Functional sub-regions for optic flow processing in the posteromedial lateral suprasylvian cortex of the cat.

During locomotion, an observer sees a large and complex pattern of visual motion called optic flow. This phenomenon is characterized by elements in the environment accelerating and expanding as they move peripherally. In cats, previous studies have indicated that the posteromedial part of the lateral suprasylvian (PMLS) cortex may be involved in the processing of optic flow fields. We further addressed this issue by studying the importance of specific parameters of the optic flow patterns and investigating whether cell responses to these stimuli depend on receptive field (RF) location in the visual field. Results can be summarized as follows: approximately two-thirds of PMLS cells responded to optic flow fields and a subset of these (84/153) showed a clear direction selectivity for motion along the frontal axis. Of these units, the majority responded preferentially to expansion rather than contraction of the pattern. Cells' responses depend on RF location in the visual field. For centrally located RFs, tested both when the origin of motion was within the RF or at the area centralis, responses were generally comparable whether or not size or speed gradients were removed from the optic flow pattern. A different tendency was observed for peripherally located RFs. In general, these cells exhibited a preferred direction almost exclusively when the origin of motion was placed at the area centralis, and neuronal discharges and direction selectivity for many of them were reduced when the optic flow cues were removed from the pattern. The results of this study suggest that there may be functional differences in response properties between PMLS cells located in the central and peripheral parts of the visual field that may reflect a specialization of the PMLS cortex in optic flow processing.     

Neuronal responses to optic flow in the monkey parietal area PEc

Area PEc, a high order association area, is located in the dorsocaudal portion of the superior parietal cortex. PEc neurons encode visual motion signals, especially the direction of stimulus motion. The present study tested if PEc neurons also process visual correlates of self-motion. The extracellular activity of single neurons in response to optic flow stimuli was recorded in two monkeys (Macaca fascicularis) trained in a fixation task. The stimuli were produced by random dots simulating planar motion, radial expansion and radial contraction. A substantial number of PEc neurons were specifically activated by radial optic flow and were selective for the position of the focus of expansion with respect to the fovea. Eccentric positions of the focus of expansion were preferred. Almost all neurons showed opponent excitatory-inhibitory activity to expanding-contracting visual fields. Planar motion elicited very weak responses. Optic flow responsiveness is not entirely explained by classical bar sensitivity in PEc neurons, suggesting that optic flow and classical bar responses could serve different mechanisms in the integration of visuo-motor signals to prepare body movements.     

Modulation of responses to optic flow in area 7a by retinotopic and oculomotor cues in monkey

Perception of two- and three-dimensional optic flow critically depends upon extrastriate cortices that are part of the 'dorsal stream' for visual processing. Neurons in area 7a, a sub-region of the posterior parietal cortex, have a dual sensitivity to visual input and to eye position. The sensitivity and selectivity of area 7a neurons to three sensory cues - optic flow, retinotopic stimulus position and eye position - were studied. The visual response to optic flow was modulated by the retinotopic stimulus position and by the eye position in the orbit. The position dependence of the retinal and eye position modulation (i.e. gain field) were quantified by a quadratic regression model that allowed for linear or peaked receptive fields. A local maximum (or minimum) in both the retinotopic fields and the gain fields was observed, suggesting that these sensory qualities are not necessarily linearly represented in area 7a. Neurons were also found that simply encoded the eye position in the absence of optic flow. The spatial tuning for the eye position signals upon stationary stimuli and optic flow was not the same, suggesting multiple anatomical sources of the signals. These neurons can provide a substrate for spatial representation while primates move in the environment.      

Behavioral influences on cortical neuronal responses to optic flow

Optic flow selectively activates neurons in medial superior temporal (MST) cortex. We find that many MST neurons yield larger and more selective responses when the optic flow guides a subsequent eye movement. Smaller, less selective responses are seen when optic flow is preceded by a flashed precue that guides eye movements. Selectivity can decrease by a third (32%) after a flashed precue is presented at a peripheral location as a small spot specifying the target location of the eye movement. Smaller decreases in selectivity (18%) occur when the precue is presented centrally with its shape specifying the target location. Shape precues presented centrally, but not linked to specific target locations, do not appear to alter optic flow selectivity. The effects of spatial precueing can be reversed so that the precue leads to larger and more selective optic flow responses: A flashed precue presented as a distracter before behaviorally relevant optic flow is associated with larger optic flow responses and a 45% increase in selectivity. Together, these findings show that spatial precues can decrease or increase the size and selectivity of optic flow responses depending on the associated behavioral contingencies.     

