Binocular Segregation in the Dorsal Lateral Geniculate Nucleus

ABSTRACT In most mammals the optic nerve fibres from the two eyes terminate in separate layers in the dorsal lateral geniculate nucleus (LGNd). The layers of retinal input in the LGNd can be revealed by transneuronal degeneration of cells in the LGNd or anterograde optic fibre degeneration following removal of one eye, by transport of radioactive amino acids to the optic terminals in the LGNd or by electrophysiological recording of single neurons in the LGNd. The laminar organisation of the LGNd is described in man, rhesus monkey, cat and some Australian marsupials. In Australian possums and kangaroos the optic fibres from the two eyes terminate in separate layers in the LGNd whereas in the Australian carnivorous marsupials there is significant overlap of optic fibres from the two eyes in the LGNd. The development of the normal laminar organisation of the LGNd is described for a number of species. Early in development there is little evidence of lamination in the LGNd and optic fibres from both eyes spread over the whole LGNd. The adult pattern of lamination generally appears at birth or shortly afterwards. Binocular interaction in the LGNd is described in the cat, where it arises both from intrageniculate circuits and via binocular cortico-geniculate fibres. It is suggested that the function of lamination in the LGNd is to provide separate channels for some of the different types of information brought in from the retina.     

X- and Y-mediated synaptic potentials in neurons of areas 17 and 18 of cat visual cortex

When a cuff-shaped electrode is placed on the optic nerve of the cat, X and Y axons, by virtue of their different diameters, exhibit different thresholds to electrical stimulation. Large-diameter Y axons have low thresholds, while smaller-diameter X axons have high thresholds. There is very little overlap between the two populations. Given this segregation, the strength of stimulation of the optic nerve required to evoke synaptic potentials in cortical neurons becomes a reliable indicator of the type of visual input a cortical neuron receives. Potentials with thresholds below the thresholds of X axons must be mediated by Y cells of the retina and LGN. Potentials with thresholds above the Y axons of the optic nerve must be mediated by X cells. From previous experiments, one would expect to find ample input via both types of axon to area 17 of the visual cortex. This was not the case. Of 58 neurons distributed throughout the layers of area 17 from which intracellular records were taken, in only four could substantial Y excitation be detected. Three of these four were located near the border with area 18. All four received large X inputs as well. The 24 neurons studied in area 18 all received large Y inputs but no detectable X input.     

Retinal projections in the ground squirrel (Citellus tridecemlineatus)

The retinal projections of the thirteen-lined ground squirrel were determined by tracing anterograde transport of intravitreally injected horseradish peroxidase (HRP) or wheat-germ conjugated horseradish peroxidase (WGA-HRP). Label was seen in the suprachiasmatic nucleus and adjacent anterior hypothalamic area, the accessory optic system (the medial, dorsal, and lateral terminal nuclei), the dorsal and ventral lateral geniculate nuclei, the intergeniculate leaflet, the pretectal nuclei (the anterior, posterior, and olivary pretectal nuclei and the nucleus of optic tract), and the superior colliculus. Most of these structures were labeled bilaterally, with dense contralateral label and sparse ipsilateral label, a pattern typical for animals with laterally placed eyes. However, the suprachiasmatic nucleus and the nucleus of the optic tract received input only from the contralateral eye. In contrast to previous degeneration studies, the sensitive HRP tracers (in conjunction with cytochrome-oxidase reactivity) revealed an elaborate organization within the lateral geniculate nucleus (dorsal LGN, ventral LGN, and intergeniculate leaflet) that is consistent with existing organizational schemes for other mammalian species.     

Cat-301 antibody selectively labels neurons in the Y-innervated laminae of the cat superior colliculus

Cat-301 is a monoclonal antibody which recognizes a cell surface associated antigen of selected neurons in the central nervous system (CNS). In the visual system, cat-301 selectively labels Y-like cells in several visual structures, including portions of the lateral geniculate nucleus complex and visual cortex. The cat superior colliculus (SC) also receives Y input and contains cells driven by Y input which are selectively distributed in the deep superficial gray and deeper laminae. If cat-301 is selective to the Y-cell system in SC, labeled cells should be restricted to those laminae. To test this hypothesis, we have examined quantitatively the laminar distribution, percentage, size, and morphology of cells in SC labeled by the cat-301 antibody. Cat-301 labeled a variety of cells in the cat SC. Labeled cells were found within the deep portion of the superficial gray layer (6.6%), optic layer (27.6%), intermediate gray layer (26.9%), and the deep gray and white layers (38.5%). By contrast, only 2 of 667 labeled cells (0.3%) were found within that part of the upper superficial gray layer innervated exclusively by W input and thought to contain only W-driven cells. When considered as a percentage of the total cell population, cat-301 labeled cells represented less than 3% of cells in the superficial gray layer and approximately 15% in the deeper layers. Neurons labeled by cat-301 were all of medium to large size (mean average diameter = 33.3 microns; range = 15-84 microns) and included vertical fusiform and stellate cells in the upper layers and the very large neurons found in the intermediate gray and deeper layers. These results provide further evidence that the cat-301 antibody selectively recognizes the Y channel of the cat visual system.     

