Development of reflexes in aquarium fish. III

III. Examples of motor reflexes.

1. Muscular reflexes of stretching and inhibition.

Consider the muscle stretch reflex. It is designed to regulate the position of the limbs, to ensure the immobility of the body, to support the body while it is standing, lying or sitting. This reflex maintains the constancy of muscle length. Stretching the muscle causes the activation of muscle spindles and contraction, i.e. shortening of the muscle, which counteracts its stretching. For example, when a person is sitting, the abdominal muscles are stretched and their tone is increased, counteracting the flexion of the back. Conversely, too much muscle contraction weakens the stimulation of its stretch receptors, muscle tone weakens.

Consider the passage of a nerve impulse along a reflex arc. It should be immediately noted that the muscle stretch reflex refers to the simplest reflexes. It passes directly from the sensory neuron to the motor neuron (Fig. 1). The signal (irritation) comes from the muscle to the receptor. Through the dendrites of the sensory neuron, the impulse passes to the spinal cord and there the shortest way passes into the motor neuron of the somatic nervous system, and then the impulse enters the effector (muscle) along the axon of the motor neuron. Thus, the muscle stretch reflex is carried out.

Fig.1. 1 - muscle; 2 - muscle receptors; 3 - sensory neuron; 4 - motor neuron; 5 - effector.

Another example of a motor reflex is the inhibition reflex. It occurs as a response to the stretch reflex. The inhibitory reflex arc includes two central synapses: excitatory and inhibitory. We can say that in this case we observe the work of antagonist muscles in a pair, for example, the flexor and extensor in the joint. The motor neurons of one muscle are inhibited during the activation of the other component of the pair. Consider knee flexion. At the same time, we observe stretching of the extensor muscle spindles, which increases the excitation of motor neurons and inhibition of the flexor motor neurons. In addition, a decrease in stretching of the flexor muscle spindles weakens the excitation of homonymous motor neurons and reciprocal inhibition of extensor motor neurons (disinhibition). By homonymous motor neurons we mean all those neurons that send axons to the same muscle or excite the muscle from which the corresponding path from the periphery to the nerve center originates. And reciprocal inhibition is a process in the nervous system, based on the fact that the same afferent pathway excites some groups of cells and inhibits other groups of cells through intercalated neurons. Ultimately, extensor motor neurons fire and flexor motor neurons contract. Thus, there is a regulation of the length of the muscle.

Consider the passage of a nerve impulse along a reflex arc. The nerve impulse originates on the extensor muscle and travels along the axons of the sensory neuron to the spinal cord. Since this reflex arc belongs to the disynaptic type, the impulse bifurcates, one part hits the extensor motor neuron to maintain the length of the muscle, and the other part hits the flexor motor neuron, and the extensor is inhibited. Then each part of the nerve impulse passes to the corresponding effector. Or, in the spinal cord, it is possible to switch to the motoneuron of the knee flexors through inhibitory synapses, which allow you to change the length of the muscle, and then through the motor axons to the end plates (effector, skeletal muscle). Two other options are possible, when the excitation perceives the flexor receptor, then the reflex passes along the same path.

OFig.2 1. The extensor muscle. 2. Flexor muscle. 3. Muscle receptor. 4. Sensory neurons. 5. Inhibitory interneurons. 6. Motor neuron. 7. Effector

Let's get acquainted now with more complex reflexes.

2. Flexion and cross extensor reflex.

As a rule, reflex arcs include two or more serially connected neurons, i.e., they are polysynaptic.

An example is the protective reflex in humans. When exposed to a limb, it is withdrawn by bending, for example, in knee joint. The receptors for this reflex arc are located in the skin. They provide movement aimed at removing the limb from the source of irritation.

When the limb is irritated, a flexion reflex occurs, the limb is withdrawn, and the opposite limb is straightened. This happens as a result of the passage of an impulse along a reflex arc. We act on the right leg. From the receptor of the right leg, along the axons of the sensory neuron, the impulse enters the spinal cord, then it is sent to four different interneuron circuits. Two circuits go to the motor neurons of the flexor and extensor of the right leg. The flexor muscle contracts, and the extensor muscle relaxes under the influence of inhibitory interneurons. We pull back our foot. In the left leg, the flexor muscle relaxes and the extensor muscle contracts under the influence of the excitatory interneuron.

