Reviews
Reflexes and the eye
Alex Paul Hunyor, MB BS*
Abstract
Reflexes are an essential part of protective and
homeostatic function, both in general terms and
with specific reference to ocular structures. A wide
range of stimuli and responses, with varying
degrees of central processing, is involved in such
reflexes.
The simplest reflexes are monosynaptic, such as
the stretch or myotatic reflex. More complex
polysynaptic reflexes are involved in many regula-
tory and protective functions - these include auto-
nomic as well as somatic reflexes. Ocular auto-
nomic reflexes include the oculocardiac, pupillary,
accommodative and lacrimatory reflexes. Ocular
somatic reflexes include eyelid and extra-ocular
muscle reflexes (such as Bell’s phenomenon,
vestibulo-ocular and optokinetic reflexes).
An account of the above reflexes is given in the
format of an essay, modified from the FRACO Part
I Examination in Physiology. The topic was
‘Discuss reflex activities with particular reference to
the eye’. The content is based on several of the
texts recommended for the Part I Examination, as
listed under references.
Key words: Blink, corneal, monosynaptic, poly-
synaptic, pupillary, stretch and vestibulo-ocular
reflexes.
Reflexes are involuntary responses, usually asso-
ciated with protective or regulatory functions in
the organism in which they occur. The following
From the Royal Victorian Eye and Ear Hospital, 32 Gisbome Street, East Melbourne, victoria 3002.
*Ophthalmology Registrar.
Reprint: Dr Alex Paul Hunyor.
Reflexes and the eye
is a discussion of the general physiological princi-
ples of reflex actions and an overview of the
specific reflexes pertaining to the eye and its
surrounding structures.
At the most basic level, reflexes require a
receptor (or sense organ), an afferent neuron,
one or more synapses in a central integrating
station (brain or spinal cord) or sympathetic
ganglion, an efferent neuron and an effector.
Receptors may be sensitive to a wide range of
stimuli including mechanical, thermal, nocicep-
tive, proprioceptive, pressure and chemical
stimuli. Effectors may result in motor, sympa-
thetic, parasympathetic, chemical and hormonal
responses. Afferent neurons enter via dorsal roots
or cranial nerves, have their cell bodies in dorsal
root or cranial nerve ganglia, and synapse in the
central nervous system (CNS), with their efferent
fibres leaving in the ventral roots or motor cranial
nerves.
Monosynaptic reflexes
The simplest reflexes are those with single
afferent and efferent neurons and a single
synapse (monosynaptic reflexes). The minimum
delay across one synapse is 0.5 msec. The stretch
or myotatic reflex is a typical monosynaptic
reflex. The stimulus is stretching of the muscle,
the receptor or sense organ is the muscle spindle
(composed of intrafusal muscle fibres), and the
response (reflex muscle contraction) is mediated
by the extrahsal muscle fibres (see Figure 1).
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Extrafusal
muscle fibre
Figure I Monosynaptic and polysynaptic reflexes. Stretch
reflex (solid afferent pathway) and inverse stretch reflex
(dotted afferent pathway).
Muscle spindles consist of three to 12 small
muscle fibres encased in a connective tissue
capsule which is connected to the glycocalyx of
the surrounding extrafusal skeletal muscle fibres
(which make up the vast bulk of the muscle).
They are thus in parallel with the extrafusal
fibres, with their central (essentially non-
contractile) segments acting as sensory receptors
and their end portions contracting in response to
stimulation of gamma efferent fibres (as opposed
to the alpha efferents which innervate the extra-
fusal muscle fibres). Thus the spindle’s central
portion can be stimulated either by contraction
of the whole muscle, or by contraction of the
end-portions of the intrafusal fibre alone.
Intrafusal fibres can be divided into nuclear
bag fibres (larger, more centrally placed and with
many nuclei) and nuclear chain fibres (thinner
and peripherally located). The muscle spindles
contain two types of sensory endings. The
primary or annulospiral ending encircles the
central portion of the intrafusal fibre and is inner-
vated by a type l a neuron with average diameter
17 km and conduction velocity 70 to 120 msec.
