1.

Eye-specific patches or stripes normally develop in the visual cortex and superior colliculus of many (but not all) mammals and are also formed, after surgically produced binocular innervation, in the optic tectum of fish and frogs. The segregation of ocular dominance patches or columns has been studied using a variety of anatomical pathway-tracing techniques, by electrophysiological recording of postsynaptic units or field potentials, and by the 2-deoxyglucose method following visual stimulation of only one eye.




2.

In the tectum of both fish and frogs and in the cortex and colliculus of mammals, eye-specific patches develop from initially diffuse, overlapping projections.




3.

Of the various mechanisms that might cause such segregation, the evidence favors an activity-dependent process that stabilizes synapses from the same eye because of their correlated activity. First, several environmental manipulations affect the segregation of afferents in visual cortex: strabismus and alternate monocular exposure apparently enhance segregation, whereas dark rearing slows the segregation process, and monocular deprivation causes the experienced eye to form larger patches at the expense of those of the deprived eye. Second, blocking activity in both eyes is effective in preventing the segregation both in the tectum of fish and frog and in the visual cortex of cat. With the eyes blocked, alternate stimulation of the optic nerves permits the segregation of ocular dominance, at least onto single cells in the cat visual cortex.




4.

These findings are discussed in terms of an activity-dependent stabilization of those synapses having correlated activity (those from neighboring ganglion cells within one eye) but not of those lacking correlated activity (those from left and right eyes). We suggest that the eye-specific patches represent a compromise between total segregation of the projections from the two eyes and the formation of a single continuous retinotopic map across the surface of the cortex or tectum.

Eye-specific segregation of optic afferents in mammals, fish, and frogs: The role of activity

1.

In monocularly deprived (MD) cats, many cells in the lateral geniculate nucleus (LGN) but few cells in the visual cortex respond to input from the deprived eye, suggesting that the connections to visual cortex from the deprived geniculate laminae may have been disrupted.




2.

It has been known for some time that the afferents representing the deprived eye terminate over a smaller percentage of layer IV than do those representing the experienced eye, but it is becoming increasingly clear that this alone cannot explain the inability of the deprived pathway to activate cortical cells.




3.

2-Deoxyglucose studies of ocular dominance columns in MD cats have shown that the columns are often (a) restricted to layer IV, suggesting that intracortical connections may be disrupted, and (b) very faint, suggesting that MD alters the efficacy of the deprived pathway in addition to restricting its territory.




4.

Electron microscopy has shown that both deprived and experienced afferents end in terminals that contain mitochondria and round synaptic vesicles and that make asymmetric contacts with dendritic profiles. However, the terminals of deprived afferents differ from those of experienced afferents: they are 25% smaller, contain 33% fewer mitochondria, are more likely to make synapses that are presynaptically convex (and thus, perhaps, immature), make fewer perforated synapses, and synapse onto smaller spines. Further, the geniculocortical axons from deprived laminae appear to end in fewer synaptic terminals, than do those from the experienced laminae.




5.

The finding that the synaptic terminals of deprived afferents are both abnormal morphologically and fewer in number can help to explain the reduced effectiveness of the deprived eye in driving cortical cells but does not rule out additional effects such as suppression and loss of intracortical connectivity.

The anatomy of geniculocortical connections in monocularly deprived cats.

Goodhill (1993) has recently suggested that the spacing of ocularity domains in visual cortex is not solely an intrinsic property of cortex, but is determined, at least in part, by the degree of correlation in the activity of the two eyes. In support of this model, Lowel (1994) has shown that strabismus, which decorrelates the activity of the two eyes, increases the spacing of ocular dominance columns in area 17, but not area 18, of the cat. As a further test of Goodhill's model, in this paper we examine the effects of another rearing procedure that decorrelates the activity of the two eyes, namely alternating monocular exposure (AME). Cats were reared either normally (9 cats) or with AME (21 cats). We labeled their ocularity domains by one of three methods: ocular dominance columns by 2-deoxyglucose (14 cats), and ocular dominance patches by transneuronal transport (14 cats), or by injections of tracer into single layers of the lateral geniculate nucleus (LGN; 2 cats). The spacing of ocular dominance was 11% greater in the AME cats than in the normal cats (0.976 vs. 0.877 mm). These results are similar to those previously reported for strabismic cats, although the effect is less striking. We thus confirm that decorrelating the activity of the two eyes increases the spacing of cortical ocularity domains. Our results further suggest that the degree of decorrelation affects the extent of that increase.

https://www.cambridge.org/core/services/aop-cambridge-core/content/view/S0952523800011640

Alternating monocular exposure increases the spacing of ocularity domains in area 17 of cats

Quantitative cytoarchitectural analysis of cellular degeneration in the dorsal lateral geniculate nuclei of cats and kittens with cerebral hemispherectomy

Monocular deprivation in kittens differently affects crossed and uncrossed visual pathways

In unequal alternating monocular exposure, each eye receives normal patterned input, but on alternate days and for unequal periods. This imbalance in stimulation produces a behavioral deficit for the less-experienced eye and alters the ability of that eye to activate cortical cells. To determine whether unequal alternating exposure also affects the sizes of cells in the lateral geniculate nucleus (LGN), we measured the cross-sectional areas of geniculate neurons in seven normally reared cats, 14 cats reared with equal alternating exposure, and 17 cats reared with unequal alternating exposure. We found that, in the LGNs of cats reared with unequal alternating monocular exposure, cells in layers that received their input from the less-experienced eye were smaller than those in layers that received their input from the more-experienced eye. This effect was restricted to the binocular segments of the nucleus, and the difference in cell size was a function of the imbalanced exposure, rather than the length of exposure per se. In control groups given balanced alternating exposure, cell size was not correlated with the length of daily exposure. In cats reared with unequal exposure, the change in cell size was greater in the nucleus ipsilateral to the less-experienced eye. Further, the size of the effect was correlated with the size of the imbalance imposed during rearing: Cats reared with a moderate imbalance (8 hours/day vs. 4 hours/day) showed less change in cell size than cats reared with a large imbalance (8 hours/day vs. 1 hour/day). These results are consistent with those of behavorial and physiological studies and strongly suggest (1) that unequal alternating monocular exposure affects the sizes of cells in the LGN by altering the normal competitive balance between the retinogeniculocortical pathways from the left and right eyes, and (2) that the contralateral pathway has some inherent advantage in this competition. We also found a slight shrinkage of cells in the LGNs of cats reared with equal alternating monocular exposure. Since this effect was restricted to the binocular segments of the nucleus, and was not related to the length of exposure given, it was probably caused by the imbalanced binocular competition that occurred during each day's monocular exposure.

N. Tumosa

Papers

Effects of unequal alternating monocular exposure on the sizes of cells in the cat's lateral geniculate nucleus