RiceBlack - inhibitory interneurons; red stimulants. 2. Motor neurons. 3. Effectors of relaxed flexor and extensor muscles. 4. Effectors of contracted flexor and extensor muscles.

3. Tendon reflex.

Tendon reflexes serve to maintain the constancy of muscle tension. Each muscle has two regulatory systems: length regulation, with the help of muscle spindles as receptors, and tension regulation, tendon organs act as receptors in this regulation. The difference between the tension regulation system and the length regulation system, in which the muscle and its antagonist are involved, is the use of the muscle tone of the entire limb by the tendon reflex.

The strength developed by the muscle depends on its preliminary stretching, speed of contraction, and fatigue. Deviation from muscle tension from the desired value is recorded by the tendon organs and corrected by the tendon reflex.

The receptor (tendon) of this reflex is located in the tendon of the limb at the end of the flexor or extensor muscle. From there, along the axons of the sensory neuron, the signal passes to the spinal cord. There, the signal can travel down an inhibitory interneuron to an extensor motor neuron, which will send a signal to the extensor muscle to keep the muscle tensed. Also, the signal can go to the excitatory interneuron, which will send a signal through the motor axon to the flexor effector to change muscle tension and perform a certain action. In the case when the excitation perceives the receptor (tendon) of the flexor, the signal passes through the axon of the sensory neuron to the interneuron, and from there, to the motor motor neuron, which sends a signal along the axons of the motor neuron to the flexor muscle. In the reflex arc of the flexor, the path is possible only through the inhibitory interneuron.

Fig. Tendon receptor. 2. Sensory neuron. 3. Inhibitory interneuron. 4. Excitatory interneuron. 5. Motor neuron. 6. Receptor.

In the Black Sea, as probably in others warm seas, there is an amazing way of amateur fishing "for tyrant". A fisherman, accustomed to cautious and capricious freshwater fish, is taken aback when he first gets on sea fishing. The tackle, in other words, the “tyrant” itself, is a long fishing line, to one end of which four or five hooks are attached to short leashes. Nothing else is required - no rod, no bait. The fisherman goes to deep place, lowers the hooks into the water, and wraps the other end of the fishing line around his finger. He sits in the boat and from time to time tugs at the line until he feels that it is getting heavier. Then drags. And what do you think, pulls out a fish, and sometimes not one, but two or three at once. True, a fish, as a rule, does not take empty hooks in its mouth, but hooks on them with its belly, gills, even tail. And still it seems that you need to be completely stupid to fall for such a frankly dangerous tackle, and even not promising any benefits.

Maybe, indeed, fish are very stupid creatures. Let's try to figure it out. Main criterion mind is the ability to learn. Pisces are diligent students. They easily develop different skills. Everyone can verify this for himself. At home, many keep tropical fish. In two or three days, it is easy to teach the inhabitants of the aquarium to swim up to the glass, if you first tap it lightly with your finger, and then throw some tasty food there. After fifteen or twenty such procedures, the fish, having heard the call, will drop all their fish business and rush to the appointed place, hoping to get a portion of worms for diligence.

The skills acquired by bees, ants and fish are not similar to those developed in quite primitive animals. In their complexity, in the duration of their retention, they rarely differ from habituation reactions and from summation reflexes. The high perfection of the nervous system of these animals allowed them to develop adaptive reactions of a new type. They are called conditioned reflexes.

This type of reflexes was discovered and studied by I.P. Pavlov on dogs. The name is not given by chance. The formation, preservation or elimination of these reflexes occurs only under special conditions.

For the emergence conditioned reflexes it is necessary that the action of two specific stimuli coincide several times in time. One of them - it is necessary that he act first - should not be of any particular importance to the animal, neither to frighten him, nor to cause him a food reaction. Otherwise, it is absolutely indifferent what kind of irritant it will be. It may be some sound, the sight of any object or other visual stimulus, any smell, heat or cold, touching the skin, and so on.

The second stimulus, on the contrary, should cause some kind of innate reaction, some kind of unconditioned reflex. This may be a food or defensive reaction. After several combinations of such stimuli, the first of them, previously a completely indifferent stimulus for the animal, begins to evoke the same reaction as the unconditioned one. It was in this way that I developed a food conditioned reflex in the inhabitants of my aquarium. The first stimulus, tapping on glass, was at first absolutely indifferent to the fish. But after it coincided fifteen to twenty times with the action of a food irritant - ordinary fish food - the tapping acquired the ability to cause a food reaction, forcing the fish to rush to the feeding place. Such a stimulus is called a conditioned stimulus.

