Optic flow direction coding in area PEc of the behaving monkey.

The cortical representation of heading perception derives from several functional processes distributed across many cortical areas. The aim of the present study was to assess if the optic flow motion directions, expansion and contraction, differently modulate the firing activity of area PEc neurons. We determined the influence of the eye position and/or the spatial position of the focus of expansion (FOE) on this activity. Single neuron activity during radial optic flow stimulation was recorded in three behaving monkeys. The retinal FOE position and the spatial eye position were examined in order to study eye position's influence upon the directional selectivity for the radial stimuli. We observed that the neurons able to discriminate the retinotopic FOE position are differently modulated by expansion and contraction. One class of neurons exhibited a different preferred FOE position during expansion and contraction. A second class showed the same preferred position with similar firing activity in the two stimuli. A third class showed the same preferred position but different firing activity. Eye position affected the directional selectivity of most PEc cells. The main result of this study is that there is a continuum in cell modulation by optic flow direction, and it can be modified by the angle of gaze with respect to the FOE. These results shed light on potential cellular integrative mechanisms of area PEc in heading perception.

Multimodal representation of optic flow in area PEc of macaque monkey.

The visual perception of self-motion is mainly provided by optic flow. Eyes usually scan the environment during locomotion, and the gaze is not always directed to the focus of expansion (FOE) of the flow field. Such eye movements change the retinal FOE position with respect to the fovea. Here, we assess if optic flow selective neurons in parietal area PEc are modulated by eye position. We recorded single neuron activity during radial optic flow stimulation in two monkeys, varying eye and retinal FOE positions. We found that the majority of PEc neurons are modulated by the FOE retinotopic position with different tuning for expansion and contraction. Although many neurons did not show any gaze field without visual stimulation, the eye position modulated optic flow responses in about half of the cells. These novel results suggest that PEc neurons integrate both visual and eye position signals, and allow us to hypothesize their role in guiding locomotion as a part of a cortical network involved in FOE representation during self-motion. Visual and eye position interaction in this area could be seen as a contribution to the building of the invariant space representation necessary to motor planning.

Neuronal responses in macaque area PEc to saccades and eye position.

Neurons in area PEc in the superior parietal cortex encode signals from different modalities, such as visual, extraretinal and somatosensory, probably combining them to encode spatial parameter of extrapersonal space to prepare body movements. This study reports the characterization of the functional properties of PEc non-visual neurons that showed saccade-related activity. We analyzed the pre- and post-saccadic firing activity in 189 neurons recorded in five hemispheres of three behaving monkeys. Spiking activity of PEc single neurons was recorded while the monkeys performed visually-guided saccades in a reaction time task. We found that 84% of neurons recorded from area PEc showed pre-saccadic activity with directional tuning. In 26% of neurons, we found inhibition of activity in the pre-saccadic period. The onset of this "pause" always started before the saccade and, in 51% of neurons, it was invariant among different gaze directions. The post-saccadic activity in these cells was either a phasic response with directional tuning (77%) and/or an eye position tuning (75%). The analysis of the preferred direction did not show hemispheric preference, however, for the majority of neurons, the angular difference in the preferred direction, in the pre- and post-saccadic period, was more than 60 degrees . By confirming, therefore, that PEc neurons carry information about eye position, these novel findings open new horizons on PEc function that, to date, is not well documented. The pre-saccadic activity may reflect an involvement in saccade control, whereas post-saccadic activity may indicate a role in informing on the new eye position. These novel results about saccade and eye position processing may imply a role of area PEc in gaze direction mechanisms and, possibly, in remapping visual space after eye movements.

Evaluation of an occupational therapy program for patients with spinal cord injury.

STUDY DESIGN: Clinical controlled trial. OBJECTIVES: To evaluate the effectiveness of an occupational therapy (OT) program combined with neuromotor rehabilitation, by assessing the degree of functional independence reached by patients with spinal cord injuries at first hospitalization. SETTINGS: Subjects selected from the Spinal Cord Unit of the Rehabilitation Institute of Montecatone (Imola, Italy). PARTICIPANTS: Thirty-six male patients below age 60, with complete paraplegia (ASIA-A) in thoracic-lumbar level, at first hospitalization. METHODS: Patients were divided into experimental and control groups. Subjects in the experimental group underwent neuromotor rehabilitation coupled with an OT program, whereas those in the control group followed neuromotor rehabilitation only. Increase in functional independence at discharge was evaluated by the Valutazione Funzionale Mielolesi (VFM) assessment scale. RESULTS: Patients in the experimental group showed a significant increase in the total VFM score, and in domains concerning transfers and wheelchair use. A significant improvement was observed in unmarried patients as compared to married ones. CONCLUSION: An OT service within a Spinal Cord Unit allows us to achieve a higher level of functional independence.

