The visual analyzer is a set of structures that perceive light energy in the form of electromagnetic radiation with a wavelength of 400-700 nm and discrete particles of photons, or quanta, and form visual sensations. With the help of the eye, 80 - 90% of all information about the world around us is perceived.

Rice. 2.1

Thanks to the activity of the visual analyzer, the illumination of objects, their color, shape, size, direction of movement, the distance at which they are removed from the eye and from each other are distinguished. All this allows you to evaluate the space, navigate the world around you, and perform various types of purposeful activities.

Along with the concept of a visual analyzer, there is the concept of an organ of vision (Fig. 2.1)

This is an eye that includes three functionally different elements:

1) the eyeball, in which the light-perceiving, light-refracting and light-regulating apparatuses are located;

2) protective devices, i.e. outer shells of the eye (sclera and cornea), lacrimal apparatus, eyelids, eyelashes, eyebrows; 3) the motor apparatus, represented by three pairs of eye muscles (external and internal rectus, superior and inferior rectus, superior and inferior oblique), which are innervated by III (oculomotor nerve), IV (trochlear nerve) and VI (abducens nerve) pairs of cranial nerves.

Structural and functional characteristics

Receptor (peripheral) department The visual analyzer (photoreceptors) is subdivided into rod and cone neurosensory cells, the outer segments of which are, respectively, rod-shaped ("rods") and cone-shaped ("cones") forms. A person has 6-7 million cones and 110-125 million rods.

The exit point of the optic nerve from the retina does not contain photoreceptors and is called the blind spot. Lateral to the blind spot in the region of the fovea lies the area of ​​​​best vision - the yellow spot, containing mainly cones. Towards the periphery of the retina, the number of cones decreases, and the number of rods increases, and the periphery of the retina contains only rods.

Differences in the functions of cones and rods underlie the phenomenon of dual vision. Rods are receptors that perceive light rays in low light conditions, i.e. colorless or achromatic vision. Cones, on the other hand, function in bright light conditions and are characterized by different sensitivity to the spectral properties of light (color or chromatic vision). Photoreceptors have a very high sensitivity, which is due to the peculiarity of the structure of the receptors and the physicochemical processes that underlie the perception of light stimulus energy. It is believed that photoreceptors are excited by the action of 1-2 light quanta on them.

Rods and cones consist of two segments - outer and inner, which are interconnected by means of a narrow cilium. The rods and cones are oriented radially in the retina, and the molecules of photosensitive proteins are located in the outer segments in such a way that about 90% of their photosensitive groups lie in the plane of the disks that make up the outer segments. Light has the greatest exciting effect if the direction of the beam coincides with the long axis of the rod or cone, while it is directed perpendicular to the discs of their outer segments.

Photochemical processes in the retina. In the receptor cells of the retina there are light-sensitive pigments (complex protein substances) - chromoproteins, which become discolored in the light. The rods on the membrane of the outer segments contain rhodopsin, the cones contain iodopsin and other pigments.

Rhodopsin and iodopsin consist of retinal (vitamin A 1 aldehyde) and glycoprotein (opsin). Having similarities in photochemical processes, they differ in that the absorption maximum is located in different regions of the spectrum. Rods containing rhodopsin have an absorption maximum in the region of 500 nm. Among the cones, three types are distinguished, which differ in the maxima in the absorption spectra: some have a maximum in the blue part of the spectrum (430-470 nm), others in the green (500-530), and others in the red (620-760 nm) part, which is due to the presence of three types of visual pigments. The red cone pigment is called iodopsin. Retinal can be in various spatial configurations (isomeric forms), but only one of them, the 11-CIS isomer of retinal, acts as the chromophore group of all known visual pigments. The source of retinal in the body are carotenoids.

Photochemical processes in the retina proceed very economically. Even under the action of bright light, only a small part of the rhodopsin present in the sticks (about 0.006%) is cleaved.

In the dark, resynthesis of pigments takes place, proceeding with the absorption of energy. The recovery of iodopsin proceeds 530 times faster than that of rhodopsin. If the content of vitamin A in the body decreases, then the processes of resynthesis of rhodopsin weaken, which leads to impaired twilight vision, the so-called night blindness. With constant and uniform illumination, a balance is established between the rate of disintegration and resynthesis of pigments. When the amount of light falling on the retina decreases, this dynamic balance is disturbed and shifted towards higher pigment concentrations. This photochemical phenomenon underlies dark adaptation.

Of particular importance in photochemical processes is the pigment layer of the retina, which is formed by an epithelium containing fuscin. This pigment absorbs light, preventing its reflection and scattering, which determines the clarity of visual perception. The processes of pigment cells surround the light-sensitive segments of rods and cones, taking part in the metabolism of photoreceptors and in the synthesis of visual pigments.

Due to photochemical processes in the photoreceptors of the eye, under the action of light, a receptor potential arises, which is a hyperpolarization of the receptor membrane. This is a distinctive feature of the visual receptors, the activation of other receptors is expressed in the form of depolarization of their membrane. The amplitude of the visual receptor potential increases with increasing intensity of the light stimulus. So, under the action of red, the wavelength of which is 620-760 nm, the receptor potential is more pronounced in the photoreceptors of the central part of the retina, and blue (430-470 nm) - in the peripheral.

The synaptic endings of the photoreceptors converge to the bipolar neurons of the retina. In this case, the photoreceptors of the fovea are associated with only one bipolar.

Conductor department. The first neuron of the conductive section of the visual analyzer is represented by bipolar cells of the retina (Fig. 2.2).