Even in ants and fish, conditioned reflexes persist for a very long time, and in higher animals - almost all their lives. And if at least occasionally the training of the conditioned reflex is carried out, it is able to serve the fish indefinitely. However, when the conditions that led to the formation of the conditioned reflex change, if the action of the conditioned stimulus no longer follows the unconditioned stimulus, the reflex is destroyed.

In fish, conditioned reflexes are easily formed even without our help. My fish immediately swim out of all corners as soon as I find myself near the aquarium, although no one specially accustomed them to this. They firmly know that I will not approach them empty-handed. Another thing is if the aquarium is crowded with children. Kids like to knock on glass more, scare the inhabitants of the aquarium, and the fish hide in advance. This is also a conditioned reflex, only the reflex is not food, but defensive.

There are many types of conditioned reflexes. Their names emphasize some one feature of the reaction, developed in such a way that everyone immediately understands what is at stake. Most often, the name is given in accordance with the reaction that the animal performs. A food conditioned reflex, when a fish swims up to a feeding place, and if it hurries to hide in the thick of underwater plants, they say that it has formed a defensive conditioned reflex.

When studying the mental abilities of fish, they often resort to the development of both food and defensive conditioned reflexes. Usually, the subjects come up with a task a little more difficult than the ability to quickly arrive at the feeding place or hastily escape. The scientists of our country love to make fish grab a bead with their mouths. If you lower a small red ball tied to a thin thread into the water, it will definitely interest the fish. In general, red color attracts them. The fish will certainly grab the ball with its mouth in order to taste it, and, pulling the thread, will try to take it away with it, in order to calmly figure out somewhere on the sidelines whether this thing is edible or not. A conditioned reflex is developed to light or to a call. While the fish swims up to the bead, the light is on, and as soon as the bead is in the mouth of the fish, they throw a worm to it. One or two procedures are enough for the fish to continuously grab the bead, but if the development of the reflex is continued, it will eventually notice that the worm is being given as long as the light is on. Now, as soon as the light comes on, the fish will hastily rush to the bead, and the rest of the time it will not pay any attention to it. She remembered the connection between light, bead and worm, which means that she developed a food reflex to light.

Pisces are capable of solving more complex problems. Three beads are immediately lowered into the aquarium to the minnow, and on the outside, a simple picture is attached to the glass opposite each of them, for example, a black triangle, the same square and circle. The minnow, of course, will immediately become interested in the beads, and the experimenter is closely watching his actions. If they are going to develop a conditioned reflex to a circle, then as soon as the fish swims up to this picture and grabs the bead hanging opposite it, they throw a worm at it. Pictures during the experiment are constantly swapped, and soon the minnow will understand that the worm can only be obtained by pulling the bead hanging against the circle. Now he will not be interested in other pictures and other beads. He developed a food conditioned reflex to the image of a circle. This experience convinced scientists that fish are able to distinguish pictures and remember them well.

To develop a defensive conditioned reflex, the aquarium is divided into two parts by a partition. A hole is left in the partition so that the fish can move from one part to another. Sometimes a door is hung on a hole in the partition, which the fish can easily open by pushing with its nose.

The development of the reflex is carried out according to the usual scheme. A conditioned stimulus is turned on, for example, a bell, and then for a moment they turn on the electric current and continue to spur the fish on with the current until it guesses to open the door in the partition and go to another part of the aquarium. After several repetitions of this procedure, the fish will understand that soon after the start of the sound of the call, very unpleasant and painful effects await it, and, not waiting for them to begin, hastily swims away behind the partition. Conditioned defensive reflexes are often developed faster and last much longer than food ones.

In this chapter, we met with animals that have well-developed conditioned reflexes. In my own way mental development animals are about the same. True, some of them, namely social insects, are the highest representatives of their branch of the animal kingdom, the highest link in the development of arthropods. There are no smarter arthropods than bees, wasps, ants and termites. Another thing is the fish. They stand at the very first steps in the development of their branch - vertebrates. Among them, they are the most primitive, underdeveloped creatures.

Both ants and fish are able to learn, they are able to notice the patterns of the world around them. Their training, acquaintance with various natural phenomena proceeds through the formation of simple conditioned reflexes. For them, this is the only way to know the world.

