Ординатура / Офтальмология / Английские материалы / The Neurology of Eye Movements_Leigh, Zee_2006
.pdf190 The Properties and Neural Substrate of Eye Movements
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Chapter 05
GAZE HOLDING AND THE
NEURAL INTEGRATOR
NEURAL CODING OF THE OCULAR MOTOR SIGNAL
The Need for a Neural Integrator of Ocular Motor Signals
Special Demands on the Neural Integrator QUANTITATIVE ASPECTS OF NEURAL
INTEGRATION
NEURAL SUBSTRATE FOR GAZE HOLDING
Contribution of the Nucleus Prepositus Hypoglossi and Medial Vestibular Nucleus to Gaze Holding
The Interstitial Nucleus of Cajal and Vertical Gaze Holding
Contribution of the Cerebellum to Gaze Holding
How a Network of Neurons Could Function as the Neural Integrator
CLINICAL EVALUATIONOF GAZE
HOLDING
ABNORMALITIES OF THE NEURAL INTEGRATOR
Pathogenesis of Deficient Neural Integration
Pathogenesis of Centripetal Nystagmus and Rebound Nystagmus
SUMMARY
This chapter deals with the neural mechanism that holds gaze steady when the eyes are turned away from the central position. Clinicians traditionally test the stability of gaze with the patient's eyes at the limits of the horizontal or vertical range. Holding
the eyes steady under such circumstances calls upon more than visual fixation (discussed in Chap. 4), since eccentric gaze remains relatively steady in darkness.5'74 In Chapter 1, we pointed out that the orbital contents impose elastic restoring forces that tend to pull the eyes back to central position. To counteract these forces and hold the eyes steady in an eccentric position in the orbit, the extraocular muscles must contract tonically. Such a tonic contraction is achieved by a sustained rate of discharge of the ocular motoneurons.
The mechanical forces that act on the eye are illustrated in the experiment shown in Figure 5-1.76 The subject viewed a stationary visual target with one eye while vision from the other eye was occluded with a sheet of opaque paper at a distance of about 5 cm. After applying topical anesthetic to the nonfixating eye, it was mechanically displaced, using ophthalmic forceps, into eccentric positions of (^4) intorsion, (B) extorsion, or (C) horizontal abduction. After the eye was suddenly released from each of these eccentric positions, it sprang back to a "central" position of rest. The time course of this return was determined by the mechanical forces acting on the eye, which differed according to the prior direction in which it had been displaced. The brain must take into account these mechanical forces in programing all types of eye movements.
Our approach in this chapter will be first, to explore what neural signals the oc-
198
Gaze Holding and the Neural Integrator |
1 9 9 |
Figure 5-1. Experimental data from a normal human subject to show the time course of the return to resting position after the eye was mechanically displaced into an eccentric position in the orbit and then suddenly released. If the time course of return were fit by a single exponential function, the time constants would be (A) 323 msec after release from intorsion; (B) 58 msec after release from extorsion; and (C) 183 msec after release from abduction. The asterisk in C indicates a blink. (Adapted from Vision Research, volume 35, Seidman SH, Leigh RJ, Tomsak RL, Grant MP, Dell'Osso LF. Dynamic properties of the human vestibulo-ocular reflex during head rotations in roll, page 679-89, 1995, with permission from Elsevier Science.)
ular motoneurons must generate to hold the eye in an eccentric position; second, to outline quantitative aspects of this gaze holding function; third, to identify what anatomical pathways are important for normal gaze holding; fourth, to apply these principles to the clinical examination; and finally, to review clinical disorders that impair the ability to hold steady, eccentric gaze.
NEURAL CODING OF THE
OCULAR MOTOR SIGNAL
The Need for a Neural Integrator of Ocular Motor Signals
To understand the neural basis for the gaze-holding mechanism, it is helpful to consider the way that brain stem neurons