Visual motion responses of neurons in the caudal area pe of macaque monkeys.

Area PE of macaques has traditionally been considered a somatosensory association cortex. Recent studies, however, suggest that neurons of this and neighboring areas are involved in the visual control of movement, especially arm movement. We investigated the neuronal sensitivity to local visual stimuli of this region by recording neuronal activity in two behaving macaque monkeys trained in a simple visual fixation task. Recordings were performed from the dorsal surface of the caudal pole of the superior parietal lobule (SPL). Classical receptive fields (RFs) were mapped by using conventional static or moving luminous figures. We found that many neurons in this area were selectively activated by moving visual stimuli. Cell responses were tuned to the movement direction. RFs were usually large; their mean surface covered some 30 x 30 degrees of the visual field. The fovea was often included into RF, in many cases it was along a RF side. The center of RFs was mainly located in the contralateral hemifield, although RFs having the center ipsilaterally sited were also found. No evident retinotopy was found. Visual neurons were especially concentrated in a region of the SPL likely corresponding to area PEc. These results suggest that the caudal part of area PE contains neuronal populations specifically signaling local visual motion, possibly encoding the direction of moving objects. These signals might well be suited for sensorimotor integration mechanisms aimed at motor acts.

Early coding of reaching: frontal and parietal association connections of parieto-occipital cortex.

The ipsilateral association connections of the cortex of the dorsal part of the rostral bank of the parieto-occipital sulcus and of the adjoining posterior part of the superior parietal lobule were studied by using different retrograde fluorescent tracers. Fluoro-Ruby, Fast blue and Diamidino yellow were injected into visual area V6A, and dorso-caudal (PMdc, F2) and dorso-rostral (PMdr, F7) premotor cortex, respectively. The parietal area of injection had been previously characterized physiologically in behaving monkeys, through a variety of oculomotor and visuomanual tasks. Area V6A is mainly linked by reciprocal projections to parietal areas 7m, MIP (medial intraparietal) and PEa, and, to a lesser extent, to frontal areas PMdr (rostral dorsal premotor cortex, F7) and PMdc (F2). All these areas project to that part of the dorsocaudal premotor cortex that has a direct access to primary motor cortex. V6A is also connected to area F5 and, to a lesser extent, to 7a, ventral (VIP) and lateral (LIP) intraparietal areas. This pattern of association connections may explain the presence of visually-related and eye-position signals in premotor cortex, as well as the influence of information concerning arm position and movement direction on V6A neural activity. Area V6A emerges as a potential 'early' node of the distributed network underlying visually-guided reaching. In this network, reciprocal association connections probably impose, through re-entrant signalling, a recursive property to the operations leading to the composition of eye and hand motor commands.

Corticocortical connections between frontal periarcuate regions and visual areas of the superior temporal sulcus and the adjoining inferior parietal lobule in the macaque monkey.

In macaque monkeys, corticocortical connections between distinct parietotemporal visual areas (areas MST-FST, DP, and 7a) and frontal periarcuate areas are studied using tritiated aminoacids and WGA-HRP. While labeling within the banks of the principal sulcus, the dorsal part of the arcuate concavity, and the banks of the upper arcuate limb were present in both 7a and MST-FST injected animals; in the latter cases, additional projections were found towards frontal regions including the dorsomedial frontal cortex and the posterior bank of the arcuate ventral limb. Our results point to widespread frontal connections of the MST-FST complex, involving both prefrontal and premotor cortical regions.

Encoding of smooth pursuit direction and eye position by neurons of area MSTd of macaque monkey.

The spike activity of neurons was recorded from the dorsal bank of the superior temporal sulcus (area MSTd) of alert behaving macaque monkeys performing visual fixation or target tracking tasks, with the aim of studying the tuning features of these neurons with both the direction of slow eye movement and the position of gaze. One hundred thirty-two neurons were tested for several fixation points and tracking directions. Many of them (43%) tuned to the direction of pursuit, regardless of the angle of gaze. Some (18%) showed a tonic discharge modulated by the static position of the eyes without pursuit direction specificity. A substantial number of cells (22%) were characterized by a discharge rate tuned to pursuit direction but influenced also by angle of gaze. Tuning curves for eye movement direction presented an average bandwidth of 130 degrees and turned out to be continuously overlapping, suggesting a sort of vector coding of smooth pursuit direction. Gaze fields of eye position (EP) neurons were mostly ramp-like, with center of ramps shifted away from the straight ahead, implying a form of scalar coding of gaze eccentricity. The different categories of cells were intermingled and close to each other, suggesting possible reciprocal interactions within the same cortical area. These results show that EP and pursuit direction are signaled mainly by separate neuronal elements in area MSTd. Moreover, some cells can integrate both signals. Taking into account the visual responses of MSTd neurons to large, textured, moving fields, it is suggested that this area could be the site of interaction between visuo-oculomotor signals related to visual motion detection, slow eye movement direction, and EP. This signal interaction may be important for integrative functions such as analysis of external or self-induced visual motion, cortical control of pursuit eye movements, and eye/head coordination.