All accumulated knowledge is stored in their brain in the form of visual, sound, olfactory and gustatory images, that is, as if duplicates (or copies) of those impressions that were formed at the time of perception of the corresponding stimuli. The light above the aquarium lit up - and revived in the animal's brain the image of a bead, the image of its own motor reactions, the image of a worm. Obeying this chain of images, the fish swims up to the bead, grabs it and waits for the due reward.

The peculiarity of the knowledge acquired by animals due to the formation of simple conditioned reflexes is that they can notice only those patterns of the surrounding world that are of direct importance to them. The minnow will certainly remember that after a flash of light, under certain conditions, delicious food may appear, and after the sound of the bell, you will feel pain if you do not immediately clean up to another room. My pet fish don't care what I wear when I go to their tank, as it doesn't involve any special benefit or trouble, and they don't care about my clothes. But my dog ​​instantly perks up as soon as I go to the hanger and take the coat. She has long noticed that I go out into the street in a coat, and every time she hopes that they will take her for a walk.

Conditioned reflexes are easily formed and persist for a long time, even if they are not trained, but they can just as easily be destroyed, destroyed. And this is not a defect, but a great advantage of conditioned reflexes. Due to the fact that it is possible to make changes in the developed reflexes and even destroy them, the knowledge acquired by the animal is constantly refined and improved. After the flash of light, the experimenters stopped throwing worms into the aquarium, you see, after a few days the crucian stopped grabbing the bead. The reaction became useless, no reward was given for it, and the conditioned reflex, as scientists say, faded away. They stopped giving the minnow a worm when he pulls a bead hanging against the circle, and the conditioned reflex will soon fade away. They began to give food when he grabs a bead hanging against a square, and a new conditioned reflex is developed in the fish.

From early childhood to very old age, the animal can form more and more conditioned reflexes, and those that have become unnecessary are extinguished. Thanks to this, knowledge is constantly accumulated, refined and polished. They are very necessary for animals, helping to find food, escape from enemies, in general, to survive.

HIGH NERVOUS ACTIVITY OF LARVAR-CHORD CYCLOSTOMES AND FISH

The higher nervous activity of vertebrates reflects one of the important trends in their evolution - individual perfection. This trend is manifested in increasing life expectancy, a reduction in the number of offspring, an increase in body size, and an increase in the conservatism of heredity. An expression of the same tendency is that, on the basis of a limited number of species instincts, each individual, in the order of personal life experience can form a greater number of diverse conditioned reflexes.

In such lower chordates as larval-chordates and cyclostomes, conditioned reflexes are of a primitive nature. With the development of the analytical-synthetic activity of the brain and the use of more and more subtle signals in fish, conditioned reflexes begin to play an increasingly significant role in their behavior.

Conditioned reflexes of larval-chordates

Despite the regression of its nervous system, the ascidia can form a conditioned protective reflex of closing the siphons to a sound, or rather, a vibration-mechanical signal.

To develop such a reflex, a dropper was installed over the ascidian sitting in the aquarium. With each impact of a drop on the surface of the water, the ascidia quickly closed the siphons, and with stronger irritation (drop falling from a great height), it pulled them in. The source of the conditioned signals was an electric bell mounted on a table next to the aquarium. Its isolated action lasted 5 s, at the end of which a drop fell. After 20–30 combinations, the bell itself could already evoke defensive movements of the siphons.

Removal of the central ganglion destroyed the developed reflex and made it impossible for the formation of new ones. Persistent attempts to develop similar conditioned reflexes to light in healthy animals were unsuccessful. Obviously, the absence of reactions to light signals is explained by the living conditions of ascidians.

In these experiments it was also found that as a result of combinations of a signal with an unconditioned reaction, the latter was more and more easily evoked by the unconditioned stimulus. It is possible that such a conditional increase in the excitability of the signaled reaction is the initial summation form of a temporary connection, from which more specialized ones developed later.

cyclostomes

The sea lamprey reaches a meter in length. The sexual instinct every spring makes her, like many marine fish, leave the depths of the sea and rise into the rivers for spawning. However, inhibition can be developed for this instinctive reaction (lampreys stopped entering rivers where they encountered polluted water).

The conditioned reflexes of the river lamprey were studied during reinforcement with electric shocks. A light signal (2 lamps of 100 W), to which, after 5–10 s of isolated action, a 1–2 second unconditioned electrocutaneous stimulation was added, after 3–4 combinations, it itself began to cause a motor defensive reaction. However, after 4–5 repetitions, the conditioned reflex decreased and soon disappeared. After 2–3 hours, it could be developed again. It is noteworthy that, simultaneously with a decrease in the conditioned defensive reflex, the value of the unconditioned one also decreased. The threshold of electrocutaneous stimulation to evoke a defensive reaction was increased in this case. It is possible that such changes depended on the traumatic nature of the electrical stimulation.