The primary ending innervates both nuclear bag
and nuclear chain fibres. The secondary or flow-
erspray ending innervates the intrafusal fibre
receptor region to one side of the primary ending,
and is innervated by a type I1 fibre with average
diameter 8 pm and conduction velocity 30 to
70 msec.
The static response of the spindle receptor is
such that the discharge of the primary and
secondary endings increases in proportion to the
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degree of stretch of the nuclear chain fibres. The
dynamic response to the rate of change of
receptor length is mediated by the nuclear bag
fibres and the primary (but not secondary) fibres.
The muscle stretch reflex is thus divided into the
dynamic stretch reflex, which causes rapid,
forceful reflex contraction of the stretched
muscle; and the weaker static stretch reflex which
operates over a prolonged timecourse, for as long
as the muscle remains stretched. The knee jerk is
a clinical example of the dynamic stretch reflex.
It has a central delay (time taken to traverse the
spinal cord) of 0.6 to 0.9 msec. When a muscle is
suddenly shortened, dynamic and static reflex
inhibition occurs; this is called the negative stretch
reflex.
Responses to changes in muscle tension (as
opposed to muscle length) are mediated via the
Gdlgi tendon organ (GTO), which is contained
in the muscle tendon, is in series with 10 to 15
muscle fibres, and is innervated by type l a affer-
ents. Like the muscle spindle, the G T O has both
a dynamic and static response to changes in
muscle tension. The GTO is the receptor for the
inverse stretch reflex (a polysynaptic reflex): up to
a certain level, as muscle stretch increases, so
does reflex contraction; beyond this, the muscle
relaxes (Figure 1).
Polysynaptic reflexes
Polysynaptic reflexes involve varying numbers of
interneurons and thus increased synaptic delays.
An example is the flexor or withdrawal reflex,
where noxious stimulation of a limb results in
protective reflex withdrawal via activation of
flexor muscles of that limb. Some 0.2 to 0.5
seconds after ipsilateral limb flexion there is
extension of the opposite limb, the crossed extensor
reflex. The numerous reflexes that subserve regu-
latory and protective functions (including those
discussed below) are also clearly polysynaptic
reflexes.
Autonomic reflexes
The responses of the autonomic nervous system,
namely the sympathetic (typically concerned
with reactions to stress) and the parasympathetic
(more associated with restorative and vegetative
functions) usually have opposing actions which
are reciprocal rather than antagonistic in main-
Australian and New Zealand Journal of Ophthalmology 1994; 22(3)
 Pretectal
’ optic nerve
Figure 2 Schematic diagram of the pathway for the pupillary light reflex.
taining homeostasis. Autonomic reflex control
occurs at different levels.
1. Local reflexes such as bladder contraction,
where pelvic sensory nerves form the afferent
limb, relaying in the spinal cord at S2-S4 with
parasympathetic preganglionic fibres, which in
turn synapse in pelvic parasympathetic ganglia
with postganglionic fibres which innervate the
detrusor muscle.
2. Brainstem reflexes such as the pupillary reflex
(see below) and vital reflexes, e.g., vagal afferents
from the lungs influence the output from pontine
respiratory centres (pneumotaxic and apneustic
centres).
3 . Higher control of autonomic activity by the
hypothalamus includes integration with
endocrine and immune systems and higher cere-
bral functions, and maintenance of chemical,
thermal and nutritional homeostasis.
Autonomic reflexes and the eye
Oculocardiac rejex
This reflex occurs with pressure on the eye or
orbit, or stretching of extraocular muscles, as
may occur during ophthalmic surgery. Stimuli
such as pain or pressure are transmitted via affer-
ents in the nasociliary branch of the ophthalmic
Reflexes and the eye
Edinger-Westphal nucleus
Medial geniculate body
nerve, to the trigeminal nerve, thence to the
trigeminal spinal nucleus which projects to the
reticular formation in the area of the visceral
motor nuclei of the vagus nerve, and vagal
efferent outflow results in nausea, bradycardia
and faintness (via muscarinic cholinergic recep-
tors). Syncope and even cardiac arrest have been
described with this reflex, which may be
prevented by prior muscarinic blocking (with
intravenous atropine) or by retrobulbar or
peribulbar anaesthesia. Thus, it may still occur
during general anaesthesia.