Rice. 2.2

It is believed that action potentials arise in bipolar cells similar to receptor and horizontal HCs. In some bipolars, when the light is turned on and off, a slow long-term depolarization occurs, while in others, when the light is turned on, hyperpolarization occurs, and when the light is turned off, depolarization occurs.

Axons of bipolar cells, in turn, converge to ganglion cells (the second neuron). As a result, about 140 rods and 6 cones can converge for each ganglion cell, while the closer to the macula, the fewer photoreceptors converge per cell. In the area of ​​the macula, there is almost no convergence and the number of cones is almost equal to the number of bipolar and ganglion cells. This explains the high visual acuity in the central parts of the retina.

The retinal periphery is highly sensitive to weak light. This is due, apparently, to the fact that up to 600 rods converge here through bipolar cells to the same ganglion cell. As a result, the signals from many rods are summed up and cause more intense stimulation of these cells.

In ganglion cells, even with complete blackout, a series of impulses with a frequency of 5 per second are spontaneously generated. This impulsation is detected by microelectrode examination of single optic fibers or single ganglion cells, and in the dark it is perceived as "the own light of the eyes."

In some ganglion cells, an increase in background discharges occurs when the light is turned on (on-response), in others, when the light is turned off (off-response). The reaction of the ganglion cell may also be due to the spectral composition of the light.

In the retina, in addition to vertical connections, there are also lateral connections. Lateral interaction of receptors is carried out by horizontal cells. Bipolar and ganglion cells interact with each other due to numerous lateral connections formed by the collaterals of the dendrites and axons of the cells themselves, as well as with the help of amacrine cells.

Horizontal cells of the retina provide regulation of the transmission of impulses between photoreceptors and bipolars, regulation of color perception and adaptation of the eye to different illumination. During the entire period of illumination, horizontal cells generate a positive potential - a slow hyperpolarization, called the S-potential (from the English slow - slow). According to the nature of the perception of light stimuli, horizontal cells are divided into two types:

1) L-type, in which the S-potential occurs under the action of any wave of visible light;

2) C-type, or "color" type, in which the sign of the potential deviation depends on the wavelength. So, red light can cause them to depolarize, and blue light can cause hyperpolarization.

It is believed that the signals of horizontal cells are transmitted in an electrotonic form.

Horizontal as well as amacrine cells are called inhibitory neurons because they provide lateral inhibition between bipolar or ganglion cells.

The set of photoreceptors that send their signals to one ganglion cell forms its receptive field. Near the macula, these fields have a diameter of 7-200 nm, and on the periphery - 400-700 nm, i.e. in the center of the retina, the receptive fields are small, while at the periphery of the retina they are much larger in diameter. The receptive fields of the retina are rounded, built concentrically, each of them has an excitatory center and an inhibitory peripheral zone in the form of a ring. There are receptive fields with on-center (excited when the center is illuminated) and off-center (excited when the center is darkened). The inhibitory rim is currently thought to be formed by horizontal retinal cells by the mechanism of lateral inhibition, i.e. the more excited the center of the receptive field, the greater the inhibitory effect it has on the periphery. Thanks to these types of receptive fields (RP) of ganglion cells (with on- and off-centers), light and dark objects in the field of view are detected already at the level of the retina.

In the presence of color vision in animals, the color-opponent organization of the RP of retinal ganglion cells is isolated. This organization consists in the fact that a certain ganglion cell receives excitatory and inhibitory signals from cones that have different spectral sensitivity. For example, if the "red" cones have an excitatory effect on a given ganglion cell, then the "blue" cones inhibit it. Various combinations of excitatory and inhibitory inputs from different classes of cones have been found. A significant proportion of color-opponent ganglion cells are associated with all three types of cones. Due to this organization of RP, individual ganglion cells become selective for illumination of a certain spectral composition. So, if excitation arises from “red” cones, then excitation of blue- and green-sensitive cones will cause inhibition of these cells, and if a ganglion cell is excited from blue-sensitive cones, then it is inhibited from green- and red-sensitive, etc.

Rice. 2.3

The center and periphery of the receptive field have maximum sensitivity at opposite ends of the spectrum. So, if the center of the receptive field responds with a change in activity to the inclusion of red light, then the periphery responds with a similar reaction to the inclusion of blue. A number of retinal ganglion cells have the so-called directional sensitivity. It manifests itself in the fact that when the stimulus moves in one direction (optimal), the ganglion cell is activated, while in the other direction of movement, there is no reaction. It is assumed that the selectivity of the reactions of these cells to movement in different directions is created by horizontal cells that have elongated processes (teledendrites), with the help of which ganglion cells are inhibited in a direction. Due to convergence and lateral interactions, the receptive fields of adjacent ganglion cells overlap. This makes possible the summation of the effects of light exposure and the emergence of mutual inhibitory relationships in the retina.

Electrical phenomena in the retina. In the retina, where the receptor section of the visual analyzer is localized and the conductive section begins, complex electrochemical processes occur in response to the action of light, which can be recorded in the form of a total response - an electroretinogram (ERG) (Fig. 2.3).

ERG reflects such properties of a light stimulus as color, intensity and duration of its action. ERG can be recorded from the whole eye or directly from the retina. To obtain it, one electrode is placed on the surface of the cornea, and the other is applied to the skin of the face near the eye or on the earlobe.

On the ERG recorded when the eye is illuminated, several characteristic waves are distinguished. The first negative wave a is a small amplitude electrical oscillation reflecting the excitation of photoreceptors and horizontal cells. It quickly turns into a steeply growing positive wave b, which occurs as a result of excitation of bipolar and amacrine cells. After wave b, a slow electropositive wave c is observed - the result of excitation of pigment epithelium cells. With the moment of cessation of light stimulation, the appearance of an electropositive wave d is associated.