As was shown above with the example of ascidians, the formation of a conditioned reflex can manifest itself in an increase in the excitability of the signaled reaction. In this case, using the lamprey as an example, it can be seen how, when the conditioned reflex is inhibited, the causative agent of the signaled reaction decreases. Easily forming a conditioned defensive reflex to the light of a lamp, lampreys were unable to develop it to the sound of a bell. Despite 30–70 combinations of the bell with electric shocks, it never became a signal for defensive movements. This indicates a predominantly visual orientation of lampreys in the environment.

Lamprey perceives light stimuli not only with the help of the eyes. Even after transection of the optic nerves or complete removal of the eyes, the reaction to light persisted. It disappeared only when, in addition to the eye, the parietal organ of the brain, which has light-sensitive cells, was also removed. Some nerve cells of the diencephalon and cells located in the skin near the anal fin also have a photoreceptor function.

Having achieved high perfection in adaptation to an aquatic lifestyle, fish have significantly expanded their receptor capabilities, in particular, due to the mechanoreceptors of the lateral line organs. Conditioned reflexes constitute an essential part of the behavior of cartilaginous and especially bony fish.

Cartilaginous fish. The shark's gluttony is proverbial for a reason. Its powerful food instinct is difficult to slow down even with strong pain stimuli. Thus, whalers claim that the shark continues to tear and swallow pieces of the meat of a dead whale, even if you stick a spear into it. On the basis of such pronounced unconditioned food reactions in sharks in a natural setting, many conditioned food reflexes are apparently formed. This, in particular, is evidenced by the descriptions of how quickly sharks develop a response to escort ships and even swim at a certain time to the board from which kitchen waste is thrown.

Sharks use olfactory food signals very actively. They have been known to follow wounded prey on a trail of blood. The importance of smell for the formation of food reflexes was shown in experiments on small Mustelus laevis, floating freely in the pond. These sharks found live hidden crabs in 10–15 min, and dead and opened crabs in 2–5 min. If the sharks had their nostrils covered with Vaseline cotton, they could not find the hidden crab.

Properties of the formation of conditioned defensive reflexes in Black Sea sharks (Squalus acanthias) studied using the technique described above for lampreys. It turned out that sharks developed a conditioned reflex to a bell after 5–8 combinations, and to a lamp only after 8–12 combinations. The developed reflexes were very unstable. They did not last for 24 hours, and the next day they had to be worked out again, although this required fewer combinations than on the first day.

Similar properties of the formation of conditioned defensive reflexes were also found by other representatives of cartilaginous fish - rays. These properties reflect the conditions of their life. Yes, inhabitant sea ​​depths the spiny stingray needed 28–30 combinations to develop a reflex to a call, while a mobile inhabitant coastal waters 4–5 combinations were enough for the stingray. In these conditioned reflexes, the fragility of temporary connections also manifested itself. The conditioned reflex developed the day before disappeared the next day. It had to be restored each time with two or three combinations.

Bony fish. Due to the enormous diversity in body structure and behavior, bony fish achieved excellent adaptability to a wide variety of habitat conditions. The little one belongs to these fishes. Mistichthus luzonensis(the smallest vertebrate, 12-14 mm in size), and a giant "herring king" (Regalecus) the southern seas, reaching 7 m in length.

The instincts of fish, especially food and sex, are extremely diverse and specialized. Some fish, such as the vegetarian crucian carp, swim peacefully in muddy ponds, while others, such as the carnivorous pike, live by hunting. Although most fish leave fertilized eggs to their fate, some of them show concern for offspring. For example, blennies guard the laid eggs until the juveniles hatch. The nine-spined stickleback builds a real nest of blades of grass, sticking them together with its mucous secretions. Having completed the construction, the male drives the female into the nest and does not release it until she spawns. After that, he waters the eggs with seminal fluid and guards at the entrance to the nest, from time to time ventilating it with special movements of the pectoral fins.

Freshwater fish from the family cichlidae in case of danger, they hide the hatched juveniles in their mouths. They describe the special "calling" movements of adult fish, with which they collect their fry. Pinagora leads fry, which can be attached to the father's body with special suckers.