Cortical neuronal responses to optic flow are shaped by visual strategies for steering

We hypothesized that neuronal responses to virtual self-movement would be enhanced during steering tasks. We recorded the activity of medial superior temporal (MSTd) neurons in monkeys trained to steer a straight-ahead course, using optic flow. We found smaller optic flow responses during active steering than during the passive viewing of the same stimuli. Behavioral analysis showed that the monkeys had learned to steer using local motion cues. Retraining the monkeys to use the global pattern of optic flow reversed the effects of the active-steering task: active steering then evoked larger responses than passive viewing. We then compared the responses of neurons during active steering by local motion and by global patterns: Local motion trials promoted the use of local dot movement near the center of the stimulus by occluding the peripheral visual field midway through the trial. Global pattern trials promoted the use of radial pattern movement by occluding the central visual field midway through the trial. In this study, identical full-field optic-flow stimuli evoked larger responses in global-pattern trials than in local motion trials. We conclude that the selection of specific visual cues reflects strategies for active steering and alters MSTd neuronal responses to optic flow.     

A more biologically plausible learning rule than backpropagation applied to a network model of cortical area 7a.

Area 7a of the posterior parietal cortex of the primate brain is concerned with representing head-centered space by combining information about the retinal location of a visual stimulus and the position of the eyes in the orbits. An artificial neural network was previously trained to perform this coordinate transformation task using the backpropagation learning procedure, and units in its middle layer (the hidden units) developed properties very similar to those of area 7a neurons presumed to code for spatial location (Andersen and Zipser, 1988; Zipser and Andersen, 1988). We developed two neural networks with architecture similar to Zipser and Andersen's model and trained them to perform the same task using a more biologically plausible learning procedure than backpropagation. This procedure is a modification of the Associative Reward-Penalty (AR-P) algorithm (Barto and Anandan, 1985), which adjusts connection strengths using a global reinforcement signal and local synaptic information. Our networks learn to perform the task successfully to any degree of accuracy and almost as quickly as with backpropagation, and the hidden units develop response properties very similar to those of area 7a neurons. In particular, the probability of firing of the hidden units in our networks varies with eye position in a roughly planar fashion, and their visual receptive fields are large and have complex surfaces. The synaptic strengths computed by the AR-P algorithm are equivalent to and interchangeable with those computed by backpropagation. Our networks also perform the correct transformation on pairs of eye and retinal positions never encountered before. All of these findings are unaffected by the interposition of an extra layer of units between the hidden and output layers. These results show that the response properties of the hidden units of a layered network trained to perform coordinate transformations, and their similarity with those of area 7a neurons, are not a specific result of backpropagation training. The fact that they can be obtained by a more biologically plausible learning rule corroborates the validity of this neural network's computational algorithm as a plausible model of how area 7a may perform coordinate transformations.     

Area MST and heading perception in macaque monkeys

The macaque medial superior temporal area (MST) is proposed to be specialized for analyzing complex 'optic flow' information. Such space-varying motion patterns provide a rich source of information about self motion, scene structure and object shape. We report the performance of rhesus macaques on a two-alternative 'heading' task, in which they reported whether horizontally varying, simulated trajectories were to left or right of center. Monkeys were sensitive to small heading angles; thresholds averaged 1.5-3 degrees. Heading estimates were stable in the face of changing stimulus location and smooth pursuit eye movements. In addition, we tested the role of area MST in heading judgements by electrically activating columns of neurons in this area while the monkeys performed the heading task. Activation of MST frequently affected performance, usually causing choice biases. These induced biases were often large and usually concordant with the preference of the neurons being activated. In addition, the induced biases were often larger in the presence of smooth pursuit eye movements. These results favor the hypothesis that MST is involved in recovering self-motion direction from optic flow cues and in the process by which heading perception is compensated for ongoing eye movements.     

