The simplicity of the motor systems involved in controlling eye musculature make them ideal for illustrating the mechanisms and principals you have been studying in the preceding material on motor systems. They involve the action of few muscles and of well defined neural circuits.
We use our eyes to monitor our external environment and depend on our ocular motor systems to protect and guide our eyes. The ocular motor systems control eye lid closure, the amount of light that enters the eye, the refractive properties of the eye, and eye movements. The visual system provides afferent input to ocular motor circuits that use visual stimuli to initiate and guide the motor responses. Neuromuscular systems control the muscles within the eye (intraocular muscles); the muscles attached to the eye (extraocular muscles) and the muscles in the eyelid. Ocular motor responses include ocular reflexes and voluntary motor responses to visual and other stimuli. The complexity of the circuitry (the chain or network of neurons) controlling a ocular motor response increases with the level of processing involved in initiating, monitoring, and guiding the response.
In this chapter we will start at the level of reflex responses and move onto more complex voluntary responses in the following lecture. The eye blink reflex is the simplest response and does not require the involvement of cortical structures. In contrast, voluntary eye movements (i.e., visual tracking of a moving object) involve multiple areas of the cerebral cortex as well as basal ganglion, brain stem and cerebellar structures.
7.2 Ocular Reflex Responses
The ocular reflexes are the simplest ocular motor responses. Ocular reflexes compensate for the condition of the cornea and for changes in the visual stimulus. For example, the eye blink reflex protects the cornea from drying out and from contact with foreign objects. The pupillary light reflex compensates for changes in illumination level, whereas the accommodation responses compensate for changes in eye-to-object-viewed distance. Note that reflex responses are initiated by sensory stimuli that activate afferent neurons (e.g., somatosensory stimuli for the eye blink reflex and visual stimuli for the pupillary light reflex and accommodation responses).
In general, ocular reflexes are consensual (i.e., the response is bilateral involving both eyes). Consequently, a light directed in one eye elicits responses, pupillary constriction, in both eyes. In this chapter you will learn of the structures normally involved in performing these ocular responses and the disorders that result from damage to components of neural circuit controlling these responses.
A. The Eye Blink Reflex
Tactile stimulation of the cornea results in an irritating sensation that normally evokes eyelid closure (an eye blink). The response is consensual (i.e., bilateral) - involving automatic eyelid closure at both eyes.
The corneal eye blink reflex neural circuit: This neural circuit (Figure 7.1) is relatively simple, consisting of the
- trigeminal1° afferent (free nerve endings in the cornea, trigeminal nerve, ganglion, root, and spinal trigeminal tract), which end on
- trigeminal 2° afferent in the spinal trigeminal nucleus, some of which send their axons to
- reticular formation interneurons, which send their axons bilaterally to
- facial motor neurons in the facial nucleus, which send their axons in the facial nerve to
- orbicularis oculi, which functions to lower the eyelid
B. Pupillary Light Reflex
The pupillary light reflex involves adjustments in pupil size with changes in light levels.
- The reflex is consensual: Normally light that is directed in one eye produces pupil constriction in both eyes.
- The direct response is the change in pupil size in the eye to which the light is directed (e.g., if the light is shone in the right eye, the right pupil constricts).
- The consensual response is the change in pupil size in the eye opposite to the eye to which the light is directed (e.g., if the light is shone in the right eye, the left pupil also constricts consensually).
The pupillary light reflex allows the eye to adjust the amount of light reaching the retina and protects the photoreceptors from bright lights. The iris contains two sets of smooth muscles that control the size of the pupil (Figure 7.2).
- The sphincter muscle fibers form a ring at the pupil margin so that when the sphincter contracts, it decreases (constricts) pupil size.
- The dilator muscle fibers radiate from the pupil aperture so that when the dilator contracts, it increases (dilates) pupil size.
Both muscles act to control the amount of light entering the eye and the depth of field of the eye1.
- The iris sphincter is controlled by the parasympathetic system, whereas the iris dilator is controlled by the sympathetic system.
- The action of the dilator is antagonistic to that of the sphincter and the dilator must relax to allow the sphincter to decrease pupil size.
Normally the sphincter action dominates during the pupillary light reflex.
