Striate Projection Research Articles

Injuries to the Immature Optic Radiation Show Correlated Thinning of the Macular Ganglion Cell Layer.

Injuries to the immature optic radiation (OR) are associated with thinning of the retinal nerve fiber layer and corresponding visual field (VF) defects. The aim of the current study was to seek evidence for causal retrograde trans-synaptic degeneration by exploring the correspondence between the localization and extension of the injury to the OR and the structure of the macular ganglion cell complex, and the relation to VF function. Seven adults (age range 18–35) with visual dysfunction secondary to white-matter damage of immaturity and six healthy adults (age range 22–33) underwent magnetic resonance imaging (MRI). Fiber tractography was used to generate the geniculate projections to the dorsal and ventral striate cortex, delineated by retinotopic functional MRI mapping. The structure of the macular ganglion cell complex was measured with optical coherence tomography. The tractography showed overlaps between the dorsal and ventral geniculo-striate projections. However, in four patients with inferior VF defects, the dorsal projections were to a large extent traversing the space normally solely occupied by ventral projections. This is consistent with structural changes to the OR and suggests of re-organization upon injury. Diffusion parameters were significantly different between patients and controls, and most pronounced in the dorsal geniculo-striate projections, with a pattern indicating primary injury. The macular ganglion cell complex was significantly thinner in the patients and most pronounced in the superior sectors; a pattern particularly evident in the four patients with inferior VF defects. The ratio of the mean thickness of the macular ganglion cell complex in the superior and inferior sectors significantly correlated with the axial and mean diffusivities in the contra- and ipsilateral dorsal striate projections. The results suggest a causal link between injuries to the superior portion of the immature OR and secondary thinning in the macular ganglion cell complex, resulting in inferior VF defects.

Role of retinal input on the development of striate-extrastriate patterns of connections in the rat

Previous studies have shown that retinal input plays an important role in the development of interhemispheric callosal connections, but little is known about the role retinal input plays on the development of ipsilateral striate-extrastriate connections and the interplay that might exist between developing ipsilateral and callosal pathways. We analyzed the effects of bilateral enucleation performed at different ages on both the distribution of extrastriate projections originating from restricted loci in medial, acallosal striate cortex, and the overall pattern of callosal connections revealed following multiple tracer injections. As in normal rats, striate-extrastriate projections in rats enucleated at birth consisted of multiple, well-defined fields that were largely confined to acallosal regions throughout extrastriate cortex. However, these projections were highly irregular and variable, and they tended to occupy correspondingly anomalous and variable acallosal regions. Moreover, area 17, but not area 18a, was smaller in enucleates compared to controls, resulting in an increase in the divergence of striate projections. Anomalies in patterns of striate-extrastriate projections were not observed in rats enucleated at postnatal day (P)6, although the size of area 17 was still reduced in these rats. These results indicate that the critical period during which the eyes influence the development of striate-extrastriate, but not the size of striate cortex, ends by P6. Finally, enucleation did not change the time course and definition of the initial invasion of axons into gray matter, suggesting that highly variable striate projections patterns do not result from anomalous pruning of exuberant distributions of 17-18a fibers in gray matter.

Effect of stimulus localisation on motion-onset VEP

Reliable motion-onset visual evoked potentials (result of the dorsal stream activation) were recorded to motion stimuli with the temporal frequency of five cycles per seconds in 20 different locations with eccentricity up to 42° to periphery of the visual field. Amplitudes and latencies of the positive–negative–positive (P1–N1–P2; 84–144–208 ms) complex were evaluated in occipital (O Z and two derivations 5 cm to the left and right from O Z) and central region (C Z) in 10 subjects. We observed: (1) Shortening of the N1 latency toward periphery of the visual field. (2) The N1 amplitude maximum and latency minimum moved from occipital into central region (C Z derivation) as stimulus moved from centre toward periphery of visual field. (3) The P1 and N1 peaks displayed significantly greater amplitudes and shorter latencies when the lower part of the visual field was stimulated. (4) The N1 peak changed lateralisation of its maximum amplitude in dependence on the eccentricity. Up to 17°, it corresponds to striate projection of the “optic radiation” whilst more in periphery, there was paradoxical lateralisation of higher amplitude and shorter latency. The retinotopic dependence shows that the motion response includes position information and that the motion-onset VEPs are not generated solely in the higher extrastriate areas (MT or MST).