Shape selectivity for camouflage-breaking dynamic stimuli in dorsal V4 neurons

Motion is a potent cue for breaking camouflage in the natural world. To understand the neural basis of this phenomenon, one must utilize moving shapes defined by coherent motion of random texture elements against a similar, but stationary texture. To investigate how well neurons in area V4 process this novel, ecologically relevant stimulus and to compare shape selectivity for these shapes with static and other moving shapes, we tested V4 neurons with 5 static or moving shapes defined either by luminance or kinetic cues. The kinetic cues included a temporal frequency cue due to the difference in temporal frequencies of the moving dots inside the shape boundary and stationary dots outside the boundary. Therefore, static opponent motion-defined shapes without this cue were tested as an additional control. Approximately 44% (95/216) of V4 neurons showed shape selectivity. Analyses of these selective neurons both at single-neuron and population levels revealed that the shape-selective V4 neurons responded selectively to the moving kinetic shapes and that these neurons demonstrated robust invariance for shape preference across different shape conditions. Cue-invariant shape selectivity was more pronounced when kinetic shapes included the temporal frequency cue. This invariance may be rooted in nonlinearities occurring early in the visual pathway.     

A possible regulatory role of glyoxalase I in cell viability of human prostate cancer

The medial parieto-occipital cortex is a central node in the dorsomedial visual stream. Recent physiological studies in the macaque monkey have demonstrated that the medial parieto-occipital cortex contains two areas, the visual area V6 and the visuomotor area V6A. Area V6 is a retinotopically organized visual area that receives form and motion information directly from V1 and is heavily connected with the other areas of the dorsal visual stream, including V6A. Area V6A is a bimodal visual/somatosensory area that elaborates visual information such as form, motion and space suitable for the control of both reaching and grasping movements. Somatosensory and skeletomotor activities in V6A affect the upper limbs and involve both the transport phase of reaching and grasping movements. Finally, V6A is strongly and reciprocally connected with the dorsal premotor cortex controlling arm movements. The picture emerging from these data is that the medial parieto-occipital cortex is well equipped to control both proximal and distal movements in the online visuomotor guidance of prehension. In agreement with this view, selective V6A lesions in monkey produce misreaching and misgrasping with the arm contralateral to the lesion in visually guided movements. These deficits are similar to those observed in optic ataxia patients and suggest that human and monkey superior parietal lobules are homologous structures, and that optic ataxia syndrome is the result of the lesion of a 'human' area V6A.     

Changes in prefrontal neuronal activity after learning to perform a spatial working memory task

The prefrontal cortex is considered essential for learning to perform cognitive tasks though little is known about how the representation of stimulus properties is altered by learning. To address this issue, we recorded neuronal activity in monkeys before and after training on a task that required visual working memory. After the subjects learned to perform the task, we observed activation of more prefrontal neurons and increased activity during working memory maintenance. The working memory-related increase in firing rate was due mostly to regular-spiking putative pyramidal neurons. Unexpectedly, the selectivity of neurons for stimulus properties and the ability of neurons to discriminate between stimuli decreased as the information about stimulus properties was apparently present in neural firing prior to training and neuronal selectivity degraded after training in the task. The effect was robust and could not be accounted for by differences in sampling sites, selection of neurons, level of performance, or merely the elapse of time. The results indicate that, in contrast to the effects of perceptual learning, mastery of a cognitive task degrades the apparent stimulus selectivity as neurons represent more abstract information related to the task. This effect is countered by the recruitment of more neurons after training.     

Three-dimensional structure-from-motion selectivity in the anterior superior temporal polysensory area, STPa, of the behaving monkey

Human and non-human primates are able to perceive three-dimensional structure from motion displays. Three-dimensional structure-from-motion (object-motion) displays were used to test the hypothesis that neurons in the anterior division of the superior temporal polysensory area (STPa) of monkeys can selectively respond to three-dimensional structure-from-motion. Monkeys performed a reaction time task that required the detection of a change in the fraction of structure in three-dimensional transparent sphere displays. Neurons were able to distinguish structured and unstructured three-dimensional optic flow. These cells could differentiate the change in structure-from-motion at stimulus presentation and when the animal was detecting the amount of structure in the display. Some of these neurons were also tuned for characteristics of the sphere stimuli. Cells were also tested with navigational motion and many were found to respond both to three-dimensional structure-from-motion and navigational motion. These results suggest that STPa neurons represent specific aspects of three-dimensional surface structure and that neurons within STPa contribute to the perception of three-dimensional structure-from-motion.     