Figure 7.2 |
The pupillary light reflex neural circuit: The pathway controlling pupillary light reflex (Figure 7.3) involves the
- retina, optic nerve, optic chiasm, and the optic tract fibers that join the
- brachium of the superior colliculus, which terminate in the
- pretectal area of the midbrain, which sends most of its axons bilaterally in the posterior commissure to terminate in the
- Edinger-Westphal nucleus of the oculomotor complex, which contains parasympathetic preganglionic neurons and sends its axons in the oculomotor nerve to terminate in the
- ciliary ganglion, which sends its parasympathetic postganglionic axons in the
- short ciliary nerve, which ends on the
- iris sphincter
Recall that the optic tract carries visual information from both eyes and the pretectal area projects bilaterally to both Edinger-Westphal nuclei: Consequently, the normal pupillary response to light is consensual. That is, a light directed in one eye results in constriction of the pupils of both eyes.
The pupils normally dilate (increase in size) when it is dark (i.e., when light is removed). This response involves the relaxation of the iris sphincter and contraction of the iris dilator. The iris dilator is controlled by the sympathetic nervous system.
The pupillary dark reflex neural circuit: The pathway controlling pupil dilation involves the
- retina and the optic tract fibers terminating on neurons in the hypothalamus and the
- axons of the hypothalamic neurons that descend to the spinal cord to end on the
- sympathetic preganglionic neurons in the lateral horn of spinal cord segments T1 to T3, which send their axons out the spinal cord to end on the
- sympathetic neurons in the superior cervical ganglion, which send their
- sympathetic postganglionic axons in the long ciliary nerve to the
- iris dilator.
Axons from the superior cervical ganglion also innervate the face vasculature, sweat and lachrymal glands and the eyelid tarsal muscles. When the superior cervical ganglion or its axons are damaged, a constellation of symptoms, known as Horner's syndrome, result. This syndrome is characterized by miosis (pupil constriction), anhidrosis (loss of sweating), pseudoptosis (mild eyelid droop), enopthalmosis (sunken eye) and flushing of the face.
D. The Accommodation Response
The accommodation response is elicited when the viewer directs his eyes from a distant (greater than 30 ft. away) object to a nearby object (Nolte, Figure 17-40, Pg. 447). The stimulus is an “out-of-focus” image. The accommodation (near point) response is consensual (i.e., it involves the actions of the muscles of both eyes). The accommodation response involves three actions:
Pupil accommodation: The action of the iris sphincter was covered in the section on the pupillary light reflex. During accommodation, pupil constriction utilizes the "pin-hole" effect and increases the depth of focus of the eye by blocking the light scattered by the periphery of the cornea (Nolte, Figure 17-39, Pg. 447). The iris sphincter is innervated by the postganglionic parasympathetic axons (short ciliary nerve fibers) of the ciliary ganglion (Figure 7.3).
Lens accommodation: Lens accommodation increases the curvature of the lens, which increases its refractive (focusing) power. The ciliary muscles are responsible for the lens accommodation response. They control the tension on the zonules, which are attached to the elastic lens capsule at one end and anchored to the ciliary body at the other end (Figure 7.4).
Figure 7.4 |
Figure 7.5 |
Convergence in accommodation: When shifting one's view from a distant object to a nearby object, the eyes converge (are directed nasally) to keep the object's image focused on the foveae of the two eyes. This action involves the contraction of the medial rectus muscles of the two eyes and relaxation of the lateral rectus muscles. The medial rectus attaches to the medial aspect of the eye and its contraction directs the eye nasally (adducts the eye). The medial rectus is innervated by motor neurons in the oculomotor nucleus and nerve.
The accommodation neural circuit: The circuitry of the accommodation response is more complex than that of the pupillary light reflex (Figure 7.6).
The afferent limb of the circuit includes the
- retina (with the retinal ganglion axons in the optic nerve, chiasm and tract),
- lateral geniculate body (with axons in the optic radiations), and
- visual cortex.
Ocular motor control neurons are interposed between the afferent and efferent limbs of this circuit and include the
- visual association cortex, which
- determines the image is "out-of-focus"
- sends corrective signals via the internal capsule and crus cerebri to the
- determines the image is "out-of-focus"
- supraoculomotor nuclei, which
- is located immediately superior to the oculomotor nuclei
- generates motor control signals that initiate the accommodation response
- sends these control signals bilaterally to the oculomotor complex.