ERG indicators are widely used in the clinic of eye diseases to diagnose and control the treatment of various eye diseases associated with retinal damage.

The conduction section, starting in the retina (the first neuron is bipolar, the second neuron is ganglion cells), is anatomically represented further by the optic nerves and, after a partial intersection of their fibers, by the optic tracts. Each optic tract contains nerve fibers coming from the inner (nasal) surface of the retina of the same side and from the outer half of the retina of the other eye. The fibers of the optic tract are sent to the optic tubercle (the thalamus proper), to the metathalamus (external geniculate bodies) and to the pillow nuclei. The third neuron of the visual analyzer is located here. From them, the optic nerve fibers are sent to the cortex of the cerebral hemispheres.

In the outer (or lateral) geniculate bodies, where fibers from the retina come, there are receptive fields that are also rounded, but smaller than those in the retina. The responses of neurons here are phasic in nature, but more pronounced than in the retina.

At the level of the external geniculate bodies, the process of interaction of afferent signals coming from the retina of the eye with efferent signals from the region of the cortical part of the visual analyzer takes place. With the participation of the reticular formation, interaction with the auditory and other sensory systems occurs here, which ensures the processes of selective visual attention by highlighting the most significant components of the sensory signal.

Central, or cortical, department the visual analyzer is located in the occipital lobe (fields 17, 18, 19 according to Brodmann) or VI, V2, V3 (according to the accepted nomenclature). It is believed that the primary projection area (field 17) carries out specialized, but more complex than in the retina and in the external geniculate bodies, information processing. The receptive fields of neurons in the visual cortex of small sizes are elongated, almost rectangular, rather than rounded. Along with this, there are complex and supercomplex receptive fields of the detector type. This feature allows you to select from the whole image only separate parts of lines with different locations and orientations, while the ability to selectively respond to these fragments is manifested.

In each area of ​​the cortex, neurons are concentrated, which form a column that passes vertically in depth through all layers, while there is a functional association of neurons that perform a similar function. Different properties of visual objects (color, shape, movement) are processed in different parts of the visual cortex of the large brain in parallel.

In the visual cortex there are functionally different groups of cells - simple and complex.

Simple cells create a receptive field, which consists of excitatory and inhibitory zones. This can be determined by examining the reaction of the cell to a small light spot. It is impossible to establish the structure of the receptive field of a complex cell in this way. These cells are detectors for the angle, tilt, and movement of lines in the field of view.

One column can contain both simple and complex cells. In the III and IV layers of the visual cortex, where the thalamic fibers end, simple cells were found. Complex cells are located in the more superficial layers of field 17; in fields 18 and 19 of the visual cortex, simple cells are an exception; complex and supercomplex cells are located there.

In the visual cortex, some neurons form "simple" or concentric color-opponent receptive fields (layer IV). The color opposition of RP is manifested in the fact that the neuron located in the center reacts with excitation to one color and is inhibited when stimulated by another color. Some neurons react with an on-response to red illumination and an ofT-response to green, while others react inversely.

In neurons with concentric RP, in addition to the opponent relations between color receivers (cones), there are antagonistic relations between the center and the periphery, i.e. there are RPs with double color opposing. For example, if an on-response to red and an off-response to green appear in the neuron upon exposure to the RP center, then its selectivity to color is combined with selectivity to the brightness of the corresponding color, and it does not respond to diffuse stimulation with light of any wavelength (from - for the opponent relations between the center and the periphery of the Republic of Poland).

In a simple RP, two or three parallel zones are distinguished, between which there is a double opposition: if the central zone has an on-response to red lighting and an off-response to green, then the edge zones give an off-response to red and an on-response to green.

From field VI, another (dorsal) canal passes through the middle temporal (mediotemporal - MT) region of the cortex. Registration of responses of neurons in this area showed that they are highly selective to disparity (non-identity), speed and direction of movement of objects in the visual world, and respond well to the movement of objects against a textured background. Local destruction sharply impairs the ability to respond to moving objects, but after a while this ability is restored, indicating that this area is not the only area where the analysis of moving objects in the visual field is performed. But along with this, it is assumed that the information extracted by the neurons of the primary visual field 17(V1) is then transferred for processing to the secondary (field V2) and tertiary (field V3) areas of the visual cortex.

However, the analysis of visual information does not end in the fields of the striate (visual) cortex (V1, V2, V3). It has been established that paths (channels) to other areas begin from field V1, in which further processing of visual signals is performed.

So, if the V4 field, which is located at the junction of the temporal and parietal regions, is destroyed in a monkey, then the perception of color and shape is disturbed. The processing of visual information about the form is also assumed to occur mainly in the lower temporal region. When this area is destroyed, the basic properties of perception (visual acuity and perception of light) do not suffer, but the analysis mechanisms of the highest level fail.

Thus, in the visual sensory system, the receptive fields of neurons become more complex from level to level, and the higher the synaptic level, the more severely the functions of individual neurons are limited.

Currently, the visual system, starting with ganglion cells, is divided into two functionally different parts (magna- and parvocellular). This division is due to the fact that in the retina of mammals there are ganglion cells of various types - X, Y, W. These cells have concentric receptive fields, and their axons form the optic nerves.