Seasonal migrations are a striking manifestation of the strength of the sexual instinct of fish. For example, salmon at certain times of the year rush from the sea to rivers to spawn. Animals and birds exterminate them in masses, many fish die from exhaustion, but the rest stubbornly continue on their way. In an irresistible rush to the upper reaches of the river, the noble salmon, meeting an obstacle, jumps on stones, breaks into blood and again rushes forward until it overcomes it. He jumps rapids and climbs waterfalls. Protective and food instincts are completely inhibited, everything is subordinated to the task of reproduction.

The relationship of fish in a flock reveals a certain hierarchy of subordination to the leader, which can take various forms. Thus, observations are made of a flock of Malabar zebrafish, where the leader swims almost horizontally, which allows him to be the first to see and grab an insect that has fallen to the surface of the water. The rest of the fish are distributed according to ranks and swim with an inclination of 20 to 45 °. A large role in the behavior of fish is played by the pheromones they secrete. For example, when the minnow's skin is damaged, toribons, chemical alarm signals, enter the water. It was enough to drop such water into an aquarium with minnows so that they rushed to flight.

Conditioned reflexes to sound stimuli. Aquarium hobbyists know well how to train fish to gather at the surface of the water at the signal of tapping on the wall, if you practice this tapping before each feeding. Apparently, such a conditioned food reflex determined the behavior of the famous fish of the monastery pond in Krems (Austria), which attracted the attention of tourists by the fact that they swam to the shore at the sound of a bell. Researchers who deny hearing in fish claim that the fish swam only when they saw a person coming to the pond or when his steps caused the ground to shake. However, this does not exclude the participation of sound as one of the parts of the complex stimulus.

The question of the hearing of fish has long been controversial, especially since the fish has neither a cochlea nor the main membrane of the organ of Corti. It was resolved positively only by the objective method of conditioned reflexes (Yu. Frolov, 1925).

The experiments were carried out on freshwater (crucian carp, ruff) and marine (cod, haddock, goby) fish. In a small aquarium, the test fish swam on a leash tied to an air transfer capsule. The same thread was used to bring electric current to the body of the fish, the second pole was a metal plate lying on the bottom. The source of the sound was a telephone receiver. After 30–40 combinations of sounds with electric shocks, an auditory conditioned protective reflex was formed. When the phone was turned on, the fish dived without expecting an electric shock.

In this way, it was possible to develop conditioned reflexes also to various kinds of water vibrations and other signals, such as light.

The defensive reflexes developed on reinforcement with electric current turned out to be very strong. They persisted for a long time and were difficult to extinguish. At the same time, it was not possible to develop reflexes for traces of signals. If the beginning of the unconditioned reinforcement lagged behind the end of the action of the conditioned signal by at least 1 s, the reflex did not form. It was also found that the development of one conditioned reflex facilitated the formation of subsequent ones. Based on the results of these experiments, one can judge a certain inertia and weakness of temporary connections, which, however, are capable of training.

It is not difficult to develop a conditioned food reflex to sound in a golden orphan fish, accompanying the sound signal by lowering a bag of chopped worms into the aquarium. At the fish Umbra limi not only was a similar conditioned positive reflex to a tone of 288 oscillations/s formed, but also a differentiation of the tone of 426 oscillations/s was developed, which was accompanied by the supply of a lump of filter paper moistened with camphor alcohol instead of food.

In order to completely exclude the participation of vision, sound conditioned reflexes were developed on previously blinded pygmy catfish, minnows and loaches. In this way, the upper limit of the audibility of sounds was established, which turned out to be about 12,000 oscillations / s for the catfish, about 6000 for the minnow, and about 2500 for the char. When determining the lower limit of the audibility of sounds, it turned out that fish perceive very slow (2–5 vibrations / s) and even single vibrations of water, which for the human ear are not sounds. These slow fluctuations can be made conditioned stimuli of the food reflex and their differentiation can be worked out. Transection of the nerves of the lateral line organ destroys reflexes to low sounds, the lower limit of audibility rises to 25 Hz. Consequently, the lateral line organ is a kind of infrasonic hearing organ in fish.