- is located immediately superior to the oculomotor nuclei
The efferent limb of this system has two components: the
- Edinger-Westphal nucleus, which
- sends its axons in the oculomotor nerve to
- control the ciliary ganglion, which
- sends it axons in the short ciliary nerve to
- control the iris sphincter and the ciliary muscle/zonules/lens of the eye
- sends it axons in the short ciliary nerve to
- sends its axons in the oculomotor nerve to
- oculomotor neurons, which
- sends its axons in the oculomotor nerve to
- control the medial rectus
- converge the two eyes.
- sends its axons in the oculomotor nerve to
Figure 7.6 |
An excellent way to test your knowledge of the material presented thus far is by examining the effects of damage to structures within the ocular motor pathways. The observed motor loss(s) provide clues to the pathway(s) affected; and the muscle(s) and eye affected provide clues to the level of the damage.
Cranial nerve damage: Damage to cranial nerves may result in sensory and motor symptoms. The sensory losses would involve those sensations the cranial nerve normally conveys (e.g., taste from the anterior two thirds of the tongue and somatic sensations from the skin of the ear - if facial nerve is damaged). The motor losses may be severe (i.e., a lower motor neuron loss that produces total paralysis) if the cranial nerve contains all of the motor axons controlling the muscles of the normally innervated area.
The cranial nerves involved in the eye blink response and pupillary response are the optic, oculomotor, trigeminal and facial nerves.
- The optic nerve carries visual information from the eye.
- The oculomotor nerve contains
- lower motor axons innervating
- extraocular muscles: the medial, superior and inferior rectus muscles, the inferior oblique muscle,
- eyelid muscle: the superior levator palpebrae,
- extraocular muscles: the medial, superior and inferior rectus muscles, the inferior oblique muscle,
- as well as parasympathetic preganglionic axons to the ciliary ganglion.
- lower motor axons innervating
- The trigeminal nerve contains
- the 1° somatosensory afferents for the face, dura, oral and nasal cavities
- the lower motor axons for the jaw muscles.
- the 1° somatosensory afferents for the face, dura, oral and nasal cavities
- The facial nerve contains
- the lower motor neurons innervating the superficial muscles of the face,
- the 1° gustatory afferents to the anterior tongue
- the parasympathetic preganglionic axons to parasympathetic ganglia for the lachrymal and salivary glands.
- the lower motor neurons innervating the superficial muscles of the face,
Symptoms. The patient, who appears with a bloodshot left eye, complains of an inability to close his left eye. When asked to rise his eyebrows, he can only elevate the right eyebrow. When asked to close both eyes, the right eyelid closes but the left eyelid is only partially closed. Touching the right or left cornea with a wisp of cotton elicits the eye blink reflex in the right eye, but not the left eye (Figure 7.7). However, the patient reports he can feel the cotton when it touches either eye. He can smile, whistle and show his teeth, which indicates his lower facial muscles are functioning normally. Physical examination determines that touch, vibration, position and pain sensations are normal over the entire the body and face. There are no other motor symptoms.
Figure 7.7 |
Observation: You observe that the patient
- has not lost cutaneous sensation in the upper left face area
- does not blink when his left cornea is touched
- cannot close his left eye voluntarily
You conclude that his left eye's functional loss is
- not sensory
- a lower motor neuron dysfunction
Pathway(s) affected: You conclude that structures in the following motor pathway have been affected
- the eye blink pathway (Figure 7.8)
Figure 7.8 |
Side & Level of damage: As the eye blink loss involves
- only motor function
- both reflex and voluntary motor functions
- the upper part of the face
- only one eye lid
- eyelid closure
Conclusion: You conclude that the damage involves
- the facial nerve
- a branch of the nerve innervating the upper face
- a lower motor neuron paralysis of the left orbicularis oculi
- motor innervation on the left side (i.e., the symptoms are ipsilesional)
When lower motor neurons are damaged, there is a flaccid paralysis of the muscle normally innervated. The action of the muscle will be weakened or lost depending on the extent of the damage. There will be a weakened or no reflex response and the muscle will be flaccid and may atrophy with time.