First order connections of the visual sector of the thalamic reticular nucleus in marmoset monkeys (Callithrix jacchus)

The thalamic reticular nucleus (TRN) supplies an important inhibitory input to the dorsal thalamus. Previous studies in non-primate mammals have suggested that the visual sector of the TRN has a lateral division, which has connections with first-order (primary) sensory thalamic and cortical areas, and a medial division, which has connections with higher-order (association) thalamic and cortical areas. However, the question whether the primate TRN is segregated in the same manner is controversial. Here, we investigated the connections of the TRN in a New World primate, the marmoset (Callithrix jacchus). The topography of labeled cells and terminals was analyzed following iontophoretic injections of tracers into the primary visual cortex (V1) or the dorsal lateral geniculate nucleus (LGNd). The results show that rostroventral TRN, adjacent to the LGNd, is primarily connected with primary visual areas, while the most caudal parts of the TRN are associated with higher order visual thalamic areas. A small region of the TRN near the caudal pole of the LGNd (foveal representation) contains connections where first (lateral TRN) and higher order visual areas (medial TRN) overlap. Reciprocal connections between LGNd and TRN are topographically organized, so that a series of rostrocaudal injections within the LGNd labeled cells and terminals in the TRN in a pattern shaped like rostrocaudal overlapping "fish scales." We propose that the dorsal areas of the TRN, adjacent to the top of the LGNd, represent the lower visual field (connected with medial LGNd), and the more ventral parts of the TRN contain a map representing the upper visual field (connected with lateral LGNd).     

Changes in the receptive fields of cat retinal ganglion cells affected by pressure on the optic nerve

In the cat optic nerve a lesion was induced by brief application of pressure. It selectively blocked impulse conduction in large diameter fibres of the retinal ganglion cells. Electrophysiological examination of single optic axons several weeks later demonstrated a gross alteration of the visual properties of the affected BT/Y ganglion cells. It is suggested that the alteration of receptive field properties may reflect the cellular and dendritic response to distant focal injury of the axon.     

Abnormal retinogeniculate projections in the congenitally microphthalmic cat

The retinogeniculate projections from the normal eye of a unilaterally microphthalmic cat are abnormal in that optic tract fibers cross laminar borders and end, inappropriately, in geniculate layers that would normally receive input from the microphthalmic eye. This congenitally induced abnormal retinogeniculate projection is quite similar to that seen in cats with one eye surgically removed shortly after birth. Although most cells are shrunken in the laminae normally innervated by the microphathalmic eye, cells in the region of the abnormal projection appear normal. The normal pattern of geniculate lamination is also disrupted in that cell-free interlaminar regions are considerably more difficult to define in the microphthalmic cat.     

Effects of selective pressure block of Y-type optic nerve fibers on the receptive-field properties of neurons in the striate cortex of the cat.