Per recent times accumulated information about the sounds emitted by fish. It has long been known that Malay fishermen dive into the water to learn by ear where a school of fish is. The "voices" of the fish are recorded on a tape recorder. They turned out to be different various kinds fish, higher in fry and lower in adults. Among our Black Sea fish, the croaker turned out to be the most "vociferous". It is noteworthy that in the croaker, the conditioned reflex to the sound is formed after 3–5 combinations, i.e. faster than other studied fish, such as crucian carp, which required 9–15 combinations. However, the croaker develops conditioned reflexes to light signals worse (after 6–18 combinations).

Conditioned reflexes to light stimuli. A variety of conditioned reflexes to food reinforcement were developed during training of fish in order to study their vision. Thus, in experiments with minnows, it was established that they differentiate light stimuli well in terms of brightness, distinguishing between different shades of gray, it was also possible to distinguish between hatched figures by fish, and vertical hatching acquired a signal value faster than horizontal. Experiments with perches, minnows and minnows have shown that fish can differentiate according to the shape of such figures as a triangle and a square, a circle and an oval. It also turned out that visual contrasts are characteristic of fish, reflecting induction phenomena in the brain parts of the analyzers.

If you feed macropods with red chironomid larvae, then soon the fish attacked the aquarium wall when lumps of red wool, similar in size to the larvae, were glued to the glass outside. The micropods did not respond to green and white lumps of the same size. If you feed the fish with spools of white bread crumb, then they begin to grab the white woolen lumps that are in sight.

They describe that once a coral predator was given a red-painted satin along with a jellyfish tentacle. The predatory fish at first grabbed the prey, but, having burned themselves on the stinging capsules, immediately released it. After that, she did not take red fish for 20 days.

Especially a lot of research has been carried out on the study of the properties of vision of carps. Thus, in experiments on the development of defensive conditioned reflexes to the presentation of lines as signals, it was shown that fish could differentiate them by the angle of inclination. Based on these and other experiments, suggestions were made about a possible mechanism of visual analysis in fish using detector neurons. O high development visual perception carp is evidenced by its ability to distinguish the paint of an object even in different lighting conditions. This property of perceptual constancy manifested itself in the carp in relation to the shape of the object, the reaction to which remained definite, despite its spatial transformations.

Conditioned olfactory, gustatory and temperature reflexes. Fish can develop olfactory and gustatory conditioned reflexes. After being fed musk-scented meat for some time, the minnow began to respond with a typical search response to a previously indifferent musky smell. The smell of skatole or coumarin could be used as an olfactory signal. The signal odor was differentiated from those not reinforced by feeding. Very easily becomes a positive signal for minnows the smell of mucus covering their body. It is possible that such a natural reflex explains some of the properties of the gregarious behavior of these fish.

If earthworms fed to minnows are pre-soaked in a sugar solution, then after 12–14 days the fish will pounce on cotton wool with a sugar solution lowered into the aquarium. Other sweet substances, including saccharin and glycerin, evoked the same reaction. You can develop taste conditioned reflexes to bitter, salty, sour. The threshold of irritation for minnow turned out to be higher for bitter, and lower for sweet than in humans. These reflexes did not depend on olfactory signals, since they persisted even after the removal of the olfactory lobes of the brain.

Observations are described that show that the development of chemoreceptors in fish is associated with the search for and discovery of food. Carps can develop instrumental conditioned reflexes to regulate salinity or acidity of water. In this case, the motor reaction led to the addition of solutions of a given concentration. At the fish Poecilia reticulata Peters developed conditioned food reflexes to the taste of beta-phenylethanol with differentiation to coumarin.

Convincing evidence has been obtained that salmonids, approaching the mouth of the river where they were born, use their sense of smell to find their "native" spawning ground. The high selective sensitivity of their chemoreception is indicated by the results of an electrophysiological experiment in which impulses were recorded in the olfactory bulb only when water from the “native” spawning ground was passed through the nostrils of the fish, and were absent if the water was from the “alien” one. It is known to use trout as a test object for assessing the purity of water after treatment facilities.

You can make the temperature of the water in which the fish swims a conditional food signal. At the same time, it was possible to achieve differentiation of temperature stimuli with an accuracy of 0.4 °C. There are reasons to believe that natural temperature signals play big role in the sexual behavior of fish, in particular in spawning migrations.

Complex food-procuring reflexes. For best comparison indicators of conditioned reflex activity different types Animals use natural food-procuring movements. Such a movement for fish is the grasping of a bead suspended on a string. The first random grasps are reinforced with food and combined with an auditory or visual signal, to which a conditioned reflex is formed. Such a conditioned visual reflex, for example, was formed and strengthened in crucian carp in 30–40 combinations. Differentiation by color and a conditional brake were also developed. However, repeated modifications of the signal value of positive and negative stimuli proved to be an extremely difficult task for fish and even led to disturbances in conditioned reflex activity.