The Facial Nerve. Section of the facial nerve on one side will result in paralysis of the muscles of facial expression on the ipsilesional side of the face. There will be an inability to close the denervated eyelid voluntarily and reflexively. The eyelids may have some mobility if the oculomotor innervation to the levator is unaffected.
7.5 Clinical Example #2Symptoms. The patient complains of a badly infected left eye. When he is asked to close both eyes, both eyelids close. Touching the right cornea with a wisp of cotton elicits the eye blink reflex in the both eyes (Figure 7.9, Right). However, touching the left cornea with a wisp of cotton does not elicit the eye blink reflex in the either eye (Figure 7.9, Left). The patient cannot detect pinpricks to his left forehead. However, he reports that pinpricks to rest of his face are painful. He can blink, wrinkle his brows, smile, and whistle and show his teeth, which indicates his facial muscles are functioning normally. Physical examination determines that touch, vibration, position and pain sensations are normal over the entire the body and over the lower left and right side of his face.
Figure 7.9 |
Observation: You observe that the patient
- responds with direct and consensual eye blink when his right cornea is touched
- can close his left eye voluntarily
- has lost cutaneous sensation in the upper left face area
- does not blink when his left cornea is touched
You conclude that his left eye's functional loss is
- not motor
- sensory
Pathway(s) affected: You conclude that structures in the following reflex pathway have been affected
- the eye blink pathway (Figure 7.8)
Side & Level of damage: As the eye blink loss involves
- only one eye
- a sensory loss
- the upper part of the face
Conclusion: You conclude that the damage involves
- a loss of the afferent limb of the eye blink response
- the trigeminal nerve
- a branch of the nerve innervating the upper face
- the innervation of the left side (i.e., the symptoms are ipsilesional)
The Trigeminal Nerve. Section of the trigeminal nerve will eliminate somatosensory sensation from the face and the eye blink reflex (e.g., with section of the left trigeminal nerve, light touch of the left cornea will not produce an eye blink in the left or right eye). However, light touch of the right cornea will elicit a bilateral eye blink. The effect of sectioning the trigeminal nerve is to remove the afferent input for the eye blink reflex.
Symptoms. The patient complains of pain in her left eye. Her left pupil appears dilated and is not reactive to light directed at either the left or right eye (Figure 7.10). The right pupil appears normal in size and reacts to light when it is directed in the right or left eye. Both eyelids can be elevated and lowered and both eyes exhibit normal movement. Touch, vibration, position and pain sensations are normal over the entire the body and face. There are no other motor symptoms.
Figure 7.10 |
|
Observation: You observe that the patient has
- a left pupil that does not react to light directly or consensually
- a right pupil that reacts to light directly and consensually
- normal eye movements
You conclude that his left eye's functional loss is
- not sensory (the right pupil reacts to light directed at the left eye)
- a motor dysfunction
Pathway(s) affected: You conclude that structures in the following motor pathway have been affected
- the pupillary light reflex pathway (Figure 7.11)
Figure 7.11 |
|
Side & Level of damage: As the pupillary light reflex loss
- involves only one eye
- involves only motor function
- does not involve eyelid or ocular motility
- is limited to pupil constriction in the left eye
Conclusion: You conclude that the damage
- involves the motor innervation of the left iris sphincter2
- involves structures peripheral to the oculomotor nucleus (i.e., eye movement unaffected)
- does not involve the oculomotor nerve
- involves the ciliary ganglion or the short ciliary nerve
- is on the left side (i.e., the symptoms are ipsilesional)
Parasympathetic Innervation of the Eye. Section of the parasympathetic preganglionic (oculomotor nerve) or postganglionic (short ciliary nerve) innervation to one eye will result in a loss (motor) of both the direct and consensual pupillary light responses of the denervated eye. Section of the left short ciliary nerve or a benign lesion in the left ciliary ganglion will result in no direct response to light in the left eye and no consensual response in the left eye when light is directed on the right eye (a.k.a., tonic pupil). When the damage is limited to the ciliary ganglion or the short ciliary nerve, eyelid and ocular mobility are unaffected.