In an aseptic operation under surgical anesthesia, one optic nerve of a cat was exposed and subjected to pressure by means of a special cuff. The conduction of impulses through the pressurized region was monitored by means of electrodes which remained in the animal after the operation. The pressure was adjusted to selectively eliminate conduction in the largest fibers (Y-type) but not in the medium-size fibers (X-type). The conduction block is probably due to a demyelination and remains complete for about 3 weeks. Within 2 weeks after the pressure-block operation, recordings were made from single neurons in the striate cortex (area 17, area V1) of the cat anesthetized with N2O/O2 mixture supplemented by continuous intravenous infusion of barbiturate. Neurons were activated visually via the normal eye and via the eye with the pressure-blocked optic nerve ("Y-blocked eye"). Several properties of the receptive fields of single neurons in area 17 such as S (simple) or C (complex) type of receptive-field organization, size of discharge fields, orientation tuning, direction-selectivity indices, and end-zone inhibition appear to be unaffected by removal of the Y-type input. On the other hand, the peak discharge rates to stimuli presented via the Y-blocked eye were significantly lower than those to stimuli presented via the normal eye. As a result, the eye-dominance histogram was shifted markedly towards the normal eye implying that there is a significant excitatory Y-type input to area 17. In a substantial proportion of area 17 neurons, this input converges onto the cells which receive also non-Y-type inputs. In one respect, velocity sensitivity, removal of the Y input had a weak but significant effect. In particular, C (but not S) cells when activated via the normal eye responded optimally at slightly higher stimulus velocities than when activated via the Y-blocked eye. These results suggest that the Y input makes a distinct contribution to velocity sensitivity in area 17 but only in C-type neurons. Overall, our results lead us to the conclusion that the Y-type input to the striate cortex of the cat makes a significant contribution to the strength of the excitatory response of many neurons in this area. However, the contributions of Y-type input to the mechanism(s) underlying many of the receptive-field properties of neurons in this area are not distinguishable from those of the non-Y-type visual inputs.     

Lagged Y cells in the cat lateral geniculate nucleus.

We report on the existence of lagged Y (YL) cells in the A laminae of the cat lateral geniculate nucleus (LGN) and on criteria for identifying them using visual and electrical stimulation. Like the lagged X (XL) cells described previously (Mastronarde, 1987a; Humphrey & Weller, 1988a), YL cells responded to a spot stimulus with an initial dip in firing and a delayed latency to discharge after spot onset, and an anomalously prolonged firing after spot offset. However, the cells received excitatory input from retinal Y rather than X afferents, and showed nonlinear spatial summation and other Y-like receptive-field properties. Three YL cells tested for antidromic activation from visual cortex were found to be relay cells, with long conduction latencies similar to those of XL cells. Simultaneous recordings of a YL cell and its retinal Y afferents show striking parallels between lagged X and Y cells in retinogeniculate functional connectivity, and suggest that the YL-cell response profile reflects inhibitory processes occurring within the LGN. The YL cells comprised approximately 5% of Y cells and approximately 1% of all cells in the A laminae. Although infrequently encountered in the LGN, they may be roughly as numerous as Y cells in the retina, and hence could fulfill an important role in vision.     

Prenatal disruption of binocular interactions creates novel lamination in the cat's lateral geniculate nucleus

The elimination of retinogeniculate afferents from one eye on embryonic day 44 (E44) has pronounced effects on the formation of the cellular laminae in the cat lateral geniculate nucleus (LGN). Only two laminae form: a dorsal, "magnocellular" layer, and a ventral, "parvocellular" layer. Soma size measurements and previously reported patterns of termination of retinogeniculate axons suggest that the dorsal lamina is a coalescence of the normal A-laminae and the dorsal, magnocellular division of layer C, while the ventral layer is a composite of the parvocellular sublamina of layer C and the remaining C-laminae. This is a novel pattern of lamination in the LGN that differs from that found in the normal nucleus, not only in that there are now only two cell layers rather than the normal five, but also in that the interlaminar zone occurs in an abnormal location. This result is markedly different from that observed in other species where interlaminar zones present after early monocular enucleation are a subset of the ones which would normally be present. We suggest that, in the absence of ongoing binocular interactions, interactions between functionally distinct retinal ganglion cell classes from the remaining eye may direct the formation of cell laminae in the LGN, even when such interactions are not normally operative.     

Atrophy of myelinated nerve fibres in the retina in optic neuritis.

Atrophy of the nerve fibres in the retina visible by direct ophthalmoscopy using red-free illumination, together with corresponding scotomata in the visual fields, signified retrograde degeneration of retinal nerve fibres from multiple sclerosis. Defects in medullated retinal nerve fibres were continous with arcuate nerve fibre bundle defects. These myelin defects exemplify secondary demyelination of optic axons.     

Acrylamide effects on the macaque visual system. II. Retinogeniculate morphology

Oral acrylamide dosing for 6-10 weeks produced axonal swellings with neurofilament accumulation in the distal optic tract and lateral geniculate nucleus of macaques. No swellings were seen in the retina or optic nerve. Monkeys that were killed 6-8 months after similar dosing showed a marked neuronal degeneration in the visual pathways that was more pronounced after two than after a single period of exposure. This degeneration was characterized by the following: loss of ganglion cells in central retina with relative sparing of other retinal neurons; disproportionate degeneration of temporal to central optic nerve and the dorsal optic tract; and neuronal atrophy in parvocellular layers of the lateral geniculate nucleus, with relative sparing of magnocellular layers. The pattern of neuronal loss suggests that one type of retinal ganglion cell or its axon may be especially vulnerable to damage by acrylamide. The selective neuronal damage produced by acrylamide may help explain the nature of the visual dysfunction associated with this intoxication.     