Lack of orientation and direction selectivity in a subgroup of fast-spiking inhibitory interneurons: cellular and synaptic mechanisms and comparison with other electrophysiological cell types

Neurons in cat area 17 can be grouped in 4 different electrophysiological cell classes (regular spiking, intrinsically bursting, chattering, and fast spiking [FS]). However, little is known of the functional properties of these different cell classes. Here we compared orientation and direction selectivity between these cell classes in cat area 17 and found that a subset of FS inhibitory neurons, usually with complex receptive fields, exhibited little selectivity in comparison with other cell types. Differences in occurrence and amplitude of gamma-range membrane fluctuations, as well as in numbers of action potentials in response to optimal visual stimuli, did not parallel differences observed for orientation and direction selectivity. Instead, differences in selectivity resulted mostly from differences in tuning of the membrane potential responses, although variations in spike threshold also contributed: weakly selective FS neurons exhibited both a lower spike threshold and more broadly tuned membrane potential responses in comparison with the other cell classes. Our results are consistent with the hypothesis that a subgroup of FS neurons receives connections and possesses intrinsic properties allowing the generation of weakly selective responses. The existence of weakly selective inhibitory neurons is consistent with orientation selectivity models that rely on broadly tuned inhibition.     

Eye-hand coordination during reaching. II. An analysis of the relationships between visuomanual signals in parietal cortex and parieto-frontal association projections

The relationships between the distribution of visuomanual signals in parietal cortex and that of parieto-frontal projections are the subject of the present study. Single cell recording was performed in areas PEc and V6A, where different anatomical tracers were also injected. The monkeys performed a variety of behavioral tasks, aimed at studying the visual and motor properties of parietal cells, as well as the potential combination of retinal-, eye- and hand-related signals on cell activity. The activity of most cells was related to the direction of movement and the active position of the hand. Many of these reach-related cells were influenced by eye position information. Fewer cells displayed relationships to saccadic eye movements. The activity of most neurons related to a combination of both hand and eye signals. Many cells were also modulated during preparation for hand movement. Light-dark differences of activity were common and interpreted as related to the sight and monitoring of hand motion and/or position in the visual field. Most cells studied were very sensitive to moving visual stimuli and also responded to optic flow stimulation. Visual receptive fields were generally large and extended to the periphery of the visual field. For most neurons, the orientation of the preferred directions computed across different epochs and tasks conditions clustered within a limited sector of space, the field of global tuning. This can be regarded as an ideal frame to combine spatially congruent eye- and hand-related information for different forms of visuomanual behavior. All these properties were common to both PEc and V6A. Retinal, eye- and hand-related activity types, as well as parieto-frontal association cells, were distributed in a periodic fashion across the tangential domain of areas PEc and V6A. These functional and anatomical distributions were characterized and compared through a spectral and coherency analysis, which revealed the existence of a selective 'match' between activity types and parieto-frontal connections. This match depended on where each individual efferent projection was addressed. The results of the present and of the companion study can be relevant for a re-interpretation of optic ataxia as the consequence of the breakdown of the combination of retinal-, eye- and hand-related directional signals within the global tuning fields of parietal neurons.     

Visual, somatosensory, and bimodal activities in the macaque parietal area PEc

Caudal area PE (PEc) of the macaque posterior parietal cortex has been shown to be a crucial node in visuomotor coordination during reaching. The present study was aimed at studying visual and somatosensory organization of this cortical area. Visual stimulations activated 53% of PEc neurons. The overwhelming majority (89%) of these visual cells were best activated by a dark stimulus on a lighter background. Somatosensory stimulations activated 56% of PEc neurons: most were joint neurons (73%); a minority (24%) showed tactile receptive fields, most of them located on the arms. Area PEc has not a clear retinotopy or somatotopy. Among the cells tested for both somatosensory and visual sensitivity, 22% were bimodal, 25% unimodal somatosensory, 34% unimodal visual, and 19% were insensitive to either stimulation. No clear clustering of the different classes of sensory neurons was observed. Visual and somatosensory receptive fields of bimodal cells were not in register. The damage in the human brain of the likely homologous of macaque PEc produces deficits in locomotion and in whole-body interaction with the visual environment. Present data show that macaque PEc has sensory properties and a functional organization in line with the view of an involvement of this area in those processes.     