7.7 Clinical Example #4Symptoms. The patient presents with a left eye characterized by ptosis, lateral strabismus, and dilated pupil. When asked to rise his eyelids, he can only raise the lid of the right eye. When asked to close both eyes, both eyelids close fully. His left pupil does not react to light directly or consensually (Figure 7.12). When asked to look to his right, his left eye moves to a central position, but no further. The right eye is fully mobile. When the patient is asked to look straight ahead, you note his left eye remains directed to the left and depressed. Physical examination determines that touch, vibration, position and pain sensations are normal over the entire the body and face. There are no other motor symptoms.
Observation: You observe that the patient
- has not lost cutaneous sensation in the face area
- has a left ptosis
- cannot adduct his left eye (i.e., move it toward the nose)
- has a left dilated pupil that is non reactive to light in either eye
You conclude that his left eye's functional loss is
- not sensory
- a lower motor neuron dysfunction
- involving an autonomic dysfunction
Pathway(s) affected: You conclude that structures in the following motor pathway have been affected
- the pupillary/oculomotor pathway (Figure 7.11)
Side & Level of damage: As the ocular loss involves
- only motor function
- both reflex and voluntary motor functions
- both somatic and autonomic functions
- only the left eye
Conclusion: You conclude that the damage
- involves the oculomotor nerve
- is a lower motor neuron paralysis of the superior levator palpebrae
- is a lower motor neuron paralysis of the medial, superior & inferior rectus muscles and inferior oblique muscles of the eye
- is an autonomic disorder involving the axons of the Edinger-Westphal nucleus
- is on the left side (i.e., the symptoms are ipsilesional)
The Oculomotor Nerve. Section of the oculomotor nerve produces a non-reactive pupil in the ipsilesional side as well as other symptoms related to oculomotor nerve damage (e.g., ptosis and lateral strabismus). Section of the oculomotor nerve on one side will result in paralysis of the superior levator palpebrae, which normally elevates the eyelid. It will also paralyze the medial, superior & inferior rectus muscles and the inferior oblique, which will allow the lateral rectus to deviate the eye laterally and the superior oblique to depress the eye. The parasympathetic preganglionic axons of the Edinger-Westphal nucleus, which normally travel in the oculomotor nerve, will be cut off from the ciliary ganglion, disrupting the circuit normally used to control the iris sphincter response to light.
7.8 Clinical Example #5Symptoms. The patient complains of reduced vision in the left eye. Pupil size in both eyes appears normal. However, both pupils do not appear to constrict as rapidly and strongly when light is directed into his left eye (Figure 7.13). That is, compared to the response to light in the left eye, light in the right eye produces a more rapid constriction and smaller pupil in both eyes. Physical examination determines that touch, vibration, position and pain sensations are normal over the entire the body and over the lower left and right side of his face.
Figure 7.13 |
|
Observation: You observe that the patient's pupils
- respond when light is directed into either eye
- has weaker direct and consensual responses to light directed in the left eye
You conclude that his left eye's functional loss is
- not motor
- sensory (because the responses in both eyes are weaker when light is directed in the left eye)
Pathway(s) affected: You conclude that structures in the following motor pathway have been affected
- the pupillary light reflex pathway (Figure 7.11)
Side & Level of damage: As the pupillary light response deficit involves
- only stimulation of one eye
- a sensory loss
- the left eye
Conclusion: You conclude that the damage
- is in the afferent limb of the pupillary light response
- involves the optic nerve or retina
- is on the left side (i.e., the symptoms are ipsilesional)
- produced a left pupillary afferent defect
The Optic Nerve. Partial damage of the retina or optic nerve reduces the afferent component of the pupillary reflex circuit. The reduced afferent input to the pretectal areas is reflected in weakened direct and consensual pupillary reflex responses in both eyes (a.k.a., a relative afferent pupillary defect).
Section of one optic nerve will result in the complete loss of the direct pupillary light reflex but not the consensual reflex of the blinded eye. That is, if the left optic nerve is sectioned, light directed on the left (blind) eye will not elicit a pupillary response in the left eye (direct reflex) or the right eye (consensual response). However, light directed in the right eye will elicit pupillary responses in the right eye and the left (blind) eye. The effect of sectioning one optic nerve is to remove the afferent input for the direct reflex of the blinded eye and the afferent input for the consensual reflex of the normal eye. Section of one optic tract will not eliminate the direct or consensual reflex of either eye as the surviving optic tract contains optic nerve fibers from both eyes. However, the responses to light in both eyes may be weaker because of the reduced afferent input to the ipsilesional pretectal area.
