. In some birds, as well as in spiders, the male brings a kind of gift to the female. Such pacification is essential for spiders, because without a gift, the male at risk of being eaten during grooming.
The next phase in sexual behavior is the discovery of the marriage partner. For this there are a huge number of different ways. In birds and insects, singing most often serves this purpose. Usually, the male sings the songs, in his repertoire there are a lot of various sound signals, from which rival males and females receive exhaustive information about his social and physiological status. In birds, male bachelors sing most intensely. Singing stops when the sexual partner is found. Butterflies often use smells to attract and discover a marriage partner. For example, in hawk moths, females are attracted to males with the secretion of an odorous gland. Males perceive this smell even in very small doses and can fly to the female for a distance of up to 11 km.
The next stage of sexual behavior is the recognition of the marriage partner. It is most developed in higher vertebrates, in particular birds and mammals. The irritants on which recognition is based are weaker than stimuli-releasers, and, as a rule, they are individual. It is believed that the birds that form a constant pair, distinguish partners in appearance and voice. Некоторые утки (шилохвосты) способны распознать партнера на расстоянии 300 м, у большинства же птиц порог узнавания снижен до 20–50 м. У некоторых птиц формируется довольно сложный ритуал узнавания, например у голубей приветственный ритуал сопровождается поворотами и поклонами, и малейшее его изменение вызывает у партнера беспокойство. У белых аистов церемония приветствия сопровождается щелканьем клювом, причем голос партнера птицы узнают на значительном расстоянии.
Как правило, брачные ритуалы млекопитающих отличаются меньшим разнообразием, чем ритуалы рыб и птиц. Самцов чаще всего привлекает запах самок, кроме того, основная роль при поиске партнера принадлежит зрению и кожной чувствительности головы и лап.
Практически у всех животных сближение с половым партнером стимулирует многочисленные нейрогуморальные механизмы. Большинство этологов считает, что смысл сложных брачных ритуалов у птиц заключается в общей стимуляции механизма спаривания. Практически у всех земноводных, у которых брачные ритуалы довольно бедны, важная роль в стимуляции нейрогуморальных механизмов принадлежит тактильным раздражителям. У млекопитающих овуляция может наступать как после спаривания, так и до него. Например, у крыс копуляция не влияет на механизмы, связанные с созреванием яйцеклеток, а у крольчих овуляция наступает только после спаривания. У некоторых млекопитающих, например у свиней, для полового созревания самки достаточно одного присутствия самца.
Оборонительное поведение у животных впервые описано еще Ч. Дарвином. Обычно оно характеризуется определенным положением ушей, шерсти у млекопитающих, кожных складок у пресмыкающихся, перьев на голове у птиц, т. е. характерной мимикой животных. Оборонительное поведение – это реакция на изменение во внешней среде. Оборонительные рефлексы могут возникать в ответ на любые факторы внешней или внутренней среды: звуковые, вкусовые, болевые, термические и другие раздражители. Оборонительная реакция может носить как местный характер, так и принимать характер общей поведенческой реакции животного. Поведенческая реакция может выражаться и в активной обороне или нападении, и в пассивном замирании на месте. Двигательные и оборонительные реакции у животных разнообразны и зависят от образа жизни индивида. Одиночно живущие животные, например заяц, убегая от врага, старательно запутывают след. Животные, обитающие в группах, например скворцы, при виде хищника перестраивают свою стаю, стараясь занять наименьшую площадь и избежать нападения. Проявление оборонительной реакции зависит как от силы и характера действующего раздражителя, так и от особенностей нервной системы. Любой раздражитель, достигающий известной силы, может вызвать оборонительную реакцию. В природе чаще всего оборонительное поведение связано with conditioned (signal) stimuli that have formed in different species in the process of evolution.
Another form of defensive behavior is represented by physiological changes during a passive-defensive reaction. In this case, the inhibition dominates, the movements of the animal slow down sharply, and more often it lurks. In some animals, a special musculature is involved in the passive-defensive reflex. For example, the hedgehog at the time of danger collapses into a ball, his breathing is sharply limited, the tone of skeletal muscles decreases.
A special form of defensive behavior includes avoidance reactions, by which animals minimize their exposure to dangerous situations. In some animals, fear signaling stimuli induce such a reaction without prior experience. For example, for small birds, the silhouette of a hawk serves as a signal stimulus, and for some mammals, the characteristic color and smell of poisonous plants. Avoidance also refers to highly specific reflexes.