Distinct central representations for sensory fibers innervating either the conjunctiva or cornea of the rat

The laminar sheet of epithelium (e.g., skin and mucous membrane) enclosing our bodies is represented in the dorsal horns of the medulla and spinal cord. The eyeball however indents this laminar sheet and is shrouded by different layers: the cornea/sclera, the conjunctiva, and hairy skin. This involution of the orb confounds defining the central representation of the cornea and its surrounding mucosa and skin. We used herein the transganglionic transport of a cocktail of HRP conjugated to cholera toxin and wheat germ agglutinin to determine the central representation of these epithelia in the dorsal horns of the rat. The HRP cocktail was injected either into the stroma of the cornea, the mucosa of the conjunctiva, or the supraorbital and infraorbital nerves. Injections of the cornea produced dense label in the interstitial islands in the ventral medullary dorsal horn (MDH), probably lamina I, and in neuropil in the ventromedial tip of the MDH, probably lamina II. There sometimes was variable, diffuse label in the C1 dorsal horn after corneal injections but more rostral parts of the trigeminal sensory complex were never labeled. Injections of the conjunctiva densely labeled laminae I-III in the C1 dorsal horn, while laminae IV-V were diffusely labeled. Sparser reaction product also was seen in lamina I in positions similar to the cornea projection. Label was seen ventrally in subnuclei interpolaris and oralis, as well as the principal trigeminal nucleus. Projections of the infraorbital nerve included all laminae in the trigeminocervical complex as well as large portions of the rostral subnuclei in the spinal trigeminal nucleus. The projections of the supraorbital nerve were similar, but were restricted to ventral parts of the trigeminal sensory complex. In other cases the cornea was injected either after cutting the supraorbital and infraorbital nerves or the conjunctiva was injected after enucleating the eyeball. Any reaction product from corneal injections was reduced dramatically in the C1 dorsal horn after transection of the infraorbital and supraorbital nerves. Injecting the conjunctiva after enucleating the eyeball densely labeled the C1 projection to the dorsal horn, a small patch in lamina I in the MDH, as well as the rostral trigeminal complex. We propose that the cornea has but a single representation in the trigeminocervical complex in its ventral part near the caudal end of the medulla. We also propose the palpebral conjunctiva mucosa is represented in the C1 dorsal horn, and speculate that the bulbar conjunctiva overlaps with that of the cornea in lamina I. We discuss these projections in relation to the circuitry for the supraorbital-evoked and corneal-evoked blink reflexes. The relationship of the cornea and conjunctiva is intimate, and investigators must be very careful when attempting to stimulate them in isolation.     

X- and Y-mediated current sources in areas 17 and 18 of cat visual cortex

X- and Y-mediated input to areas 17 and 18 of the cat visual cortex was studied using current-source-density analysis of field potentials evoked by stimulation of the optic nerves. A cuff-shaped electrode was used for stimulation so that Y axons, by virtue of their larger diameters, would have lower electrical thresholds than X axons. The effect in each cortical area of activating Y axons alone could therefore be determined by low-amplitude stimulation of the optic nerves. Current-source densities were calculated by two separate methods. (1) In five experiments, field potentials were measured sequentially at different cortical depths with a single tungsten electrode. Current densities were then calculated by computer. (2) In two experiments, current densities were derived in real time from field potentials recorded simultaneously from three sites with a multi-electrode probe. The calculation was performed by an analog circuit specially designed for this purpose. This method has several advantages over the standard, single-electrode method. At stimulus strengths sufficient to activate the majority of Y axons in the optic nerves, but subthreshold to most X axons, the field potentials evoked in area 17 changed little from layer to layer. When the current-source-density analysis was applied to these potentials, no significant sources or sinks were detectable. Only when the stimulus strength was raised to the point that both X and Y axons were activated by the stimulus were any current sources or sinks detected in area 17. The currents were similar in time course and laminar pattern to those recorded after stimulation of the optic chiasm. In area 18, large sources and sinks were evoked by stimulation of Y axons alone. These currents changed little when the stimulus strength was increased to activate X axons as well. Area 18, therefore, in contrast to area 17, seems to be dominated by Y input and receives little X input. These results support the conclusions of the accompanying paper in which synaptic potentials were recorded intracellularly from cortical neutrons. The intracellular experiments failed to show substantial Y input to area 17. The projections of X and Y axons may therefore be much more highly segregated into areas 17 and 18 than previously thought. Alternatively, the nature of the Y input to area 17 may be very different from that to area 18 in that it cannot be easily detected with intracellular or current-source-density techniques.     