Studies of the behavior of fish in labyrinths have shown their ability to develop a reaction of unmistakably choosing the right path.

Yes, dark-loving fish Tundulus after 12–16 trials for two days, she began to swim through the openings of the screens, without going into dead ends, right into the corner where food was waiting. In similar experiments with goldfish, the time for searching for a way out of the maze for 36 trials decreased from 105 to 5 minutes. After a 2-week break in work, the acquired skill has changed only slightly. However, with more complex mazes, such as those used for rats, the fish could not cope, despite hundreds of trials.

Predatory fish can develop a conditioned reflex suppression of the hunting instinct.

If you place crucian carp behind a glass partition in an aquarium with a pike, then the pike will immediately rush at him. However, after several headbutts on the glass, the attacks stop. After a few days, the pike no longer tries to grab the crucian. The natural food reflex is completely extinguished. Then the partition is removed, and crucian carp can swim next to the pike. A similar experiment was carried out with predatory perches and minnows. Predators and their usual victims lived peacefully together.

Another example of a conditioned reflex transformation of instinctive behavior was shown by an experiment with cichlid fish, which, during their first spawning, had their eggs replaced with caviar of an alien species. When the fry hatched, the fish began to take care of them and protect them, and when they brought fry of their own species to the next spawning, they drove them as strangers. Thus, the developed conditioned reflexes turned out to be very conservative. On the basis of reinforcement with food and defensive reactions, various motor conditioned reflexes were developed in fish. For example, a goldfish was taught to swim through a ring, to make “dead loops”, a brilliant betta fighting fish, accustomed to pass through a hole in a barrier, began to jump into it even when it was raised above the water.

The behavior of fish, their unconditioned and conditioned reflexes are largely determined by environmental factors habitat, which leaves its mark on the development of the nervous system and the formation of its properties.

Development of defensive conditioned reflexes in fry. The regulation of the flow of rivers, the construction of hydroelectric dams and reclamation systems, to a greater or lesser extent, makes it difficult for fish to reach natural spawning grounds. Therefore, more and more economic importance acquires artificial fish farming.

Every year, billions of fry hatched at hatcheries are released into lakes, rivers and seas. But only a small part of them survive to commercial age. Grown in artificial conditions, they often turn out to be poorly adapted to life in the wild. In particular, fry that have not had life experience in the formation of protective reactions easily become the prey of predatory fish, from which they do not even try to escape. In order to increase the survival rate of fry released by fish breeding stations, experiments were undertaken to artificially develop in them protective conditioned reflexes to the approach of predatory fish.

In preliminary tests, the properties of the formation of such reflexes to visual, auditory and vibrational signals were studied. If, among the roach fry, shiny metal plates shaped like the body of a bee-eater are placed, and a current is passed through these plates, then the fry begin to avoid these figures even in the absence of a current. The reflex is developed very quickly (Fig. 84).

Rice. 84. Development of a conditioned defensive reflex in roach fry to the appearance of a predatory fish model for 1 hour (according to G.V. Popov):

1 - 35 day fry, 2 - 55 days

To assess how much the development of artificial defensive reflexes can increase the survival rate of juveniles, we compared the rate at which a predator eats fry that have undergone training and fry that have not had such training.

For this, cages were installed in the pond. One predatory fish was placed in each cage - a chub and a precisely counted number of fish fry. After 1 or 2 days, we counted how many fry remained alive and how many were eaten by the predator. It turned out that of the fry that did not develop defensive reflexes, almost half perished during the first day. It is noteworthy that the second day adds little in this regard. It can be assumed that the surviving fry have time to form natural conditioned defensive reflexes and successfully escape from the persecution of the predator. Indeed, if they are taken after such a natural preparation in special experiments, then the percentage of death turns out to be either relatively small, or even zero.

Fry with artificially developed conditioned defensive reflexes both to the appearance of the figure of a predatory fish and to the shaking of the water, imitating its movements, suffered the least from the chub. In most of the experiments, the predator, even within two days, was unable to catch a single one of them.

A recently developed simple technique for educating protective reflexes in fry commercial fish during their cultivation can bring significant practical benefits to the business of fish breeding.