Binocular vision

Binocular vision is vision in which both eyes are used together. The word binocular comes from two Latin roots, bini for double, and oculus for eye. ^[1] Having two eyes confers at least four advantages over having one. First, it gives a creature a spare eye in case one is damaged. Second, it gives a wider field of view. For example, a human has a horizontal field of view of approximately 200 degrees with two eyes but only 160 degrees with one.^[2]. Third, it gives binocular summation in which the ability to detect faint objects is enhanced. ^[3] Fourth it can give stereopsis in which parallax provided by the two eyes' different positions on the head give precise depth perception. ^[4] Such binocular vision is usually accompanied by singleness of vision or binocular fusion, in which a single image is seen despite each eye's having its own image of any object. ^[4] Other phenomena of binocular vision include utrocular discrimination, eye dominance, allelotropia, and binocular rivalry.

Field of view and eye movements

Some animals, usually prey animals, have their two eyes positioned on opposite sides of their heads to give the widest possible field of view. Examples include rabbits, buffalos, and antelopes. In such animals, the eyes often move independently to increase the field of view. Even without moving their eyes, some birds have a 360-degree field of view.

Other animals, usually predatory animals, have their two eyes positioned on the front of their heads, thereby reducing field of view in favour of stereopsis. Examples include eagles, cats, and snakes.

Some predator animals, such as sperm whales, have their two eyes positioned on opposite sides of their heads. Other animals that are not necessarily predators, such as fruit bats and some primates also have forward facing eyes. These are usually animals that need fine depth discrimination, for example, to pick fruit or to find a branch.

When the eyes move laterally, in the same direction, this is called a version. When the eyes move in opposite directions, to an object closer than where the eyes are pointing or farther than where the eyes are pointing, this is called a vergence. When the eyes move in, it is a convergence eye movement; when the eyes move out, it is a divergence eye movement.

Some animals (including some humans, notably exotropes) use both of the above strategies. A starling, for example, has laterally placed eyes to cover a wide field of view, but can also move them together to point to the front so their fields overlap giving stereopsis. A remarkable example is the chameleon, whose eyes appear to be mounted on turrets, each moving independently of the other, up or down, left or right. Nevertheless, the chameleon can bring both of its eyes to bear on a single object when it is hunting, showing vergence and stereopsis.

Binocular summation

Binocular summation means that the detection threshold for a stimulus is lower with two eyes than with one. There are two forms. First, when trying to detect a faint signal, there is a statistical advantage of using two detectors over using one. Mathematically, the advantage is equal to the square root of 2, about 1.41. Second, when some cells in the visual cortex receive input from both eyes simultaneously, they show binocular facilitation, a greater level of activity than the sum of the two activities evoked separately from each eye. Any advantage in using two eyes in detection task over 1.41 is credited to this sort of mechanism, dubbed neural summation.

Binocular interaction

Apart from binocular summation, the two eyes can influence each other in at least three ways.

Utrocular discrimination

Utrocular discrimination is the ability to tell, when both eyes are open, to which eye a monocular stimulus was shown. ^[5]

Singleness of vision

Once the fields of view overlap, there is a potential for confusion between the left and right eye's image of the same object. This can be dealt with in two ways: one image can be suppressed, so that only the other is seen, or the two images can be fused. If two images of a single object are seen, this is known as double vision or diplopia. Fusion of the images from the two eyes is considered to be separate from stereopsis for at least two reasons. First, some disorders of binocular vision, such as strabismus can spare fusion but abolish stereopsis. Second, the depth of an object either much nearer to or farther from where the eyes are fixating can be accurately judged despite the images of the object appearing double.

Fusion of images occurs only in a small volume of visual space around where the eyes are fixating. Running through the fixation point in the horizontal plane is a curved line for which objects there fall on corresponding retinal points in the two eyes. This line is called the empirical horizontal horopter. There is also an empirical vertical horopter, which is effectively tilted away from the eyes above the fixation point and towards the eyes below the fixation point. The horizontal and vertical horopters mark the centre of the volume of singleness of vision. Within this thin, curved volume, objects nearer and farther than the horopters are seen as single. The volume is known as Panum's fusional area (it's presumably called an area because it was measured by Panum only in the horizontal plane). Outside of Panum's fusional area (volume), double vision occurs.

