The representation of visual salience in monkey parietal cortex.

When natural scenes are viewed, a multitude of objects that are stable in their environments are brought in and out of view by eye movements. The posterior parietal cortex is crucial for the analysis of space, visual attention and movement. Neurons in one of its subdivisions, the lateral intraparietal area (LIP), have visual responses to stimuli appearing abruptly at particular retinal locations (their receptive fields). We have tested the responses of LIP neurons to stimuli that entered their receptive field by saccades. Neurons had little or no response to stimuli brought into their receptive field by saccades, unless the stimuli were behaviourally significant. We established behavioural significance in two ways: either by making a stable stimulus task-relevant, or by taking advantage of the attentional attraction of an abruptly appearing stimulus. Our results show that under ordinary circumstances the entire visual world is only weakly represented in LIP. The visual representation in LIP is sparse, with only the most salient or behaviourally relevant objects being strongly represented.

Intrinsic circuitry and physiological properties of pyramidal neurons in rat barrel cortex.

Pyramidal neurons in the rat posteromedial barrel subfield (PMBSF) were characterized physiologically and filled with biocytin in in vitro brain slices. Intrinsic axons belonging to supragranular neurons projected horizontally and vertically, arborizing in layers II/III and V, but had few or no projections to layers IV or VI. These axons projected horizontally for up to 2 mm, spanning two to seven barrel columns. Layer V neurons had more diffuse axon arbors that projected either vertically, arborizing in layers III to V, or horizontally, branching profusely in layers V and VI. The basal dendritic trees of neurons in layers II/III, V and VI spanned one or two barrel columns without being skewed toward particular barrel columns. Physiologically, regular-spiking neurons were classified as "RS1" or "RS2" according to their degree of late spike frequency adaptation. RS1 neurons were found in superficial and deep layers, whereas RS2 neurons were significantly more prevalent in the latter. Infragranular, but not supragranular neurons showed slow, inward rectification at hyperpolarized potentials. All neurons generated fast and medium afterhyperpolarizations following individual spikes; however, only infragranular pyramids had depolarizing afterpotentials interposed between the two afterhyperpolarizations. RS1 neurons had larger cell bodies, longer total basal dendritic lengths, and more densely branched proximal dendritic trees than RS2 neurons. These findings indicate that pyramidal neurons in the deep and superficial layers of the rat PMBSF have distinct patterns of intracortical axon arbors and distinct physiological properties. These features are probably involved in shaping and modulating the response properties of PMBSF neurons.

Neural responses related to smooth-pursuit eye movements and their correspondence with electrically elicited smooth eye movements in the primate frontal eye field.

1. Intracortical microstimulation of a portion of the monkey frontal eye field (FEF) lying in the floor and posterior bank of the arcuate sulcus evokes smooth, rather than saccadic eye movements. To further explore this region's involvement in pursuit, we recorded from FEF neurons in the vicinity of sites from which smooth eye movements (SEMs) were elicited electrically and studied their responses during smooth-pursuit and saccadic tasks. In this report, we describe the neurons' responses during visually guided smooth pursuit and compare their locations and response properties with those of elicited SEMs. 2. One hundred and ninety-three neurons, recorded from the FEF region in six hemispheres of three rhesus monkeys, were classified as "pursuit neurons". These neurons responded during smooth-pursuit tracking of moving visual stimuli but had no, or only minimal, responses in conjunction with visually guided saccades. Pursuit neurons were located in a small region of the arcuate fundus and posterior bank that overlapped, and extended slightly beyond, the region from which SEMs were elicited with microstimulation. 3. All pursuit neurons had a preferred pursuit direction, and all directions were represented with no strong bias toward ipsilateral, contralateral, up, or down. The directional tuning of 80 pursuit cells was measured quantitatively by testing pursuit in several directions and fitting the responses to a Gaussian function. Tuning indices (the sigma parameter of the Gaussian fit) varied between 13 degrees and 136 degrees. The median tuning index, 44.5 degrees, corresponds to a full width at half maximum of 105 degrees. The ubiquity of selectivity for pursuit direction and the wide distribution of preferred directions indicates that pursuit direction uses a place-code type of representation in FEF; however, the broad directional tuning of most neurons suggests that pursuit direction is given by a weighted average of optimal directions across the population of pursuit neurons active at any given time. 4. In general, the responses of pursuit neurons increased with pursuit velocity. Of 13 neurons formally tested with 2 s of constant-velocity tracking in their preferred direction across a range of target speeds, pursuit velocity sensitivity ranged from 0.24 to 1.42 spikes.s-1.deg-1.s-1, with an average sensitivity of 0.70. This relationship suggests that pursuit neurons represent pursuit magnitude using a rate code; this parallels our previous observation that at most SEM sites, the velocity and acceleration of the electrically elicited eye movements increased as a function of the stimulation current.(ABSTRACT TRUNCATED AT 400 WORDS)

Islands and striosomes in the neostriatum of the rhesus monkey: non-equivalent compartments.