Агрессивное поведение. Агрессивным чаще всего называют поведение, адресованное другим особям, которое приводит к нанесению повреждений и зачастую связано с установлением иерархического статуса, получением доступа к какому-либо объекту или права на определенную территорию. Различают внутривидовые столкновения и конфликты, возникающие в ситуации «хищник – жертва». Чаще всего эти формы поведения вызываются различными внешними раздражителями, состоят из различных организованных комплексов движений и определяются разными нервными механизмами. Агрессивное поведение направлено на другую особь, раздражители могут быть зрительными, слуховыми и обонятельными. Агрессия возникает в первую очередь из-за близости другой особи.
According to many researchers, aggression may occur as a result of the conflict between other types of activity. This has been proven in numerous laboratory experiments. For example, in domestic pigeons, aggressive behavior directly depended on food reinforcement: the more hungry the birds, the greater the increased aggressiveness.
В естественных условиях агрессия чаще всего является реакцией на близость другого животного, которая возникает либо при нарушении индивидуальной дистанции, либо при приближении к важным для животного объектам (гнездо, индивидуальная территория). В этом случае приближение другого животного может вызвать как оборонительную реакцию с последующим бегством, так и агрессивную в зависимости от иерархического положения особи. Агрессия зависит также от внутреннего состояния животного. Например, у многих воробьинообразных наблюдаются кратковременные стычки в зимних стаях, где птицы в зависимости от внутреннего состояния поддерживают индивидуальную дистанцию от нескольких метров до нескольких десятков метров.
У большинства видов животных агрессивные конфликты происходят в весеннее время, когда активны половые железы. Интенсивность конфликтов напрямую зависит от стадии брачного цикла. В пик брачной активности почти у всех птиц агрессию вызывает соперник, появившийся в непосредственной близости от участка. Подобные явления наблюдаются и у некоторых территориальных видов рыб.
В результате многочисленных исследований было выяснено, что для вызывания агрессии внешние раздражители играют более важную роль, чем внутреннее состояние. Последнее чаще всего влияет на избирательность восприятия раздражителей, а не на интенсивность агрессивного поведения. Большинство этих данных получено при изучении поведения птиц из отряда воробьинообразных, но подобное явление наблюдалось и у раков-отшельников, а также у некоторых территориальных видов рыб.
Extensive research on aggressive activities was conducted by K. Lorenz, who devoted a number of scientific works to this phenomenon. He conducted a large number of experiments on the study of the aggressive behavior of rats, which helped to derive the basic laws of aggressive behavior of man as a biological species.
Территориальное поведение впервые появляется у кольчатых червей и низших моллюсков, у которых все процессы жизнедеятельности приурочены к участку, где располагается убежище. Однако такое поведение еще не может считаться полноценным территориальным, потому что животное никак не маркирует территорию, не дает знать другим особям о своем присутствии на ней, не защищает ее от вторжения. Для того чтобы можно было говорить о полноценно развитом территориальном поведении, необходимо развитие у животного перцептивной психики, оно должно иметь возможность дать остальным особям информацию о своих правах на эту территорию. В этом процессе чрезвычайную важность приобретает маркировка территории. Территория может маркироваться нанесением пахучих меток на предметы по периферии участка, звуковыми и оптическими сигналами, а в качестве оптических сигналов могут выступать вытоптанные участки травы, обгрызенная кора деревьев, экскременты на ветках кустарников и другие. Животные с настоящим территориальным поведением, как правило, активно защищают свой участок от остальных особей. Особенно эта реакция проявляется у животных по отношению к особям своего вида и того же пола. Как правило, такое поведение приурочено или проявляется в особо яркой форме в период размножения.
In a fairly developed form, territorial behavior manifests itself in dragonflies. And Heimer was observing the male equal winged dragonfly beauties. It was noted that the males of these insects occupy individual areas in which functional recreation and reproduction areas are distinguished. Egg laying occurs in the breeding zone, the male attracts the female to this zone with the help of a special ritualized flight. Males perform all their functions within their territory, except for the evening rest, which takes place outside its borders. The male marks his plot, actively protects him from other males. It is interesting to note that battles between them take place in the form of rituals, and, as a rule, they do not reach the present clash.