Area 3a neuron response to skin nociceptor afferent drive

Area 3a neurons are identified that respond weakly or not at all to skin contact with a 25-38 degrees C probe, but vigorously to skin contact with the probe at > or =49 degrees C. Maximal rate of spike firing associated with 1- to 7-s contact at > or =49 degrees C occurs 1-2 s after probe removal from the skin. The activity evoked by 5-s contact with the probe at 51 degrees C remains above-background for approximately 20 s after probe retraction. After 1-s contact at 55-56 degrees C activity remains above-background for approximately 4 s. Magnitude of spike firing associated with 5-s contact increases linearly as probe temperature is increased from 49-51 degrees C. Intradermal capsaicin injection elicits a larger (approximately 2.5x) and longer-lasting (100x) increase in area 3a neuron firing rate than 5-s contact at 51 degrees C. Area 3a neurons exhibit enhanced or novel responsivity to 25-38 degrees C contact for a prolonged time after intradermal injection of capsaicin or alpha, beta methylene adenosine triphosphate. Their 1) delayed and persisting increase in spike firing in response to contact at > or =49 degrees C, 2) vigorous and prolonged response to intradermal capsaicin, and 3) enhanced and frequently novel response to 25-38 degrees C contact following intradermal algogen injection or noxious skin heating suggest that the area 3a neurons identified in this study contribute to second pain and mechanical hyperalgesia/allodynia.     

Speed of information processing in traumatic brain injury: modality-specific factors

OBJECTIVE: To assess speed of information processing by two serial addition tests (one visual, one auditory) in individuals with moderate-to-severe traumatic brain injuries (TBIs) and in a healthy, normal control group (NC). The tasks were designed to equate and control for accuracy of performance across the TBI and NC groups, thus allowing for quantification of information processing speed. DESIGN: Performance across groups and tasks were compared using 2 x 2 repeated measure analyses of variance (ANOVAs). In addition, each individual's processing speed was used to adjust rate of stimulus presentation on a subsequent "rehabilitation" trial to determine further whether this adjustment equated accuracy of performance. SETTING: Rehabilitation hospital. PATIENTS: 22 outpatients with moderate-to-severe TBI (6 women, 16 men; mean age = 34.6 years; duration of loss of consciousness = 22.6 days) and 20 age- and education-matched healthy controls. RESULTS: Processing speed was slower in TBI subjects, relative to controls and was significantly related to measures of executive functioning for those with TBI. Relative to controls, speed of processing in the TBI group was disproportionately slower when information was presented in the auditory, relative to the visual, modality. CONCLUSIONS: Speed of information processing is a major impairment in those with TBI when unconfounded by performance accuracy. The modality-specific impairment observed in the TBI group may, in part, be due to a greater within-modality interference effect created by the auditory version of the task. By manipulating information at a pace customized for an individual through compensatory strategies and environmental modifications, information-processing performance of TBI participants can be enhanced significantly.     

Cognitive deficits in patients after soft tissue injury of the cervical spine.

Fifty-one patients suffering from soft tissue injury of the cervical spine underwent clinical and psychometric examination. Clinical interview evaluated subjective complaints and formal testing of self-estimated cognitive impairment, divided attention, and speed of information processing. Results indicated at least two different syndromes: 1) the "cervicoencephalic syndrome," characterized by headache, fatigue, dizziness, poor concentration, disturbed accommodation, and impaired adaptation to light intensity; and 2) the "lower cervical spine syndrome," which is accompanied by cervical and cervicobrachial pain. When comparing patients with either of these two syndromes, those suffering from cervicoencephalic syndrome had significantly poorer results when tested for divided attention. Speed of information processing was reduced to a comparable extent in both syndromes. These findings were not related to the length of the post-traumatic interval. Reduced processing of working memory is assumed, which may account for more global cognitive problems as well as secondary neurotic reaction.     

