- Anatomy of the visual apparatus
- Structures auxiliary to the eye
- The visual process
- The work of the auxiliary structures
- The work of the retina
- Some basic facts of vision
- Absolute threshold and minimum stimulus for vision
- Electrophysiology of the retina
- Colour vision
- The higher visual centres
- The visual pathway
- Some perceptual aspects of vision
- Electrophysiology of the visual centres
The visual process
The work of the auxiliary structures
The protective mechanisms
The first line of protection of the eyes is provided by the lids, which prevent access of foreign bodies and assist in the lubrication of the corneal surface. Lid closure and opening are accomplished by the orbicularis oculi and levator palpebri muscles; the orbicularis oculi operates on both lids, bringing their margins into close apposition in the act of lid closure. Opening results from relaxation of the orbicularis muscle and contraction of the levator palpebri of the upper lid; the smooth muscle of the upper lid, Müller’s muscle, or the superior palpebral muscle, also assists in widening the lid aperture. The lower lid does not possess a muscle corresponding to the levator of the upper lid, and the only muscle available for causing an active lowering of the lid, required during the depression of the gaze, is the inferior palpebral muscle, which is analogous to the muscle of Müller of the upper lid (called the superior palpebral muscle). This inferior palpebral muscle is so directly fused with the sheaths of the ocular muscles that it provides cooperative action, opening of the lid on downward gaze being mediated, in effect, mainly by the inferior rectus.
The seventh cranial nerve—the facial nerve—supplies the motor fibres for the orbicularis muscle. The levator is innervated by the third cranial nerve—the oculomotor nerve—which also innervates some of the extraocular muscles concerned with rotation of the eyeball, including the superior rectus. The smooth muscle of the eyelids and orbit is activated by the sympathetic division of the autonomic system. The secretion of epinephrine (adrenaline) during such states of excitement as fear would also presumably cause contraction of the smooth muscle, but it seems unlikely that this would lead to the protrusion of the eyes traditionally associated with extreme fear. It is possible that the widening of the lid aperture occurring in this excited state and the dilation of the pupil create the illusion of eye protrusion.
Blinking is normally an involuntary act, but it may be carried out voluntarily. The more vigorous “full closure” of the lids involves the orbital portion of the orbicularis muscle and may be accompanied by contraction of the facial muscles that have been described as accessory muscles of blinking—namely, the corrugator supercilii, which on contraction pulls the eyebrows toward the bridge of the nose, and the procerus or pyramidalis, which pulls the skin of the forehead into horizontal folds, acting as a protection when the eyes are exposed to bright light. The more vigorous full closure may be evoked as a reflex response.
Reflex blinking may be caused by practically any peripheral stimulus, but the two functionally significant reflexes are (1) that resulting from stimulation of the endings of the fifth cranial nerve in the cornea, lid, or conjunctiva—the sensory blink reflex, or corneal reflex—and (2) that caused by bright light—the optical blink reflex. The corneal reflex is rapid (0.1 second reflex time) and is the last to disappear in deepening anesthesia, impulses being relayed from the nucleus of the fifth nerve to the seventh cranial nerve, which transmits the motor impulses. The reflex is said to be under the control of a medullary centre. The optical reflex is slower; in humans, the nervous pathway includes the visual cortex (the outer substance of the brain; the visual centre is located in the occipital—rear—lobe).
In the waking hours, the eyes blink fairly regularly at intervals of two to 10 seconds, the actual rate being a characteristic of the individual. The function of this is to spread the lacrimal secretions over the cornea. It might be thought that each blink would be reflexly determined by a corneal stimulus—drying and irritation—but extensive studies indicate that this view is incorrect. The normal blinking rate is apparently determined by the activity of a “blinking centre” in the globus pallidus of the caudate nucleus, a mass of neurons between the base and the outer substance of the brain. This is not to deny that the blink rate is modified by external stimuli.
There is a strong association between blinking and the action of the extraocular muscles. Eye movement is generally accompanied by a blink, and it is thought that this aids the eyes in changing their fixation point.
Secretion of tears
The exposed surface of the globe (eyeball) is kept moist by the tears secreted by the lacrimal apparatus, together with the mucous and oily secretions of the other secretory organs and cells of the lids and conjunctiva; these have been described earlier. The secretion produces what has been called the precorneal film, which consists of an inner layer of mucus, a middle layer of lacrimal secretion, and an outer oily film that reduces the rate of evaporation of the underlying watery layer. The normal daily (24-hour) rate of secretion has been estimated at about 0.75 to 1.1 grams (0.03–0.04 ounce avoirdupois); secretion tends to decrease with age. Chemical analysis of the tears reveals a typical body fluid with a salt concentration similar to that of blood plasma. An interesting component is lysozyme, an enzyme that has bactericidal action by virtue of its power of dissolving away the outer coats of many bacteria.
Tears are secreted reflexly in response to a variety of stimuli—e.g., irritative stimuli to the cornea, conjunctiva, nasal mucosa; hot or peppery stimuli applied to the mouth and tongue; or bright lights. In addition, tear flow occurs in association with vomiting, coughing, and yawning. The secretion associated with emotional upset is called psychical weeping. Severing of the sensory root of the trigeminal (fifth cranial) nerve prevents all reflex weeping, leaving psychical weeping unaffected; similarly, the application of cocaine to the surface of the eye, which paralyzes the sensory nerve endings, inhibits reflex weeping, even when the eye is exposed to potent tear gases. The afferent (sensory) pathway in the reflex is thus by way of the trigeminal nerve. The motor innervation is by way of the autonomic (involuntary) division; the parasympathetic supply derived from the facial nerve (the seventh cranial nerve) seems to have the dominant motor influence. Thus, drugs that mimic the parasympathetic, such as acetylcholine, provoke secretion, and secretion may be blocked by such typical anticholinergic drugs as atropine. Innervation of the lacrimal gland is not always complete at birth, so that the newborn infant is generally said to cry without weeping. Because absence of reflex tearing fails to produce any serious drying of the cornea, and surgical destruction of the main lacrimal gland is often without serious consequences, it seems likely that the subsidiary secretion from the accessory lacrimal glands is adequate to keep the cornea moist. The reflex secretion that produces abundant tears may be regarded as an emergency response.
A drainage mechanism for tears is necessary only during copious secretion. The mechanism, described as the lacrimal pump, consists of alternately negative and positive pressure in the lacrimal sac caused by the contraction of the orbicularis muscle during blinking.