In X-cells - RP is small, with a well-defined inhibitory border, the speed of excitation conduction along their axons is 15-25 m/s. Y-cells have a much larger RP center and respond better to diffuse light stimuli. The speed of conduction is 35-50 m/s. In the retina, X-cells occupy the central part, and their density decreases towards the periphery. Y-cells are evenly distributed throughout the retina, so the density of Y-cells is higher than that of X-cells at the periphery of the retina. Structural features of X-cell RPs determine their better response to slow movements of the visual stimulus, while Y-cells respond better to fast-moving stimuli.

A large group of W cells has also been described in the retina. These are the smallest ganglion cells, the speed of conduction along their axons is 5-9 m/s. The cells of this group are not homogeneous. Among them are cells with concentric and homogeneous RPs and cells that are sensitive to the movement of the stimulus through the receptive field. In this case, the reaction of the cell does not depend on the direction of movement.

The division into X, Y, and W systems continues at the level of the geniculate body and the visual cortex. Neurons X have a phasic type of reaction (activation in the form of a short burst of impulses), their receptive fields are more represented in the peripheral fields of vision, the latent period of their reaction is shorter. Such a set of properties shows that they are excited by fast-conducting afferents.

Neurons X have a topical type of reaction (the neuron is activated within a few seconds), their RPs are more represented in the center of the visual field, and the latent period is longer.

The primary and secondary zones of the visual cortex (fields Y1 and Y2) differ in the content of X- and Y-neurons. For example, in the Y1 field, the lateral geniculate body receives afferent from both X- and Y-types, while the Y2 field receives afferents only from the Y-type cells.

The study of signal transmission at different levels of the visual sensory system is carried out by recording the total evoked potentials (EP) by removing a person with electrodes from the surface of the scalp in the visual cortex (occipital region). In animals, it is possible to simultaneously study the evoked activity in all parts of the visual sensory system.

Mechanisms that provide clear vision in various conditions

When considering objects located at different distances from the observer, The following processes contribute to clear vision.

1. Convergence and divergence eye movements due to which the reduction or dilution of the visual axes is carried out. If both eyes move in the same direction, such movements are called friendly.

2. pupil reaction, which occurs in sync with eye movement. So, with the convergence of the visual axes, when closely spaced objects are considered, the pupil narrows, i.e., a convergent reaction of the pupils. This response helps to reduce image distortion caused by spherical aberration. Spherical aberration is due to the fact that the refractive media of the eye have an unequal focal length in different areas. The central part, through which the optical axis passes, has a greater focal length than the peripheral part. Therefore, the image on the retina is blurred. The smaller the pupil diameter, the less distortion caused by spherical aberration. Convergent constriction of the pupil activates the accommodation apparatus, which causes an increase in the refractive power of the lens.

Rice. 2.4 The mechanism of accommodation of the eye: a - rest, b - tension

Rice. 2.5

The pupil is also an apparatus for eliminating chromatic aberration, which is due to the fact that the optical apparatus of the eye, like simple lenses, refracts light with a short wave more than with a long wave. Based on this, for a more accurate focusing of a red object, a greater degree of accommodation is required than for a blue one. That is why blue objects appear more distant than red objects, being located at the same distance.

3. Accommodation is the main mechanism that provides clear vision of objects at different distances, and is reduced to focusing the image from far or close objects on the retina. The main mechanism of accommodation is an involuntary change in the curvature of the lens of the eye (Fig. 2.4).

Due to the change in the curvature of the lens, especially the front surface, its refractive power can vary within 10-14 diopters. The lens is enclosed in a capsule, which at the edges (along the equator of the lens) passes into the lens-fixing ligament (zinn ligament), in turn, connected to the fibers of the ciliary (ciliary) muscle. With the contraction of the ciliary muscle, the tension of the zinn ligaments decreases, and the lens, due to its elasticity, becomes more convex. The refractive power of the eye increases, and the eye is tuned to the vision of nearby objects. When a person looks into the distance, the ligament of zon is in a taut state, which leads to stretching of the lens bag and its thickening. The innervation of the ciliary muscle is carried out by sympathetic and parasympathetic nerves. The impulse coming through the parasympathetic fibers of the oculomotor nerve causes muscle contraction. Sympathetic fibers extending from the upper cervical ganglion cause it to relax. The change in the degree of contraction and relaxation of the ciliary muscle is associated with the excitation of the retina and is influenced by the cerebral cortex. The refractive power of the eye is expressed in diopters (D). One diopter corresponds to the refractive power of a lens whose main focal length in air is 1 m. If the main focal length of a lens is, for example, 0.5 or 2 m, then its refractive power is 2D or 0.5D, respectively. The refractive power of the eye without the phenomenon of accommodation is 58-60 D and is called the refraction of the eye.

With normal refraction of the eye, rays from distant objects after passing through the refractive system of the eye are collected in focus on the retina in the fovea. Normal refraction of the eye is called emmetropia, and such an eye is called emmetropic. Along with normal refraction, its anomalies are observed.

Myopia (nearsightedness) is a type of refractive error in which the rays from an object, after passing through the light-refracting apparatus, are focused not on the retina, but in front of it. This may depend on the large refractive power of the eye or on the large length of the eyeball. A shortsighted person sees close objects without accommodation, distant objects are seen as unclear, vague. Glasses with diverging biconcave lenses are used for correction.

Hypermetropia (farsightedness) is a type of refractive error in which rays from distant objects, due to the weak refractive power of the eye or with a small length of the eyeball, are focused behind the retina. The far-sighted eye sees even distant objects with accommodation tension, as a result of which hypertrophy of the accommodation muscles develops. Biconvex lenses are used for correction.

Astigmatism is a type of refractive error in which the rays cannot converge at one point, at the focus (from the Greek stigme - point), due to the different curvature of the cornea and lens in different meridians (planes). With astigmatism, objects appear flattened or elongated, its correction is carried out with spherical lenses.