Gaze field properties of eye position neurones in areas MST and 7a of the macaque monkey.

The activity of parietal cortex neurones primarily related to eye position (EP neurones) was studied in macaque monkeys with the aim of precisely defining the neurones' gaze fields (GF) and comparing them in two functionally different areas, MSTd and 7a. Discharge rates of single neurones in the inferior parietal lobule and in the underlying cortex of the superior temporal sulcus were recorded in two Java monkeys while the animals fixated a steady visual target positioned at several different points on a video screen. The GFs were then drawn as a regression surface fitting the mean discharge rates. Cells tonically influenced by the angle of gaze were found in both areas. The GFs most often took the form of a nearly planar surface best characterized as a ramp tilted towards a hemifield or quadrant of the visual field, shifted eccentrically with respect to the straight ahead (primary position), and with a midpoint centred between 0 deg and 20 deg of gaze eccentricity and saturation between 10 deg and 35 deg. In a minority of cases, the discharge rate was nearly maximal at the primary position and decreased to a minimum within 35 deg of eccentricity. In other instances, the GFs were peaked surfaces, limited to a restricted part of visual space. EP neurones, while showing similar gaze fields in areas MST and 7a, were found intermingled with functionally different types of cells. The results suggest that EP neurones similar to those already described in several areas of the monkey parietal cortex are present also in area MST. These cells, by signalling the degree of gaze eccentricity from the primary position, encode gaze position in an orbito-centered frame extending up to 30-35 deg from the straight-ahead. The role of EP neurones might be to supply contiguous elements with a gaze eccentricity signal required for visuo-motor processes such as the control of tracking movements.

Projections from cortical visual areas of the superior temporal sulcus to the superior colliculus, in macaque monkeys.

1. The distribution of tectal projections of two visual areas of the superior temporal sulcus (MT and MST areas) has been studied, in five Macaca fascicularis, by means of the autoradiographic method tracing the anterograde transport of tritiated aminoacids intracortically injected. 2. In all cases the ipsilateral superior colliculi (SC) were found labelled, whereas the contralateral ones were devoid of label. 3. The three brains injected in the MT area resulted in SC labels that involved the superficial gray layer (SGS), the stratum opticum (SO) and the intermediate gray layer (SGI), sparing the layers below SGI. 4. The collicular labels found after injections within the MST area exhibited their distribution over the deep SC subdivision, whereas they spared all the superficial layers but the deep part of the SO. 5. In two animals with large uptake zones, one in MT and the other in MST, the labelling within the SGI showed a cluster-like pattern. 6. The distinct found bulk of projections of MT and MST respectively to the superficial and deep subdivisions of the SC, along with a number of peculiar connections of the MST area as mentioned in the text, contribute to depict an overall neural network in which MST appears to be more strongly involved than MT in linking sensory visual with oculomotor attentive functions.

Projections from visual cortical areas of the superior temporal sulcus to the lateral terminal nucleus of the accessory optic system in macaque monkeys.

Tritiated amino acids were injected into the striate area and in single visual areas of the superior temporal sulcus (STS) of 7 cynomolgus monkeys, in order to trace visual cortical projections to the nuclei of the accessory optic system (AOS). Injections in STS separately involved the areas MT and MST, and resulted in labels within the lateral terminal nucleus of the AOS. In no case were labels found within the AOS nuclei in the brains injected in the striate area, or within the contralateral AOS. It seems likely that the areas MT and MST contribute signals--selectively related to visual motion processing--to the AOS, which is probably involved in the neuronal pathway subserving the optokinetic reflex.

Effect of fast moving stimuli and saccadic eye movements on cell activity in visual areas V1 and V2 of behaving monkeys.

Extracellular recordings were carried out in the visual cortex of behaving monkeys trained on a fixation/detection task, during which a target light was displayed stationary or suddenly moving on a tangent translucent screen. The responses of visual cortical cells to fast moving stimuli during steady fixation and those obtained during rapid eye movements (saccades) which moved their receptive field across a stationary stimulus, were studied. Areas V1 and V2 were explored. When tested with rapidly moving stimuli (500 deg/sec) during steady fixation, neurons in each area behaved in almost the same way. About one fourth of them were activated, the remainder showing either no response (little more than a half of them) or a reduction of the spontaneous firing rate. In both areas, some of the neurons activated during steady fixation did not respond or responded very weakly during eye motion at saccadic velocity (500 +/- 50 deg/sec). Neurons of this type, which we refer to as 'real motion' cells, could somehow contribute to the maintenance of visual stability during the execution of large eye movements.