Cytoarchitectonically defined cell-dense islands and regions of low acetylcholinesterase reactivity referred to as striosomes have been regarded as equivalent markers of the non-matrix compartment in the neostriatum. We examined islands and striosomes in adjacent sections to determine the degree of correspondence between the two neostriatal compartmental markers. Islands are aggregated centrally within the caudate, whereas striosomes are located throughout the entire nucleus, including the dorsolateral and ventromedial sectors. Moreover, even within the central sector, striosomes are more prevalent than islands. The present quantitative analysis suggests that islands may be further characterized as acetylcholinesterase-poor since the vast majority of islands co-localize with striosomes. However, due to the fact that striosomes are more numerous and more widely distributed throughout the neostriatum, less than a third of all striosomes are coincident with islands in adjacent sections. Comparison of each of these compartmental markers with the patterned terminal field of the prefrontal cortical projection revealed a near one-to-one correspondence between islands and terminal-free zones in the prefrontal projection. The percentage of striosomes which are aligned with fenestrations in the prefrontal projection is also quite high; however, because more striosomes than islands are found within the prefrontal terminal domain, some striosomes that fit within terminal-free zones do not have corresponding islands. These results indicate that islands and striosomes are not entirely equivalent compartmental markers and further suggest that contemporary, two-compartment models may not adequately represent the heterogeneity of the neostriatum.

Smooth eye movements elicited by microstimulation in the primate frontal eye field.

1. We electrically stimulated the macaque monkey's frontal eye field (FEF) region to localize and to analyze the smooth pursuit eye movement representation. Rhesus monkeys were trained to fixate stationary spots of light, and trains of stimulation (usually 250-500 ms at 10-100 microA) were applied while the fixation targets remained lit and stationary. This paradigm was used in a total of 485 electrode penetrations through the arcuate sulcus region of six hemispheres in three adult monkeys. Smooth eye movements (SEMs), clearly distinct from saccades, were elicited at 86 sites in 53 of these penetrations. These SEMs had an average peak velocity of 11 degrees/s and an average latency of 39 ms. 2. The initial acceleration and peak velocity of elicited SEMs increased with stimulation intensity at any given site. On the other hand, SEM direction was characteristic of a given stimulation site and did not vary with stimulation intensity. These findings indicate that SEM amplitude is coded by the intensity of neural activity, and SEM direction is coded by the location of this activity within the cortex ("rate" vs. "place" codes). 3. SEMs elicited in the presence of a stationary fixation target (closed-loop conditions) typically reached a plateau velocity early in the stimulation and maintained that velocity throughout most of the stimulation train. However, when retinal slip was eliminated by artificially stabilizing the fixation target on the fovea (open-loop conditions), the electrical stimulation caused the eye to accelerate for longer periods and to attain higher velocities than in the closed-loop condition. Eye velocities obtained at the same site in open- and closed-loop conditions diverged from one another approximately 100 ms after SEM onset, consistent with the visual latency of the pursuit system. These findings suggest that the FEF primarily conveys an eye acceleration signal, rather than an eye velocity goal, to the pursuit system, and that this signal can be affected by visual retinal errors before effecting the smooth eye movements. 4. SEMs were elicited from a small portion of the arcuate fundus and neighboring posterior bank lying directly posterior to the principal sulcus. Functionally, this SEM region was surrounded by the saccadic FEF and by somatic premotor cortex. 5. Even though ipsilateral, contralateral, and vertical SEMs were elicited, the distribution of SEM directions was skewed toward ipsilateral movements. This tendency was more pronounced for sites in the arcuate fundus, whereas SEMs elicited from the posterior arcuate bank were often directed contralaterally and vertically.(ABSTRACT TRUNCATED AT 400 WORDS)

Smooth-pursuit eye movement representation in the primate frontal eye field.

Physiological and behavioral data reported here show an involvement of the primate frontal eye field (FEF) cortex in smooth-pursuit eye movements, in addition to its well-established role in saccadic eye movements. Microstimulation just ventral to the small saccade representation of the FEF elicits eye movements that, in contrast to elicited saccades, have low velocities, continue smoothly without interruption during prolonged stimulation, and are usually directed ipsilaterally to the stimulated hemisphere. Neurons in this region respond in association with smooth-pursuit eye movements and visual motion. Tracking deficits following experimental lesions of the FEF depend critically upon the status of this ventral region: superficial lesions sparing it leave smooth-pursuit eye movements intact, whereas lesions removing it produce substantial deficits in the anticipatory initiation, motion-induced acceleration, asymptotic velocity, and predictive continuation of ipsilateral smooth pursuit.