It should be noted that the refractive system of the eye also includes: the cornea, the moisture of the anterior chamber of the eye, the lens and the vitreous body. However, their refractive power, unlike the lens, is not regulated and does not participate in accommodation. After the rays pass through the refractive system of the eye, a real, reduced and inverted image is obtained on the retina. But in the process of individual development, the comparison of the sensations of the visual analyzer with the sensations of the motor, skin, vestibular and other analyzers, as noted above, leads to the fact that a person perceives the outside world as it really is.

Binocular vision (vision with two eyes) plays an important role in the perception of objects at different distances and determining the distance to them, gives a more pronounced sense of the depth of space compared to monocular vision, i.e. vision in one eye. When viewing an object with two eyes, its image can fall on symmetrical (identical) points of the retinas of both eyes, excitations from which are combined into a single whole at the cortical end of the analyzer, giving one image. If the image of an object falls on non-identical (disparate) areas of the retina, then a split image occurs. The process of visual analysis of space depends not only on the presence of binocular vision, a significant role in this is played by conditioned reflex interactions that develop between visual and motor analyzers. Of certain importance are convergent eye movements and the process of accommodation, which are controlled by the principle of feedback. The perception of space as a whole is associated with the definition of spatial relations of visible objects - their size, shape, relationship to each other, which is ensured by the interaction of various departments of the analyzer; acquired experience plays a significant role in this.

When moving objects The following factors contribute to clear vision:

1) voluntary eye movements up, down, left or right with the speed of the object, which is carried out due to the friendly activity of the oculomotor muscles;

2) when an object appears in a new part of the field of view, a fixation reflex is triggered - a rapid involuntary movement of the eyes, which ensures that the image of the object on the retina is aligned with the fovea. When tracking a moving object, a slow movement of the eyes occurs - a tracking movement.

When looking at a stationary object to ensure clear vision, the eye makes three types of small involuntary movements: tremor - eye trembling with a small amplitude and frequency, drift - a slow shift of the eye over a fairly significant distance, and jumps (flicks) - fast eye movements. There are also saccadic movements (saccades) - friendly movements of both eyes, performed at high speed. Saccades are observed when reading, viewing pictures, when the examined points of the visual space are at the same distance from the observer and other objects. If these eye movements are blocked, then the world around us, due to the adaptation of retinal receptors, will become difficult to distinguish, as it is in a frog. The frog's eyes are motionless, so it only distinguishes moving objects, such as butterflies, well. That is why the frog approaches the snake, which constantly throws its tongue out. The frog, which is in a state of immobility, does not distinguish, and its moving tongue takes it for a flying butterfly.

Under changing light conditions clear vision is provided by the pupillary reflex, dark and light adaptation.

Pupil regulates the intensity of the light flux acting on the retina by changing its diameter. Pupil width can vary from 1.5 to 8.0 mm. The constriction of the pupil (miosis) occurs with an increase in illumination, as well as when examining a closely located object and in a dream. Pupil dilation (mydriasis) occurs with a decrease in illumination, as well as with excitation of receptors, any afferent nerves, with emotional stress reactions associated with an increase in the tone of the sympathetic department of the nervous system (pain, anger, fear, joy, etc.), with mental excitations (psychosis, hysteria, etc.), with suffocation, anesthesia. Although the pupillary reflex improves visual perception when the illumination changes (it expands in the dark, which increases the light flux falling on the retina, narrows in the light), however, the main mechanism is still dark and light adaptation.

Tempo adaptation expressed in an increase in the sensitivity of the visual analyzer (sensitization), light adaptation- Decreased sensitivity of the eye to light. The basis of the mechanisms of light and dark adaptation is the photochemical processes occurring in cones and rods, which ensure the splitting (in the light) and resynthesis (in the dark) of photosensitive pigments, as well as the processes of functional mobility: turning on and off the activity of the receptor elements of the retina. In addition, adaptation is determined by some neural mechanisms and, above all, by the processes occurring in the nerve elements of the retina, in particular, the methods of connecting photoreceptors to ganglion cells with the participation of horizontal and bipolar cells. So, in the dark, the number of receptors connected to one bipolar cell increases, and more of them converge to the ganglion cell. This expands the receptive field of each bipolar and, of course, ganglion cells, which improves visual perception. The inclusion of horizontal cells is regulated by the central nervous system.

A decrease in the tone of the sympathetic nervous system (desympathization of the eye) reduces the rate of dark adaptation, and the introduction of adrenaline has the opposite effect. Irritation of the reticular formation of the brain stem increases the frequency of impulses in the fibers of the optic nerves. The influence of the central nervous system on adaptive processes in the retina is also confirmed by the fact that the sensitivity of the unlit eye to light changes when the other eye is illuminated and under the action of sound, olfactory, or taste stimuli.

Color adaptation. The most rapid and sharp adaptation (decrease in sensitivity) occurs under the action of a blue-violet stimulus. The red stimulus occupies a middle position.

Visual perception of large objects and their details provided by central and peripheral vision - changes in the angle of view. The most subtle assessment of the fine details of the object is provided if the image falls on the yellow spot, which is localized in the central fovea of ​​the retina, since in this case the greatest visual acuity occurs. This is explained by the fact that only cones are located in the area of ​​the macula, their sizes are the smallest, and each cone is in contact with a small number of neurons, which increases visual acuity. Visual acuity is determined by the smallest angle of view under which the eye is still able to see two points separately. A normal eye is able to distinguish between two luminous points at an angle of view of 1 ". The visual acuity of such an eye is taken as a unit. Visual acuity depends on the optical properties of the eye, the structural features of the retina and the work of the neuronal mechanisms of the conductive and central sections of the visual analyzer. Determination of visual acuity is carried out using alphabetic or various types of curly standard tables.Large objects in general and the surrounding space are perceived mainly due to peripheral vision, which provides a large field of view.