Dopamine neurons encode teaching signals for learning reward-based decision strategy

Although a critical involvement of dopamine has been implicated in the functions of the basal ganglia, especially in learning, little is known about its mechanisms. In two macaque monkeys, the nigrostriate dopamine system was unilaterally depleted by neurotoxin, 1-methyl-4-phenyl-1,2,3,6-tetrahydropyridine (MPTP). We asked the monkeys to learn sequential button press tasks. In addition to the slowness of movement, they showed specific deficits in learning action strategy and reward prediction when using contralateral arm to the dopamine depletion, but with ipsilateral arm, they learned the tasks efficiently. Activity of single, midbrain dopamine neurons was recorded from two monkeys in a task in which they chose one correct, rewarding button among three alternatives in a trial and error basis. It was found that the magnitudes of responses to beep sound informing correct and incorrect choices precisely reward prediction error signals. We present a hypothetical scheme for the mechanisms of action learning in the basal ganglia, in which the nigrostriate dopamine system provides the striatum, locus of learning, with reward prediction errors as a teaching signal for learning, and excitability of the striate projection neurons to specific cortical inputs is modified by the teaching signal so as to achieve goal-directed action selection.

Neural information transferred from the putamen to the globus pallidus during learned movement in the monkey.

1. We studied the physiology of the neuronal projection from the striatum to the external and internal segments of the globus pallidus (GPe and GPi, respectively) in macaque monkeys. The objective of the study was to answer the following specific questions. 1) Which classes of the electrophysiologically identified striate neurons project to GPe and GPi? 2) What kind of information is transferred from the striatum to GPe and GPi during learned movement? 3) What are the physiological actions of striate projection neurons on target neurons in GPe and GPi? 4) What is the spatial pattern of the striatopallidal projections? 2. Sequential arm and orofacial movements were used as behavioral tasks. Visual stimuli triggered a sequence of three flexions-extensions of the elbow joint across the target, and the click of a solenoid valve triggered repetitive licking movements. 3. Striatopallidal projection neurons were electrophysiologically identified by antidromic activation after focal stimulation of either GPe or GPi. Of two classes of striate neurons, tonically active neurons (TANs) with tonic spontaneous discharges (2-8 imp/s) and broad action potentials, and phasically active neurons (PANs) with a very low spontaneous discharge rate (< 0.5 imp/ s) and high-frequency discharges in relation to behavioral tasks, PANs were identified as the projection neurons to either GPe or GPi. In 325 TANs examined by stimulation of GPe or GPi, no neuron was activated antidromically, even in the case of TANs located in the close vicinity of PANs that were identified as striatopallidal projection neurons. 4. The physiologically identified projection neurons (52 cells) in the striatum exhibited either discharges related to movement (30 cells) or discharges related to preparation for movement (4 cells) during performance of learned motor tasks. The activities of the remaining 17 striatopallidal neurons either were not related to the behavioral tasks used or could not be characterized sufficiently in the tasks. However, all of the unidentified striatopallidal neurons were PANs, on the basis of the spontaneous discharge rate and the shape of the action potential. 5. PANs with movement-related activity and those with preparation for movement-related activity were antidromically activated from the globus pallidus (GP). Not only the PANs that show burst discharges specifically at the beginning of a sequence of movement but also PANs that show phasic discharges time-locked to each movement of a sequence were identified as putaminopallidal projection neurons. On the other hand, no neurons that showed responses to sensory stimulus were identified as putaminopallidal neurons. 6. The conduction velocities of the putaminopallidal axons were estimated at approximately 1 m/s on the basis of the latency of antidromic activation and conduction distance. The PANs with activity only at the beginning of a sequential movement were more frequently found to project to GPi than to GPe, whereas the PANs with burst activity at each movement were more frequently found to project to GPe than to GPi. Among the GPi-projecting PANs, neurons with initial activity only showed a tendency to have longer latencies of activation from GPi than neurons with activity time-locked to each movement. 7. The physiological action of the striatopallidal projection was examined by switching from recording to microstimulation after identification of striatopallidal projection neurons in the putamen while recording evoked field potentials or spike discharges of single GP neurons located where the electrical stimulation evoked antidromic activation of the striate neurons with the lowest threshold. A small majority of GP neurons that exhibited increase of discharges during motor tasks received facilitatory putaminopallidal influences, whereas the vast majority of GP neurons that exhibited decrease of discharges during motor tasks received suppressive putaminopallidal influences.