Specific transcellular staining of microglia in the adult rat after traumatic degeneration of carbocyanine-filled retinal ganglion cells.

The present work was undertaken to assess the fate of ganglion cell debris in the axotomized retina of adult rats and employed a new technique to label phagocytosing microglia via the internalized material. In the main experiment, transection axotomy was performed on the intraorbital segment of the optic nerve, and a fast-transported, vital fluorescent styryl dye (4Di-10ASP) was deposited at the ocular stump of the nerve in order to pre-label retrogradely the ganglion cells destined to die because of the axotomy. Optic nerve transection resulted in progressive degradation of ganglion cell axons, perikarya, and dendrites within the retina and in release of fluorescent material, which was then incorporated into cells identified as microglia. No other retinal cells stained, although astrocytes and Müller's cells also responded to neuron degeneration by accumulating glial fibrillary acidic protein. Incorporation of labelled material into microglia topo-chronologically paralleled the ganglion cell degeneration starting within the optic fibre layer (OFL) and proceeding towards the ganglion cell layer (GCL) and the inner plexiform layer (IPL) of the affected retina. Long-term labelling of microglia monitored up to 3 months after optic nerve transection indicated that labelled microglial cells persisted within the retina. Microglia displayed a strong territorial arrangement within the GCL and IPL, and staggered, bilaminated distribution in both layers. These studies directly prove that microglia in the retina can be transcellularly labelled during traumatic degeneration of ganglion cells. The findings suggest that microglial cells play an important role in axotomy-induced wound healing and removal of cell debris.     

Organization of reciprocal connections between the perigeniculate nucleus and dorsal lateral geniculate nucleus in the cat: a transneuronal transport study

Cells of the cat's perigeniculate nucleus (PGN), part of the visual sector of the thalamic reticular nucleus (TRN), provide GABAergic inhibition to the A and A1 layers of the dorsal lateral geniculate nucleus (LGNd) and, therefore, may control information flow from the retina to the cortex. Previous electrophysiological experiments suggested that the PGN may be subdivided on the basis of ocular dominance thus reflecting the afferent and efferent projections with lamina A and A1 of the LGNd. The present study utilized the ability of wheat germ agglutinin-horseradish peroxidase (WGA-HRP) to be transported transneuronally following intraocular injections in four cats to examine whether there is any anatomical evidence for eye specific layers within the PGN. Sections were processed with tetramethylbenzidine. Light WGA-HRP transneuronal labeling of LGNd collaterals and somata were seen in the PGN and very light labeling (but not somata) was seen in the TRN. Neither the cells of the PGN projecting to the LGNd nor the LGNd relay collaterals within the PGN were clearly organized into nonoverlapping laminae related to the eye specific layers of the LGNd. However, parts of the PGN immediately adjacent to the LGNd appear devoid of connections with lamina A1 thus creating a thin monocular segment for the contralateral eye.     

Histopathological Changes of Inner Retina,Optic Disc,and Optic Nerve in Rabbit with Advanced Retinitis Pigmentosa

We observed the histopathological changes of retinal ganglion cells (RGCs), optic disc, and optic nerve in rabbit with advanced retinitis pigmentosa (RP). Wild-type (WT) and rhodopsin transgenic (Tg) of RP rabbits were used at age 24 months. Light and electron microscopy were used to observe the retina, optic disc, and optic nerve. RGCs were also confirmed by immunofluorescent staining with a TUJ-1 monoclonal antibody. In addition to the rod and cone degeneration, we observed the astrocyte infiltration of the optic disc due to the damage of small RGCs and nerve fibres and atrophy of small optic nerve fibres. They subsequently lead to the optic disc excavation and atrophy of the optic nerve. Consequently, our histopathological study clarified that not only the outer retina but also the inner retina, the optic disc, and the optic nerve were also affected in the late stages of RP rabbit.     