Pupillary reflexes
The main pupillary reflexes are the light and dark
reflexes (which share a common afferent
pathway) and miosis associated with the near
triad or synkinesis. The pupil has opposing
dilator and sphincter muscles, innervated
predominantly by sympathetic and parasympa-
thetic fibres respectively.
The pathway for the light reflex (Figure 2) is
from the retina via optic nerve, hemi-decussating
in the optic chiasm, to the optic tract, leaving in
its proximal third, and passing between the
medial and lateral geniculate bodies, via the
brachium of the superior colliculus to the
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pretectal nuclei. There is a further hemidecussa-
tion via the posterior commissure to project to
both Edinger-Westphal (E-W) nuclei (part of the
oculomotor nuclear complex of the midbrain)
whose preganglionic fibres travel via the nasocil-
iary nerve to synapse in the ciliary ganglion, with
myelinated postganglionic fibres travelling in the
short ciliary nerves to innervate the sphincter
pupillae via M3 muscarinic receptors. There is
weak beta-adrenergic inhibition of the sphincter
pupillae. The bilateral projection of the light
reflex pathway produces both direct and consen-
sual pupillary constriction, and asymmetry of
afferent input, due to retinal or optic nerve
disease, is manifest clinically as a relative afferent
pupillary defect.
The sympathetic pathway to the dilator
pupillae has a similar afferent limb - the central
outflow is from the ventrolateral hypothalamus
(and in areas adjacent to the E-W nucleus), via
the brainstem and spinal cord to the ciliospinal
centre of Budge in the intermediolateral tract at
C8-T2, synapsing with the preganglionic neuron
which passes in the sympathetic chain to synapse
in the superior cervical ganglion, whence the
postganglionic efferents pass to the carotid
plexus and join the nasociliary branch of the
ophthalmic nerve and reach the dilator pupillae
(predominantly) via the long posterior ciliary
nerves. There is also weak cholinergic inhibition
of the dilator pupillae.
The dark reflex involves both a passive return of
the pupil to its relaxed state, and active supranu-
clear inhibition of the tonically active E-W nucleus.
There may also be reflex pupillary dilatation as part
of a generalised sympathetic response to physical
and psychosensory stimuli.
Accommodation and the near triad
The near triad of bilateral miosis, accommoda-
tion and convergence to near visual targets is a
reflex which enables particularly rapid response
to approaching or nearby objects. Miosis will
occur with accommodation in the absence of
convergence, and vice versa. The stimuli to
accommodation are blur, chromatic aberration,
proximity and convergence. In youth, conver-
gence and accommodation match in a 1:1 ratio.
Thus, a stimulus to convergence alone (i.e.,
diplopia) will be adequate to induce the correct
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amount of accommodation for distinct vision.
After the age of 24, accommodation lags behind
Convergence.
The afferent limb for accommodation is the
retinogeniculocortical pathway. The exact
central mechanism for accommodation is
unclear, but unilateral stimulation of certain
peristriate areas of cortex will result in bilateral
accommodation, miosis and convergence. Para-
sympathetic neurons, following a similar path-
way to that for miosis, have the predominant
influence over the ciliary muscle. Those
subserving accommodation are located rostrally
in the E-W nucleus with respect to pupillomotor
neurons. Cortical output to the E-W nucleus is
subject to hypothalamic autonomic influence.
Only 1% of neurons innervating the ciliary
muscle are sympathetic, resulting in weak beta-1
inhibition. Convergence is mediated by the
medial rectus subnucleus (probably subnucleus
‘C’) of the oculomotor complex of the midbrain.