Delineation of the striate cortex, and the striate-peristriate projections in the guinea pig

The size and position of the guinea pig area 17 were determined by transneuronal labeling after intraocular injections of 3H-proline or WGA-HRP. Area 17 occupies a large region of the occipital cortex located between two shallow fissures, the fissura sagittalis lateralis and the lateral groove. Area 17 extends for about 6 mm rostral from the occipital pole of the hemisphere, and encroaches occipitally for more than 1 mm upon the ventromedial surface of the hemisphere; the lateral width is up to 4.5 mm. Single injections of WGA-HRP into area 17 produced eight patches of transported tracer which formed the same general pattern in the peristriate cortex, regardless of the position of the injection within the visual field representation of area 17. Two of these patches were found in anteromedial peristriate cortex; three patches were distributed anterolateral and lateral of area 17; and three patches were located in posterolateral peristriate cortex. For several reasons, each of these patches was interpreted as representing a single striate projection onto a separate peristriate area. Comparison of these results with published findings indicates that the parcellation of the peristriate cortex into a variety of different areas, the pattern formed by these areas around area 17, and their reciprocal connections with area 17 follow a common plan in all hitherto studied terrestrial Old World and New World rodents. Lucifer Yellow injections into striate cells projecting to one of the recipient areas (AM) indicated that the pyramidal cells of this set of striate neurons are characterized by a short apical dendrite, and that the basal dendrites of the layer V pyramidal cells branch more profusely than those of the layer III pyramids.

Patterns of interhemispheric and striate-peristriate connections in visual cortex of the South American marsupial Marmosa elegans (mouse opossum)

We have analyzed the distributions of interhemispheric and striate-peristriate connections in the South American marsupial, Marmosa elegans (mouse opossum). Following multiple injections of horseradish peroxidase (HRP) into one hemisphere, we found that anterogradely labeled terminations and retrogradely labeled perikarya are distributed unevenly in the contralateral hemisphere, forming a distinct tangential pattern in striate and peristriate cortex. This pattern delineates as many as eight peristriate areas relatively poor in commissural connections in lateral peristriate cortex, and in lateral and anterolateral portions of peristriate cortex. Single injections of HRP conjugated with wheat germ agglutinin into anterior or posterior regions of striate cortex produced as many as nine discrete ipsilateral fields of labeled perikarya, and terminations distributed over a broad cortical area in lateral and anterolateral peristriate cortex. Our observations of multiple areas with little or no HRP labeling in the interhemispheric pattern, and of multiple ipsilateral striate projection fields, indicate that the topography of visual cortex in Marmosa is highly elaborate, and suggest that extrastriate cortex is subdivided into several visual areas. Furthermore, by showing that the organization of visual cortex in this marsupial is as complex as in many placental mammals, our data support the view that a basic cortical plan, consisting of multiple visual areas, appeared early in mammalian evolution.

Visual topography of striate projection zone (MT) in posterior superior temporal sulcus of the macaque.

SUMMARY AND CONCLUSIONS 1. The representation of the visual field in the striate projection zone in the posterior portion of the superior temporal sulcus of the macaque (MT) was mapped with multiunit electrodes. The animals were immobilized and anesthetized and in each animal 25-35 electrode penetrations were typically made over several recording sessions. 2. MT contains a representation of virtually the entire contralateral visual field. The representation of the vertical meridian forms its ventrolateral border and lies near the bottom of the lower bank of the superior temporal sulcus (STS). The representation of the horizontal meridian runs across the floor of STS. The upper field is located ventral and anterior and the lower field dorsal and posterior. The medial border lies at the junction of the floor of STS and its upper bank. 3. MT is similar to striate cortex in being a first-order transformation of the visual field. In both areas, receptive-field size and cortical magnification increase with eccentricity. MT is much smaller than striate cortex and has much larger receptive fields at a given eccentricity and a cruder topography. 4. The results further support the suggestion that MT in the macaque is homologous to visual area MT in New World primates.