Visual electrophysiology in the clinical evaluation of optic neuritis,chiasmal tumours,achiasmia, and ocular albinism: an overview

Background and Methods In routine clinical evaluation of optic neuritis and chiasmal tumours, pattern electroretinography and visual evoked potentials (VEPs) to pattern-reversal stimulation are useful examinations. Similarly, in achiasmia and ocular albinism, VEPs to flash and pattern-onset stimulation provide relevant information. Results The role of visual electrophysiology in these diseases is to assess potential dysfunction of the visual pathway: (a) at the acute stage of optic neuritis, to determine the magnitude of conduction block of the optic nerve fibres; (b) at the clinical recovery stage of optic neuritis, to determine optic nerve conduction delay due to demyelination, and to follow possible remyelination; (c) at the recovery of optic neuritis when visual acuity does not normalise, to define loss of optic nerve fibres and retrograde degeneration of retinal ganglion cells; (d) in tumours at the chiasm, to detect abnormal conduction along the crossed and/or uncrossed fibres; and (e) in achiasmia or albinism, which are both congenital disorders associated with nystagmus, to detect achiasmia and absence of or reduced optic nerve fibre decussation at the chiasm, or to detect ocular albinism and excess of optic nerve fibre decussation at the chiasm. In optic neuritis, two recent examinations have been used to detect retrograde axonal degeneration: photopic negative response of the electroretinogram, to assess dysfunction of ganglion cell axons; and optic coherence tomography, to measure thinning of the retinal nerve fibre layer. In optic neuritis, multifocal VEPs provide a promising clinical examination, because this can show areas that are associated with normal or abnormal optic nerve fibre function. Conclusions Visual electrophysiology defines function of the visual pathway and is relevant: (1) in optic neuritis, when visual acuity does not recover well; (2) in tumours of the chiasm with normal visual fields, as in paediatric patients who cannot adequately perform perimetry; and (3) in children with congenital nystagmus and suspected achiasmia or ocular albinism.     

Effects of selective pressure block of Y-type optic nerve fibers on the receptive-field properties of neurons in area 18 of the visual cortex of the cat.

Recordings were made from single neurons in area 18 of anesthetized cats (N2O/O2 mixture supplemented by continuous intravenous infusion of barbiturate) in which one optic nerve had been pressure blocked to selectively block conduction in the largest (Y-type) fibers. Cortical neurons were stimulated visually via the normal eye or via the eye with the pressure-blocked optic nerve ("Y-blocked eye"). Several properties of the receptive fields such as their spatial organization (S or C cells), orientation tuning, and the presence and strength of end-zone inhibition appear to be unaffected by removal of the Y input. By contrast, the removal of the Y input resulted in a small but significant reduction in the size of the discharge field and in the direction-selectivity index. In three respects, peak response discharge rate, eye dominance, and velocity sensitivity, removal of the Y input had strong and highly significant effects. Thus, the mean peak discharge frequency of responses evoked by the stimulation of binocular neurons via the Y-blocked eye was significantly lower than that of responses evoked by the stimulation via the normal eye. Accordingly, the eye-dominance histogram was shifted markedly towards the normal eye (more so than in the homologous experiment conducted on area 17-Burke et al., 1992). Finally, the mean preferred velocity of responses of cells activated via the normal eye was in the vicinity of 145 deg/s, whereas for cells activated via the Y-blocked eye the value was about 35 deg/s. Overall, the results of the present study imply that (1) apart from Y-type excitatory input there are significant excitatory non-Y-inputs to area 18; these inputs at least partially consist of indirect X-type input relayed via area 17; (2) in neurons of area 18 that receive both Y-type and non-Y-type excitatory inputs, the Y-type input has a major influence on strength of the response and velocity sensitivity and a lesser influence on the direction selectivity and size of the discharge fields; and (3) area 18 contains mechanisms determining such receptive-field properties as S- or C-type organization, orientation tuning, and direction selectivity which can be accessed either by the Y input or by non-Y input.     

Selective labeling of visual corpus callosum connections with aspartate in cat and rat

Tritiated neurotransmitter candidates were unilaterally injected in visual cortical regions with abundant corpus callosum connections. D-aspartate (Asp) or gamma-aminobutyric acid (GABA) was injected along the area 17/18 border in cat, and the area 17/18a and 17/18b borders in rat. Retrograde Asp label was found contralaterally in supragranular and infragranular laminae in areas 17, 18, 19, PMLS, and PLLS in cat, and in areas 29, 18b, 17, and 18a in rat. No contralateral GABA label was found in cat or rat. Thus, the cat and rat corpus callosum may use Asp or a related substance as a neurotransmitter.