Reflex lacrimation
Reflex tear secretion may be in response to
various stimuli, including: physical and chemical
stimuli to the cornea, conjunctiva and nasal
mucosa; bright light; and in association with
emotional upset, vomiting, coughing and
yawning. Basal tear secretion is thought to be
mediated by sympathetic regulation of blood
flow to the lacrimal gland, while reflex lacrima-
tion is predominantly via parasympathetic
secretomotor action on the acini and ducts of the
lacrimal gland. The afferent pathway (in the case
of local stimuli) is via the sensory branches of the
trigeminal nerve (as detailed above).
Parasympathetic outflow is from the lacrimal
nucleus of the pons, via the n e w s intermedius
through the geniculate ganglion, as the greater
superficial petrosal nerve, joining the deep
petrosal nerve to form the Vidian nerve, and
synapsing in the pterygopalatine ganglion.
Postganglionic fibres join the zygomatic nerve
and reach the lacrimal gland via its lacrimal
branch.
Somatic reflexes and the eye
Eyelid reflexes
Closure of the eyelids may be voluntary, by spon-
taneous blinking, or as a protective reflex
Australian and New Zealand Journal of Ophthalmology 1994; 22(3)
response to tactile, visual and auditory stimuli.
The efferent limb in all cases is the facial nerve.
The corneal rejfex is a response to tactile stimuli
on the cornea, with afferents via trigeminal nerve
branches. Similar stimulation of the eyelids,
eyelashes and conjunctiva can also produce reflex
blinking. Visual stimuli can produce: the dazzle
reflex, a response to shining a bright light into the
eye, which occurs at a subcortical level; and the
menace reflex, in response to a threatening object
suddenly entering the visual field, which requires
occipital cortical connections. Auditory stimuli
(sudden loud noises), and stretching or striking
of structures near the orbit, may also cause reflex
blinking.
Extraocular muscle reflexes
Bell’s phenomenon is the protective reflex upward
movement of the globe that occurs with forcible
eyelid closure (during blinking, eyes tend to
move towards the primary position). T h e
pathway for this phenomenon is not known, but
it differs from that for voluntary upward gaze, as
it may still occur in the presence of supranuclear
gaze palsy. It is absent in 10% of normal individ-
uals.
The vestibulo-ocular rejlex (VOR) functions to
maintain steady gaze during head rotation. Static
VOR maintains a normally orientated visual field
despite changes in head position. Head tilting,
maintained postural deviation or rectilinear
acceleration of the head will result in slow,
incomplete eye movement in the direction oppo-
site to that of head movement. The utricle and
saccule of the vestibular complex mediate the
static VOR, which forms the basis of
Bielschowsky’s head tilting test. Dynamic VOR is
Reflexes and the eye
a response to rotational acceleration, maintaining
steady eye position in space despite head rota-
tion. The semicircular canals are the sense organs
for this response, which results in eye movement
equal and opposite to the direction of head rota-
tion. For large angles, slow compensatory eye
movements are interrupted by fast movements in
the direction of rotation (vestibular nystagmus) .
By Flouren’s law, each semicircular canal gives
rise to nystagmus in the plane of that canal.
The optokinetic system attempts to maintain a
stable retinal image so that the perception of the
stationary world remains clear despite movement
of the head or the object of interest. The stimulus
is full-field retinal slip (there are both central and
peripheral components, in distinction to simple
foveal smooth pursuit) , and optokinezic nystagmus
(OKN) consists of alternating slow (compen-
satory) movements in the direction of object
movement, and rapid (anticompensatory) move-
ments in the opposite direction. OKN is medi-
ated by the geniculo-transcortical-floccular
pathway, and the presence of OKN can be used
as a test of visual function in infants and illiterate
patients.
References
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Macmillan Press, 1990.
Duane TD, Jaeger EA. Biomedical foundations of ophthal-
mology. Vol 2. Physiology of the eye and visual system.
Philadelphia: Harper and Row, 1982.
Ganong WF. Review of medical physiology. 15th ed. London:
Prentice-Hall International, 1989.
Guyton AC. Textbook of medical physiology. 7th ed.
Philadelphia: WB Saunders, 1986.
Hart WM. Adler’s physiology of the eye. 9th ed. St Louis: CV
Mosby, 1992.
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