Большой сложности, как показали исследования российского этолога А.А. Захарова, достигает территориальное поведение муравьев. У этих насекомых существует два разных типа использования кормовых участков: совместное использование угодий несколькими семьями и использование участка населением одного гнезда. Если плотность вида невелика, участки не охраняются, если же плотность достаточно высока, кормовые участки делятся на охраняемые территории, между которыми есть небольшие неохраняемые участки. Наиболее сложно поведение у рыжих лесных муравьев. Их территории, которые строго охраняются, очень велики, через них пролегает разветвленная сеть тропок. При этом каждая группа муравьев использует определенный сектор муравейника и определенные тропы, которые к нему прилегают. Таким образом, общая территория муравейника у этих насекомых делится на территории отдельных групп, между которыми располагаются нейтральные пространства. Границы таких территорий маркируются пахучими метками.
Многие высшие позвоночные, в частности млекопитающие, птицы и рыбы, держатся в центре хорошо известного им участка, границы которого они ревностно охраняют и тщательно маркируют. У высших млекопитающих хозяин участка, даже находящийся на более низкой ступени иерархической лестницы, с легкостью прогоняет сородича, нарушившего границу. Хозяину территории для этого достаточно принять угрожающую позу, и соперник отступает. Истинная территориальность встречается у грызунов, хищников и некоторых обезьян. У видов, для которых характерна беспорядочность половых отношений, нельзя выделить индивидуальную территорию.
Territoriality is expressed in many fish. Usually their territorial behavior is closely related to the process of reproduction, which is typical of many cichlids, as well as sticklebacks. The desire to choose the territory of the fish is innate, in addition, it is due to the system of landmarks used by the fish. Protection of the territory of fish is most pronounced in the sexual period.
In birds, territorial behavior has reached high development. Some scientists have developed a classification of areas in different species of birds by type of use. Such birds may have separate areas for nesting, mating, as well as separate areas for wintering or overnight. To protect the territory of the birds often use singing. The basis of territorial behavior is intraspecific competition. Selects the site and attracts the female, as a rule, more aggressive male. The size of the territory in birds is species specific. Territoriality in birds does not always exclude gregarious behavior, although most often these forms of behavior are not observed simultaneously.
Parental behavior. All animals can be divided into two groups. The first group includes animals, the females of which, even at first birth, demonstrate parental behavior. The second group includes animals whose females improve their parental behavior throughout their lives. This classification was first developed in mammals, although various forms of parental behavior are observed in other groups of animals.
Typical representatives of animals of the first group are mice and rats, they are already caring for offspring from the very first day, and many researchers did not notice significant differences in this between young and experienced females. Animals of the second group include great apes and corvids. Young female chimpanzees are helped by more experienced relatives to care for the young, otherwise the newborn may die due to improper care.
Parental behavior refers to one of the most difficult types of behavior. As a rule, it consists of a series of interrelated phases. In the lower vertebrates, recognition of the young by the parents is the main one in the parental behavior, and the young by the parents. Here, imprinting in the early stages of caring for offspring plays an important role. Fish fry instinctively huddle together and follow adults. Adults try to swim slowly and keep their young within sight. In case of danger, adults protect young.
The parental behavior of birds is much more complicated. As a rule, it begins with laying eggs, since the nest building phase refers more to sexual behavior and often coincides with a courtship ritual. The presence of the nest serves as a stimulating effect on the laying of eggs, and in some birds - its construction. In some birds, a nest with full clutch may suspend further egg laying for some time, and vice versa, incomplete clutch stimulates this process. In the latter case, the birds can lay several times more eggs than under normal conditions.
The next phase of the parental behavior of birds is egg recognition. A number of birds do not have selectivity; they can incubate eggs with any color and even dummies that have only a distant resemblance to eggs. But many birds, particularly passerine, well distinguish their eggs from the eggs of their relatives. For example, some Slavs reject the eggs of relatives, similar in color, but slightly different shape.
The next phase of parental behavior of birds is hatching. It is characterized by an exceptional variety of behaviors. Both male and female or both parents can incubate eggs at the same time. Hatching can take place from the first, second egg or after completion of laying. The hatching bird can sit on the nest tightly or throw the nest at the first sign of danger. Hatching in weedy chickens has reached the highest skill, when the male watches the thermoregulation in a kind of incubator of decaying vegetation, and its construction can take several months. In species, in which the male incubates, his striving for this action is synchronous with the time of laying eggs. In females, it is determined by physiological processes.
The next phase of parental behavior comes after hatching chicks. Parents begin to feed them with semi-digested food. The reaction of chicks is innate: they reach for the tip of the parent's beak for food. In this case, the most often favored is the coloring of the beak of an adult bird; in some birds it changes at this time. Adult birds most often react to the voice of the chick, as well as to the coloring of the pharynx in the chick asking for food. As a rule, it is the presence of chicks that makes parents take care of them. In experimental conditions in chickens, parental behavior can be maintained for many months by constantly placing chicks on it.