Symptoms. A patient who is suffering from the late stages of syphilis is sent to you for a neuro-ophthalmological work-up. His vision is normal when corrected for refractive errors. He has normal ocular mobility and his eyelids can be elevated and depressed at will. Examination of his pupillary responses indicates a loss of the pupillary light reflex (no pupil constriction to light in either eye) but normal pupillary accommodation response (pupil constricts when the patient's eyes are directed from a distant object to one nearby).
Observation: You observe that the patient has normal vision but that his pupils
- do not respond when light is directed into the either of his eyes
- do respond during accommodation
You conclude that his eye's functional loss is
- not sensory (his vision is normal)
- motor (the pupillary light responses in both eyes are absent)
- higher-order motor (because he has a normal pupillary accommodation response)
Pathway(s) affected: You conclude that structure(s) in the
- accommodation pathway have not been damaged (Figure 7.14)
- pupillary light reflex pathway have been damaged (Figure 7.11)
Side & Level of damage: As the pupillary response deficit
- does not involve a sensory loss
- does not involve the pupil accommodation response
- involves only the pupillary light reflex response
Conclusion: You conclude that the damage
- involves the pretectal area bilaterally
- spared the supraoculomotor area
- produced the Argyll Robertson response
In the Argyll Robertson response, there is an absence of the pupillary light reflex with a normal pupillary accommodation response. The Argyll Robertson response is attributed to bilateral damage to pretectal areas (which control the pupillary light reflex) with sparing of the supraoculomotor area (which controls the pupillary accommodation reflex).
The accommodation response involves many of the structures involved in the pupillary light response and, with the exception of the pretectal area and supraoculomotor area, damage to either pathway will produce common the symptoms. The most common complaint involving the accommodation response is its loss with aging (i.e., presbyopia). Recall that presbyopia most commonly results from structural changes in the lens which impedes the lens accommodation response.
7.10 Summary
This chapter described three types of ocular motor responses (the eye blink, pupillary light and accommodation responses) and reviewed the nature of the responses and the effectors, efferent neurons, higher-order motor control neurons (if any), and afferent neurons normally involved in performing these ocular responses. Table I summarizes these structures and the function(s) of these ocular motor responses. Readers should understand the anatomical basis for disorders that result from damage to components of neural circuit controlling these responses.
Spinal Trigeminal Nucleus*- Question 1
- A
- B
- C
- D
- E
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve
B. abducens nucleus
C. Edinger-Westphal nucleus
D. pretectal areas
E. supraoculomotor nucleus
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve This answer is INCORRECT.
Optic nerve is incorrect as section of one nerve would not obliterate the consensual response to stimulation of the contralesional eye.
B. abducens nucleus
C. Edinger-Westphal nucleus
D. pretectal areas
E. supraoculomotor nucleus
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve
B. abducens nucleus This answer is INCORRECT.
Abducens nucleus is incorrect as it is not involved in pupillary responses. Its motor neurons innervate the lateral rectus muscle.
C. Edinger-Westphal nucleus
D. pretectal areas
E. supraoculomotor nucleus
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve
B. abducens nucleus
C. Edinger-Westphal nucleus This answer is INCORRECT.
Edinger-Westphal is incorrect as damage to this nucleus would diminish the pupil response both to light and during accommodation.
D. pretectal areas
E. supraoculomotor nucleus
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve
B. abducens nucleus
C. Edinger-Westphal nucleus
D. pretectal areas This answer is CORRECT!
The pretectal area provide bilateral input to the Edinger-Westphal nucleus for the direct and consensual pupillary light response.
E. supraoculomotor nucleus
A patient is capable of pupillary constriction during accommodation but not in response to a light directed to either eye. The lesion is most likely present in the...
A. optic nerve
B. abducens nucleus
C. Edinger-Westphal nucleus
D. pretectal areas
E. supraoculomotor nucleus This answer is INCORRECT.
Supraoculomotor nucleus is incorrect because it is involved in the pupillary accommodation response and not in the pupillary light reflex response.