Projections from the cortex of the superior temporal sulcus to the dorsal lateral geniculate and pregeniculate nuclei in the macaque monkey.

Cortical projections from the visual region and adjacent polysensory region of the superior temporal sulcus (STs) to the lateral geniculate body (LGb) were investigated in the macaque monkey using an autoradiographic tracing method. Solutions of tritiated aminoacids were injected into different parts of the caudal half of the STs of five animals. A survival time of 7 days was allowed. Labels were found in both subdivisions of the LGb: the dorsal lateral geniculate nucleus (DLGn) and the pregeniculate nucleus (PGn). In particular, part of the visual cortical region adjacent to the middle temporal area (MT) projects into the DLGn as well as the PGn, whereas the MT itself and the superior temporal polysensory region project into the PGn only. Afferents to the DLGn terminate in the magnocellular layers and in their adjoining interlaminar zones, completely sparing the parvocellular layers. Afferents to the PGn terminate in separate regions of this nucleus; the MT and adjacent visual cortices project into the internal layer of the PGn, whereas the polysensosy region of the STs projects into the external retinorecipient layer of the PGn. Possible functional implications of these projections are discussed.

Single unit activity and visual perception of motion.

Extracellular recordings were carried out in the primary visual cortex of behaving macaque monkeys. Neurons were activated by moving a visual stimulus across their receptive fields during steady fixation and by moving their receptive fields (by visual tracking) at the same velocity across a stationary visual stimulus. Out of a total number of 123 cells studied, 111 were activated by the visual stimulation both during fixation and tracking. The remaining 12 cells showed good response during fixation and very weakened response during tracking. Our results show that a small percentage of striate neurons in macaque monkey could discriminate between the real motion of an object in the visual world and the self-induced displacement of its retinal image.

Projections from the visual cortical region of the superior temporal sulcus to the striatum and claustrum in the macaque monkey.

Solutions of tritiated aminoacids were injected into the visual cortical region hidden in the depth of the superior temporal sulcus (STs) of the macaque monkey. Injection sites involved the middle temporal visual area, extending into the surrounding visual cortices. Projections were found homolaterally in both the striatum and claustrum. In the caudate nucleus labeled material affected mainly the body and spread both to the tail and the head of the nucleus. Label was also seen in the caudal third of the putamen and in the postero-ventral claustrum. Compared with the scarcity of afferents arising from occipital visual areas, the present data point to a heavy projection system from additional visual areas of the STs to the basal ganglia in the macaque monkey.

Non-oriented cells of the striate cortex activated during smooth pursuit eye movements and steady fixations in behaving monkeys.

Extracellular recordings were carried out in the primary visual cortex of behaving monkeys. Neurons were activated by moving a visual stimulus on their receptive fields during periods of steady fixation and by moving their receptive fields (smooth pursuit eye movements) on a motionless visual stimulus. Regarding non-oriented cells, they turned out to be activated by the visual stimulation both during steady fixations and smooth pursuit eye movements. Therefore, the non-oriented cells we studied seem not to receive an extraretinal signal related to the slow eye movements.

Corticopontine projections from the visual area of the superior temporal sulcus in the macaque monkey.

1. The projections from the superior temporal sulcus (STS) visual area to pontine nuclei were studied in the macaque monkey by the autoradiographic tracing method. Microinjections of a mixture of L-[5-3H] proline and L-[4,5-3H] leucine were carried out in the posterior band and the floor of the caudal half of the STS. Survival time was always 7 days. 2. The STS visual area projects to the dorsolateral part of ipsilateral pontine nuclei. Terminal projections are distributed in patches in the whole rostro-caudal extent of the pons. 3. These findings support the view that the STS visual area in the macaque monkey is homologous to the postero-medial lateral suprasylvian area in the cat.

Bilateral projections from the visual cortex to the striatum in the cat.

Direct projections from visual areas 17, 18, 19, and lateral suprasylvian visual area (LS) to the striatum were searched for in 12 adult cats using the autoradiographic technique to detect neuronal pathways. Striatal labels were found only after injections in areas 19 and LS. Projections homolateral to the injection sites were observed from both areas to the head and body of the caudate nucleus and to the putamen. Contralateral projections were found from both areas 19 and LS: however, area 19 did not project to the contralateral putamen. The extent of contralateral projections was smaller and they were confined within the same regions as the homolateral ones. Silver grains were often arranged in cluster-like patches, which were more evident ipsilaterally, in the head of the caudate nucleus and after injections in area LS. The present data support the view of a not strictly topographical segregation of striatal projections from the cat visual cortex.