The striate projection zone in the superior temporal sulcus of Macaca mulatta: location and topographic organization.

In the rhesus monkey, the caudal portion of the superior temporal sulcus (STS) receives a direct projection from lateral striate cortex, the striate are representing central vision. The present study was undertaken to determine whether STS also receives a direct projection from areas of striate cortex representing peripheral vision, with the intent of defining the entire striate projection zone in STS as well as providing information regarding a possible topographic organization within this secondary visual area. A series of five rhesus monkeys was prepared with unilateral lesions of lateral, posterior, or medial striate cortex, such that, collectively, the lesions in the series included all of striate cortex with little or no invasion of prestriate cortex. The monkeys were sacrificed seven days after surgery and their brains were processed by the Fink-Heimer procedure. An analysis of the distribution of terminal degeneration within STS indicated: (1) All areas of striate cortex project to a restricted region along the caudal portion of STS. The ventral limit of this region can be demarcated by an imaginary line connecting the ventral tips of the lunate and intraparietal sulci; from this limit the region extends dorsocaudally for approximately 12 mm to the point at which STS frequently bifurcates, sending one spur forward into the inferior parietal lobule. (2) Within this portion of STS there is an orderly mapping of the visual field; progression from central vision to the far periphery is represented by a progression down the posterior bank of STS and continuing along the entire floor, or insula-like portion, of the sulcus. (3) Projections from striate cortex to STS terminate predominantly in layer IV and the deep part of layer III. (4) There is a distinctive pattern of myelination contained within the striate projection zone of STS. These anatomical findings concerning the striate projection zone of STS in the rhesus monkey are remarkably similar to those that have been described for the middle temporal visual area (MT) in New World monkeys, and thus support earlier proposals that the two areas are homologous.

Pathways between striate cortex and subcortical regions in Macaca mulatta and Saimiri sciureus: Evidence for a reciprocal pulvinar connection

The efferent and afferent connections of striate cortex have been compared in Macaca mulatta and Saimiri sciureus monkeys after injecting striate cortex with a mixture of horseradish peroxidase and tritiated amino acids. Three reciprocal pathways were found in Macaca; they were between striate cortex and (i) all layers of the dorsal lateral geniculate nucleus; (ii) the inferior subdivision of pulvinar; and (iii) the lateral subdivision of pulvinar. In Saimiri the only recprocal pathway involved the dorsal lateral geniculate nucleus; the pulvinar received a heavy striate input but too few peroxidase-labeled neurons were found in pulvinar to demonstrate convincingly a projection to striate cortex. In both species the reticular and posterior nucleus of the thalamus and the superior colliculus receive striate input; in Saimiri the ventral lateral geniculate also receives a striate projection. In both species, neurons in the basal forebrain project to striate cortex. Striate cortex synaptic terminals in both dorsal lateral geniculate nucleus and pulvinar are topographically organized. Inferior and lateral subdivisions of pulvinar each contain a representation of central retina, and the fiber bundles separating the two subdivisions apparently mark the vertical meridian. The results of this experiment suggest that both retino-geniculate and retino-superior colliculus-pulvinar types of visual information may converge within the striate cortex.

A comparison of the effects of inferotemporal and striate cortex lesions on the visual behaviour of rhesus monkeys.

The effects of bilateral removal of inferotemporal cortex or lateral striate cortex are compared. The former operation impairs visual pattern discrimination learning, without disturbing prompt detection and retrieval of food, visual acuity or visual fields. In contrast, animals in which the macular striate projection area has been removed are significantly superior in tests of visual pattern discrimination learning despite showing impairment of visually guided reaching, visual acuity, and visual fields. The results indicate that the inferotemporal defect is not caused by deficient visual sensitivity or acuity.