Field of view - the space that can be seen with a fixed eye. There is a separate field of view of the left and right eyes, as well as a common field of view for both eyes. The magnitude of the visual field in humans depends on the depth of the eyeball and the shape of the superciliary arches and nose. The boundaries of the visual field are indicated by the angle formed by the visual axis of the eye and the beam drawn to the extreme visible point through the nodal point of the eye to the retina. The field of view is not the same in different meridians (directions). Down - 70 °, up - 60 °, outward - 90 °, inside - 55 °. The achromatic field of view is larger than the chromatic one due to the fact that there are no color receptors (cones) on the periphery of the retina. In turn, the color field of view is not the same for different colors. Narrowest field of view for green, yellow, more for red, even more for blue. The size of the field of view varies depending on the illumination. The achromatic field of view increases at dusk and decreases in the light. The chromatic field of view, on the contrary, increases in the light, and decreases at dusk. It depends on the processes of mobilization and demobilization of photoreceptors (functional mobility). With twilight vision, an increase in the number of functioning rods, i.e. their mobilization leads to an increase in the achromatic field of view, at the same time, a decrease in the number of functioning cones (their demobilization) leads to a decrease in the chromatic field of view (PG Snyakin).

The visual analyzer also has a mechanism for differences in the wavelength of light - color vision.

Color vision, visual contrasts and sequential images

color vision - the ability of the visual analyzer to respond to changes in the wavelength of light with the formation of a sense of color. A certain wavelength of electromagnetic radiation corresponds to the sensation of a certain color. So, the sensation of red color corresponds to the action of light with a wavelength of 620-760 nm, and violet - 390-450 nm, the rest of the colors of the spectrum have intermediate parameters. Mixing all colors gives the impression of white. As a result of mixing the three primary colors of the spectrum - red, green, blue-violet - in different ratios, you can also get the perception of any other colors. The perception of colors is related to light. As it decreases, red colors cease to be distinguished first, and blue colors later than all. The perception of color is mainly due to the processes occurring in the photoreceptors. The most widely recognized is the three-component theory of color perception by Lomonosov - Jung - Helmholtz-Lazarev, according to which there are three types of photoreceptors in the retina - cones that separately perceive red, green and blue-violet colors. Combinations of excitation of different cones lead to the sensation of different colors and shades. Uniform excitation of three types of cones gives a sensation of white color. The three-component theory of color vision was confirmed in the electrophysiological studies of R. Granit (1947). Three types of color-sensitive cones were called modulators, cones that were excited when the brightness of the light changed (the fourth type) were called dominators. Subsequently, by microspectrophotometry, it was possible to establish that even a single cone can absorb rays of various wavelengths. This is due to the presence in each cone of various pigments that are sensitive to light waves of different lengths.

Despite the convincing arguments of the three-component theory in the physiology of color vision, facts are described that cannot be explained from these positions. This made it possible to put forward the theory of opposite, or contrasting, colors, i.e. create the so-called opponent theory of color vision by Ewald Hering.

According to this theory, there are three opponent processes in the eye and/or brain: one for the sensation of red and green, the second for the sensation of yellow and blue, and the third one, qualitatively different from the first two processes, for black and white. This theory is applicable to explain the transmission of information about color in the subsequent parts of the visual system: ganglion cells of the retina, lateral geniculate bodies, cortical centers of vision, where color-opposing RPs with their center and periphery function.

Thus, based on the data obtained, it can be assumed that the processes in cones are more consistent with the three-component theory of color perception, while Hering's theory of contrast colors is suitable for the neural networks of the retina and overlying visual centers.

In the perception of color, processes that take place in neurons of different levels of the visual analyzer (including the retina), which are called color-opponent neurons, also play a certain role. When the eye is exposed to radiation of one part of the spectrum, they are excited, and the other part is inhibited. Such neurons are involved in encoding color information.

Anomalies of color vision are observed, which can manifest as partial or complete color blindness. People who do not distinguish colors at all are called achromats. Partial color blindness occurs in 8-10% of men and 0.5% of women. It is believed that color blindness is associated with the absence in men of certain genes in the sexual unpaired X chromosome. There are three types of partial color blindness: protanopia(color blindness) - blindness mainly to red. This type of color blindness was first described in 1794 by the physicist J. Dalton, who had this type of anomaly. People with this type of anomaly are called "red-blind"; deuteranopia- Decreased perception of green color. Such people are called "green-blind"; tritanopia is a rare anomaly. At the same time, people do not perceive blue and purple colors, they are called "violet-blind".

From the point of view of the three-component theory of color vision, each type of anomaly is the result of the absence of one of the three cone color-receiving substrates. For the diagnosis of color perception disorders, color tables of E. B. Rabkin are used, as well as special devices called anomaloscopes. The identification of various color vision anomalies is of great importance in determining the professional suitability of a person for various types of work (driver, pilot, artist, etc.).