Assessing bone status beyond BMD: evaluation of bone geometry and porosity by quantitative ultrasound of human finger phalanges.

In an in vitro study, we found significant associations between QUS variables and properties and geometrical parameters of the compact bone of human finger phalanges. QUS variables were not only related to BMD but also to other skeletal properties, which explained 70% of the variability of speed of sound. INTRODUCTION: Transverse transmission quantitative ultrasound (QUS) measurements at the finger phalanges are known to be correlated with BMD and to predict osteoporotic fractures. To determine which other skeletal properties are affected by ultrasound, we investigated the impact of density, geometry, and porosity on QUS variables in vitro. MATERIALS AND METHODS: Ultrasound variables were correlated with density, porosity, and geometrical characteristics of cortical bone. Additionally, we tested which combinations of geometry and bone properties best predicted the ultrasound results observed. Forty-four proximal phalanges from the middle finger were investigated at their distal metaphysis, similar to the typical clinical measurement procedure. Donor age ranged from 52 to 98 years (15 males and 29 females; mean age, 80.9 +/- 9.4 years). QUS variables were measured on the metaphysis of the phalanges using the DBMSonic 1200. Quantitative CT was used for the measurement of BMD, and high-resolution MRI was used for the measurement of porosity and geometrical variables. RESULTS: Speed of sound (SOS) and the clinically used variable AD-SOS correlated significantly with area, relative area, density, and porosity of the compact bone (R2 = 0.28-0.58, p < 0.0001). Signal amplitude correlated significantly only with relative area of the compact bone and area of the medullary canal (R2 = 0.18-0.20, p < 0.01). The combination of cortical area, density, and porosity improved the determination of SOS to R2 = 0.70, with a residual unexplained variability of 54 m/s (3.2%). CONCLUSIONS: These results clarify the impact of skeletal properties on QUS variables. SOS is affected by cortical area, cortical bone density, and cortical porosity, whereas attenuation only depends on geometry of the medulla. AD-SOS, the variable routinely measured in clinical practice, is primarily affected by cortical area. QUS of the finger phalanges is not only related to BMD but also to other skeletal properties.     

Effects of surface cues on macaque inferior temporal cortical responses

Humans are able to recognize objects when surface details, such as colour, texture and luminance gradients, are not available. By systematically eliminating colour, texture, shading, contrast and inner contours from given objects, we tested whether certain shape-selective inferior temporal cortex (IT) neurons of awake rhesus monkeys remain selective for these objects as the surface information is reduced. In psychophysical experiments, we established that the rhesus monkey can identify the shape of a coloured object largely independently of its surface characteristics and, to a lesser degree, of its inner contours. Shape selectivity of the neurons does not change when texture and shading are concealed. The responsiveness of the neurons is also affected by the removal of these surface attributes. The IT neurons were found to respond highly similarly to objects brighter or darker than their background. Selectivity for shape is preserved when the contrast is reversed. Deletion of the inner contours, outlining the main parts of the objects, did not affect the responses and selectivity of the IT neurons. These findings indicate that the IT can contribute to the invariant perception of objects having different surface details.     

Cortical area MSTd combines visual cues to represent 3-D self-movement

As arboreal primates move through the jungle, they are immersed in visual motion that they must distinguish from the movement of predators and prey. We recorded dorsal medial superior temporal (MSTd) cortical neuronal responses to visual motion stimuli simulating self-movement and object motion. MSTd neurons encode the heading of simulated self-movement in three-dimensional (3-D) space. 3-D heading responses can be evoked either by the large patterns of visual motion in optic flow or by the visual object motion seen when an observer passes an earth-fixed landmark. Responses to naturalistically combined optic flow and object motion depend on their relative directions: an object moving as part of the optic flow field has little effect on neuronal responses. In contrast, an object moving separately from the optic flow field has large effects, decreasing the amplitude of the population response and shifting the population's heading estimate to match the direction of object motion as the object moves toward central vision. These effects parallel those seen in human heading perception with minimal effects of objects moving with the optic flow and substantial effects of objects violating the optic flow. We conclude that MSTd can contribute to navigation by supporting 3-D heading estimation, potentially switching from optic flow to object cues when a moving object passes in front of the observer.