Eye dominance

When each eye has its own image of objects, it becomes impossible to align images outside of Panum's fusional area with an image inside the area. This happens when one has to point to a distant object with one's finger. When one looks at one's fingertip, it is single but there are two images of the distant object. When one looks at the distant object it is single but there are two images of one's fingertip. To point successfully, one of the double images has to take precedence and one be ignored or suppressed (eye dominance). The eye of the image that takes precedence is called the dominant eye.

Stereopsis

Stereopsis is an ability to make fine depth discriminations from parallax provided by the two eye's different positions on the head. There are two sorts: quantitative stereopsis, in which the depth seen is very similar to the actual depth of the object being judged, and qualitative stereopsis, in which the depth is correctly nearer or farther than the fixation point but the amount of depth does not grow with distance of the object from the fixation point. Quantitative stereopsis holds for small distances from the fixation plane (approximately within Panum's fusional area); qualitative stereopsis holds for larger distances from the fixation plane (outside of Panum's fusional area). Eventually an object can be moved so far from the fixation plane that there is no sense of depth of the double images--instead they appear to be on the fixation plane.

Allelotropia

Because the eyes are in different positions on the head, any object away from fixation and off the plane of the horopter has a different visual direction in each eye. Yet when the two monocular images of the object are fused, creating a Cyclopean image, the object has a new visual direction, essentially the average of the two monocular visual directions. This is called allelotropia. The origin of the new visual direction is a point approximately between the two eyes, the so-called cyclopean eye. The position of the cyclopean eye is not usually exactly centred between the eyes, but tends to be closer to the dominant eye.

Binocular rivalry

When very different images are shown to the same retinal regions of the two eyes, perception settles on one for a few moments, then the other, then the first, and so on, for as long as one cares to look. This alternation of perception between the images of the two eyes is called binocular rivalry.

Disorders of binocular vision

To maintain stereopsis and singleness of vision, the eyes need to be pointed accurately. The position of each eye in its orbit is controlled by six extraocular muscles. Slight differences in the length or insertion position or strength of the same muscles in the two eyes can lead to a tendency for one eye to drift to a different position in its orbit from the other, especially when one is tired. This is known as phoria. One way to reveal it is with the cover-uncover test. To do this test, look at a cooperative person's eyes. Cover one eye of that person with a card. Have the person look at your finger tip. Move the finger around; this is to break the reflex that normally holds a covered eye in the correct vergence position. Hold your finger steady and then uncover the person's eye. Look at the uncovered eye. You may see it flick quickly from being wall-eyed or cross-eyed to its correct position. If the uncovered eye moved from out to in, the person has exophoria. If it moved from in to out, the person has esophoria. If the eye did not move at all, the person has orthophoria. Most people have some amount of exophoria or esophoria; it is quite normal. If the uncovered eye also moved vertically, the person has hyperphoria (if the eye moved from up to down) or hypophoria (if the eye moved from down to up). Such vertical phorias are quite rare. It is also possible for the covered eye to rotate in its orbit. Such cyclophorias cannot be seen with the cover-uncover test; they are rarer than vertical phorias.

During the cover-uncover test, a person with some phoria will notice a brief episode of double vision or diplopia after uncovering the eye. This is a normal consequence of the eye's being briefly misaligned. If the diplopia is enduring, that is considered a disorder.

The cover-uncover test can also be used for more problematic disorders of binocular vision, the tropias. In the cover part of the test, the examiner looks at the first eye as he or she covers the second. If the eye moves from out to in, the person has exotropia. If it moved from in to out, the person has esotropia. People with exotropia or esotropia are wall-eyed or cross-eyed respectively. These are forms of strabismus with amblyopia. When the covered eye is the non-amblyopic eye, the amblyopic eye suddenly becomes the person's only means of seeing. The strabismus is revealed by the movement of that eye to fixate on the examiner's finger. There are also vertical tropias (hypertropia and hypotropia) and cyclotropias.

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