The ability to assess the length of a light wave, which manifests itself in the ability to perceive colors, plays a significant role in human life, influencing the emotional sphere and the activity of various body systems. Red color causes a feeling of warmth, has an exciting effect on the psyche, enhances emotions, but quickly tires, leads to muscle tension, increased blood pressure, and increased breathing. Orange color evokes a feeling of fun and well-being, and promotes digestion. Yellow color creates a good, high spirits, stimulates vision and the nervous system. This is the funniest color. Green color has a refreshing and calming effect, is useful for insomnia, overwork, lowers blood pressure, general body tone and is the most favorable for a person. The blue color causes a feeling of coolness and has a calming effect on the nervous system, and stronger than green (blue is especially favorable for people with increased nervous excitability), more than green, it lowers blood pressure and muscle tone. Violet is not so much calming as it relaxes the psyche. It seems that the human psyche, following along the spectrum from red to purple, goes through the whole gamut of emotions. This is the basis for the use of the Luscher test to determine the emotional state of the body.

Visual contrasts and consistent images. Visual sensations can continue even after the irritation has ceased. This phenomenon is called successive images. Visual contrasts are an altered perception of a stimulus depending on the surrounding light or color background. There are concepts of light and color visual contrasts. The phenomenon of contrast can manifest itself in an exaggeration of the actual difference between two simultaneous or successive sensations, therefore, simultaneous and successive contrasts are distinguished. A gray stripe on a white background appears darker than a gray stripe on a dark background. This is an example of simultaneous light contrast. When viewed against a red background, gray appears greenish; when viewed against a blue background, gray appears yellow. This is the phenomenon of simultaneous color contrast. Consistent color contrast is the change in color sensation when looking at a white background. So, if you look at a red-colored surface for a long time, and then look at a white one, then it acquires a greenish tint. The cause of visual contrast is the processes that are carried out in the photoreceptor and neuronal apparatus of the retina. The basis is the mutual inhibition of cells belonging to different receptive fields of the retina and their projections in the cortical section of the analyzers.

Molecule absorption cross section

Primary photochemical transformations are molecular quantum processes. In order to understand their regularities, let us consider the process of light absorption at the molecular level. To do this, we express the molar concentration of the chromophore C in terms of the “piece” concentration of its molecules (n = N/V is the number of molecules per unit volume):

Rice. 30.3. Geometric interpretation cross section absorption

In this case, equation (28.4) takes the following form:

The ratio of the natural molar absorption index to the Avogadro constant has the dimension [m 2 ] and is called absorption cross section of the molecule:

The cross section is molecular characteristic of the absorption process. Its value depends on the structure of the molecule, the wavelength of light and has the following geometric interpretation. Imagine a circle of area s, in the center of which is a molecule of this type. If the trajectory of a photon capable of causing photoexcitation of a molecule passes through this circle, then the photon is absorbed (Fig. 30.3).

Now we can write the equation for changing the intensity of light in a form that takes into account the molecular nature of absorption:

A molecule absorbs only one light quantum. In order to take into account photonic the nature of the absorption, we introduce a special value - photon flux intensity(I f).

Photon flux intensity- the number of photons incident along the normal onto the surface of a unit area per unit of time:

The number of photons also changes accordingly due to their absorption:

Quantum yield of a photochemical reaction

In order to relate the number of absorbed photons to the number of molecules that have entered into a photochemical reaction, we find out what happens to a molecule after absorption of a photon. Such a molecule can enter into a photochemical reaction or, having transferred the received energy to neighboring particles, return to the unexcited state. The transition from excitation to photochemical transformations is a random process that occurs with a certain probability.

- Anatomy of vision

Anatomy of vision

vision phenomenon

When scientists explain vision phenomenon , they often compare the eye to a camera. Light, just as it happens with the lenses of the apparatus, enters the eye through a small hole - the pupil, located in the center of the iris. The pupil can be wider or narrower: in this way, the amount of light entering is regulated. Further, the light is directed to the back wall of the eye - the retina, as a result of which a certain picture (image, image) appears in the brain. Similarly, when light hits the back of a camera, the image is captured on film.

Let's take a closer look at how our vision works.

First, the visible parts of the eye, to which they belong, receive light. Iris("input") and sclera(white of the eye). After passing through the pupil, the light enters the focusing lens ( lens) of the human eye. Under the influence of light, the pupil of the eye constricts without any effort or control of the person. This is because one of the muscles of the iris - sphincter- sensitive to light and reacts to it by expanding. The constriction of the pupil occurs due to the automatic control of our brain. Modern self-focusing cameras do much the same thing: a photoelectric "eye" adjusts the diameter of the entrance hole behind the lens, thus metering the amount of incoming light.

Now let's turn to the space behind the eye lens, where the lens is located, a vitreous gelatinous substance ( vitreous body) and finally - retina, an organ that is truly admired for its structure. The retina covers the vast surface of the fundus. It is a unique organ with a complex structure unlike any other body structure. The retina of the eye consists of hundreds of millions of light-sensitive cells called "rods" and "cones". unfocused light. sticks are designed to see in the dark, and when they are activated, we can perceive the invisible. Film can't do that. If you use film designed for shooting in dim light, it will not be able to capture a picture that is visible in bright light. But the human eye has only one retina, and it is able to operate in different conditions. Perhaps it can be called a multifunctional film. cones, unlike sticks, work best in the light. They need light to provide sharp focus and clear vision. The highest concentration of cones is in the area of ​​the retina called the macula ("spot"). In the central part of this spot is the fovea centralis (eye fossa, or fovea): it is this area that makes the most acute vision possible.

The cornea, pupil, lens, vitreous body, as well as the size of the eyeball - all this affects the focusing of light as it passes through certain structures. The process of changing the focus of light is called refraction (refraction). Light that is more accurately focused hits the fovea, while less focused light scatters on the retina.