Mammals also differ in complex parental behavior. The initial phase of parental behavior in them is the construction of a nest, which is largely species-typical. The impetus for the construction of a nest in females is a specific phase of pregnancy. Rats can start building a nest already in the early stages of pregnancy, but usually it is not fully completed and is only a pile of building material. Present construction begins three days before delivery, when the nest takes on a certain shape, and the female rat becomes less mobile.
Immediately before the birth, the female mammals change the order of licking of separate parts of the body. For example, in the last week of pregnancy, they more often lick the perineum and, more and more rarely, the sides and front legs. Female mammals give birth in a wide variety of positions. Their behavior during childbirth can vary quite strongly. As a rule, females carefully lick newborns, snack on their umbilical cord. Most mammals, especially herbivores, greedily eat the placenta.
Mammalian behavior during feeding of pups is very complex . The female collects the young, gives them the nipples to which they stick. The feeding period of different species varies from two weeks in rodents to one year in some marine mammals. Even before the end of lactation, the young undertake brief sorties from the nest and begin to try additional feed. At the end of lactation, the young go on independent feeding, but continue to pursue the mother, try to suck her, but the female less and more often allows them to do so. She pressed her belly to the ground or trying to drastically run aside.
Another characteristic manifestation of parental behavior is dragging cubs. If conditions become unsuitable, animals can build a new nest and drag their offspring there. The instinct of dragging is especially strong in the first few days after giving birth, when the female pulls not only her own, but also foreign calves, as well as foreign objects, into the nest. However, this instinct quickly fades away, and within a few days the females well distinguish their young from others. Ways to transfer the young in different species are different. Dragging itself can be caused by various stimuli. Most often, this reaction is caused by the cries of the young, as well as their characteristic odor and body temperature.
Special forms of parental behavior include punishment, which is expressed in some predatory mammals, in particular dogs. Domestic dogs can punish puppies for various offenses. The female growls at the young, shakes them, holding the collar, or presses paw. With the help of punishment, the mother can quickly wean the puppies to look for her nipples. In addition, dogs punish puppies, when they move away from them, can separate the fighting.
Social (group) behavior. This type of behavior is represented in lower invertebrates only in the embryonic form, since they do not have special signaling actions for making contacts between individuals. Group behavior in this case is limited to the colonial lifestyle of some animals, for example coral polyps. In higher invertebrates, on the contrary, group behavior is already manifested fully. First of all, this applies to insects whose lifestyle is associated with complex communities that are highly differentiated in structure and function — bees, ants, and other social animals. All individuals belonging to the community differ in their functions, food-producing, sexual and defensive forms of behavior are distributed among them. Observed specialization of individual animals in function.
In this form of behavior, the nature of the signal is of great importance, with the help of which individuals communicate with each other and coordinate their actions. In ants, for example, these signals are of a chemical nature, other types of receptors are much less significant. It is by smell that ants distinguish individuals of their community from strangers, living individuals - from the dead. Ants larvae secrete chemicals to attract adult individuals that can feed them.
With group lifestyles, great importance is attached to the coordination of the behavior of individuals when threatened by the community. Ants, as well as bees and wasps, are guided by chemical signals. For example, in case of danger, “alarm substances” are released, which spread over the air for a short distance. Such a small radius helps to pinpoint the place where the threat comes from. The number of individuals emitting a signal, and hence its strength, increases in proportion to the increase in danger.
The transfer of information may be carried out in other ways. As an example, consider the "dancing" of bees, carrying information about food objects. The dance pattern indicates the proximity of the feed. This is how the well-known Austrian ethologist Karl von Frisch (1886–1983), who had been studying the social behavior of these insects for many years, characterized bee dances: “... if it (a feed object - Auth.) Is near a hive (at 2–5 meters from it), then produced "dance-push": a bee randomly runs along the honeycomb, wagging its abdomen from time to time; if the feed is found at a distance of up to 100 meters from the hive, then a “circular” run is performed, which consists of running around in a circle, alternately clockwise and against it. If nectar is found at a greater distance, then a wagging dance is performed. These are jogs in a straight line, accompanied by wagging movements of the abdomen, returning to the starting point either to the left or to the right. The intensity of wagging movements indicates the distance of the find: the closer the feed object is, the more intense the dance is performed ”. [eleven]
In all the examples given, it is clearly noted that information is always transmitted in a transformed, conditional form, with the spatial parameters being translated into signals. The instinctive components of communication have been most developed in such a complex phenomenon as the ritualization of behavior, especially sexual, which has already been mentioned above.