Our eyes are capable of distinguishing about ten million gradations of light intensity and about seven million shades of colors.

However, the anatomy of vision is not limited to this. Man, in order to see, uses both his eyes and his brain at the same time, and for this a simple analogy with a camera is not enough. Every second, the eye sends about a billion pieces of information to the brain (more than 75 percent of all the information we perceive). These portions of light turn in consciousness into amazingly complex images that you recognize. Light, taking the form of these recognizable images, appears as a kind of stimulant for your memories of the events of the past. In this sense, vision acts only as a passive perception.

Almost everything we see is what we have learned to see. After all, we come into life without having any idea how to extract information from the light falling on the retina. In infancy, what we see means nothing or almost nothing to us. Impulses stimulated by light from the retina enter the brain, but for the baby they are only sensations, devoid of meaning. As a person grows up and learns, he begins to interpret these sensations, tries to understand them, to understand what they mean.

The visual sensations of humans and animals are also associated with photochemical processes. Light, reaching the retina, is absorbed by photosensitive substances (rhodopsin, or visual purple, in rods and iodopsin in cones). The mechanism of decomposition of these substances and their subsequent recovery has not yet been elucidated, but it has been established that the decomposition products cause irritation of the optic nerve, as a result of which electrical impulses pass through the nerve to the brain and a sensation of light arises. Since the optic nerve has branches over the entire surface of the retina, the nature of the irritation depends on where the photochemical decomposition occurred in the retina. Therefore, stimulation of the optic nerve makes it possible to judge the nature of the image on the retina and, consequently, the picture in the external space, which is the source of this image.

Depending on the illumination of certain areas of the retina, i.e., depending on the brightness of the object, the amount of photosensitive substance decomposing per unit time, and hence the strength of the light sensation, changes. However, attention should be paid to the fact that the eye is able to perceive images of objects well, despite the huge difference in their brightness. We quite clearly see objects illuminated by the bright sun, as well as the same objects in moderate evening illumination, when their illumination, and consequently their brightness (see § 73) change tens of thousands of times. This ability of the eye to adapt to a very wide range of brightness is called adaptation. Adaptation to brightness is achieved in several ways. Thus, the eye quickly responds to a change in brightness by changing the diameter of the pupil, which can change the area of ​​the pupil, and, consequently, the illumination of the retina about 50 times. The mechanism that provides adaptation to light in a much wider range (about 1000 times) operates much more slowly. In addition, the eye, as is known, has two types of sensitive elements: more sensitive - rods, and less sensitive - cones, which are able not only to react to light, but also to perceive a color difference. In the dark (in low light), rods play the main role (twilight vision). When moving to bright light, the visual purple in the rods quickly fades and they lose their ability to perceive light; only cones work, which are much less sensitive and for which the new lighting conditions may be quite acceptable. In this case, adaptation takes time corresponding to the time of "blinding" of the rods, and usually occurs within 2-3 minutes. If the transition to bright light is too abrupt, this protective process may not have time to occur, and the eye will go blind for a while or forever, depending on the severity of the blindness. Temporary loss of vision, well known to motorists, occurs when the headlights of oncoming vehicles are blinded.

The fact that in low light (at twilight) rods work, and not cones, makes it impossible to distinguish colors at twilight (“all cats are gray at night”).

As for the ability of the eye to distinguish colors in sufficiently bright light, when the cones come into action, this question cannot yet be considered completely resolved. Apparently, it comes down to the presence in our eye of three types of cones (or three types of mechanisms in each cone), sensitive to three different colors: red, green and blue, from various combinations of which the sensations of any color are made up. It should be noted that, despite the successes of recent years, direct experiments on the study of the structure of the retina do not yet allow us to assert with complete reliability the existence of the indicated triple apparatus, which is assumed by the three-color theory of color vision.

The presence in the eye of two types of photosensitive elements - rods and cones - leads to another important phenomenon. The sensitivity of both cones and rods to different colors is different. But for cones, the maximum sensitivity lies in the green part of the spectrum, as shown in § 68, the curve of the relative spectral sensitivity of the eye, constructed for daytime, cone vision. For rods, the sensitivity maximum is shifted to shorter wavelengths and lies approximately around . In accordance with this, in strong illumination, when the “daylight apparatus” is operating, red tones will seem brighter to us than blue ones; under low illumination with light of the same spectral composition, blue tones may appear brighter due to the fact that the “twilight apparatus”, i.e., sticks, works under these conditions. So, for example, a red poppy appears brighter than a blue cornflower in daylight, and, conversely, may appear darker in low light at dusk.

"Methodological development of the program section" - Compliance of educational technologies and methods with the goals and content of the program. Socio-pedagogical significance of the presented results of the application of methodological development. Diagnostics of the planned educational results. - Cognitive - transforming - general educational - self-organizing.

"Modular educational program" - Requirements for the development of the module. In German universities, the training module consists of disciplines of three levels. Module structure. The second level training courses are included in the module on other grounds. The content of an individual component is consistent with the content of other component components of the module.

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"Summer rest" - Musical relaxation, health tea. Carrying out monitoring of the regulatory framework of the subjects of the summer health campaign. Section 2. Work with personnel. Continuation of the study of dance and practical exercises. Development of recommendations based on the results of the past stages. Expected results. Stages of program execution.

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"KSE" - Basic concepts of a systematic approach. Concepts of modern natural science (CSE). Science as critical knowledge. - Whole - part - system - structure - element - set - connection - relation - level. The concept of "concept". Humanities Psychology Sociology Linguistics Ethics Aesthetics. Physics Chemistry Biology Geology Geography.

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