Social behavior in higher vertebrates is very diverse. There are many classifications of various types of animal associations, as well as the behavior of animals within different groups. Birds and mammals have various transitional forms of organization from a single family group to a true community. Within these groups, relationships are built primarily on a variety of sexual, parental, and territorial behaviors, but some forms are characteristic only of animals living in communities. One of them is the exchange of food - trophallaxis. It is most developed in social insects, but is also found in mammals, for example, in wild dogs, which exchange food, belching it.
Social behavior also includes group care for the offspring. It is observed in penguins: young cubs are collected in separate groups, which are looked after by adults, while parents get their own food. In hoofed mammals, such as elk, the male owns a harem of several females that can jointly care for offspring.
Social behavior includes joint work, which is controlled by a system of sensory regulation and coordination. Such joint activities consist mainly in construction, which is impossible for an individual, for example, the construction of an anthill or the construction of dams by beavers on small forest rivers. In ants, as well as colonial birds (rooks, coastal swallows), there is a joint defense of colonies against predator attacks.
It is believed that for social animals only the presence and activity of a relative serve as an incentive for the start of social activities. Such stimulation causes in them a set of reactions that are impossible in single animals.
Exploratory behavior determines the desire of animals to move and inspect the environment, even in those cases where they do not experience hunger or sexual arousal. This form of behavior is innate and necessarily precedes learning.
In the event of an unexpected external influence, all higher animals react to a source of irritation, trying to explore an unfamiliar object using all available sensory organs. Once in an unfamiliar environment, the animal moves erratically, examining everything that surrounds it. At the same time, various types of behavior are used, which can be not only species-typical, but also individual. You should not identify exploratory behavior with the game, which it looks like.
Some scientists, for example R. Hind, draw a clear boundary between the orientation reaction, when the animal is motionless, and active research, when it moves relative to the object being examined. These two types of exploratory behavior mutually suppress each other. You can also highlight the superficial and deep research behavior, as well as to draw a distinction, based on the sensory systems involved in it.
Exploratory behavior, especially at first, depends on the reaction of fear and on the experience of the animal. The probability that this situation will cause either a fear reaction or exploratory behavior depends on the internal state of the animal. For example, if a stuffed owl is placed in a cage with small birds of a passerine detachment, at first they rarely approach it, experiencing a fear reaction, but gradually shorten this distance and continue to exhibit only exploratory behavior in relation to the stuffed animal.
In the initial stages of the study of the object, the animal may exhibit other forms of activity, for example, feeding behavior, cleaning of wool. Exploratory behavior depends largely on the degree of hunger experienced by animals. Usually, hunger reduces exploratory activity, but hungry mammals (rats) are much more likely than well-fed, leave the familiar environment and go to research new territories.
Investigative behavior is closely related to the internal state of the animal. The effectiveness of research reactions depends on what the animal considers familiar based on its experience. It also depends on the internal state whether the same stimulus will cause fear or a research reaction. Sometimes other types of motivations come into conflict with exploratory behavior.
Exploratory behavior can be very stable, especially in higher mammals. For example, rats may explore an unfamiliar object for several hours and, even in familiar surroundings, exhibit exploratory behavior that may enable them to research something. Some scientists believe that exploratory behavior differs from other forms of behavior in that the animal is actively looking for increased stimulation, but this is not entirely true, because both nutritional and sexual behavior includes the search for final stimuli, which brings these behaviors closer to exploratory ones.
Research behavior is aimed at eliminating inconsistencies between the familiar situation model and the central consequences of the new perception. This brings him closer, for example, with nest building, which is also aimed at eliminating inconsistencies between stimuli in the form of a complete and an unfinished nest. But with exploratory behavior, the discrepancy is eliminated not because of a change in stimuli, but as a result of a restructuring of the nervous model, after which it begins to conform to the new situation. In this case, the stimuli lose their novelty, and research behavior will be directed to the search for new stimuli.
Research behavior inherent in highly developed animals is an important step before learning and developing intelligence.
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Часть 1 3.1. The main forms of animal behavior
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Comparative Psychology and Zoopsychology
Terms: Comparative Psychology and Zoopsychology