Schluppeck, Denis, Paul Glimcher, and David J. Heeger. Topographic organization for delayed saccades in human posterior parietal cortex.

Training can significantly improve performance on even the most basic visual tasks, such as detecting a faint patch of light or determining the orientation of a bar (for reviews, see ). The neural mechanisms of visual learning, however, remain controversial. One simple way to improve behavior is to increase the overall neural response to the trained stimulus by increasing the number or gain of responsive neurons. Learning of this type has been observed in other sensory modalities, where training increases the number of receptive fields that cover the trained stimulus. Here, we show that visual learning can selectively increase the overall response to trained stimuli in primary visual cortex (V1). We used functional magnetic resonance imaging (fMRI) to measure neural signals before and after one month of practice at detecting very low-contrast oriented patterns. Training increased V1 response for practiced orientations relative to control orientations by an average of 39%, and the magnitude of the change in V1 correlated moderately well with the magnitude of changes in detection performance. The elevation of V1 activity by training likely results from an increase in the number of neurons responding to the trained stimulus or an increase in response gain.

Functional magnetic resonance imaging (fMRI) is now routinely used to map the topographic organization of human visual cortex. Mapping the detailed topography of somatosensory cortex, however, has proven to be more difficult. Here we used the increased blood-oxygen-level-dependent contrast-to-noise ratio at ultra-high field (7 Tesla) to measure the topographic representation of the digits in human somatosensory cortex at 1 mm isotropic resolution in individual subjects. A "traveling wave" paradigm was used to locate regions of cortex responding to periodic tactile stimulation of each distal phalangeal digit. Tactile stimulation was applied sequentially to each digit of the left hand from thumb to little finger (and in the reverse order). In all subjects, we found an orderly map of the digits on the posterior bank of the central sulcus (postcentral gyrus). Additionally, we measured event-related responses to brief stimuli for comparison with the topographic mapping data and related the fMRI responses to anatomical images obtained with an inversion-recovery sequence. Our results have important implications for the study of human somatosensory cortex and underscore the practical utility of ultra-high field functional imaging with 1 mm isotropic resolution for neuroscience experiments. First, topographic mapping of somatosensory cortex can be achieved in 20 min, allowing time for further experiments in the same session. Second, the maps are of sufficiently high resolution to resolve the representations of all five digits and third, the measurements are robust and can be made in an individual subject. These combined advantages will allow somatotopic fMRI to be used to measure the representation of digits in patients undergoing rehabilitation or plastic changes after peripheral nerve damage as well as tracking changes in normal subjects undergoing perceptual learning.

In a previous study, we identified three cortical areas in human posterior parietal cortex that exhibited topographic responses during memory-guided saccades [visual area 7 (V7), intraparietal sulcus 1 (IPS1), and IPS2], which are candidate homologs of macaque parietal areas such as the lateral intraparietal area and parietal reach region. Here, we show that these areas exhibit sustained delay-period activity, a critical physiological signature of areas in macaque parietal cortex. By varying delay duration, we disambiguated delay-period activity from sensory and motor responses. Mean time courses in the parietal areas were well fit by a linear model comprising three components representing responses to (1) the visual target, (2) the delay period, and (3) the eye movement interval. We estimated the contributions of each component: the response amplitude during the delay period was substantially smaller (Ͻ30%) than that elicited by the transient visual target. All three parietal regions showed comparable delay-period response amplitudes, with a trend toward larger responses from V7 to IPS1 and IPS2. Responses to the cue and during the delay period showed clear lateralization with larger responses to trials in which the target was placed in the contralateral visual field, suggesting that both of these components contributed to the topography we measured.

Electrophysiological studies in monkeys have identified effector-related regions in the posterior parietal cortex (PPC). The lateral intraparietal area, for example, responds preferentially for saccades, whereas the parietal reach region responds preferentially for arm movements. However, the degree of effector selectivity actually observed is limited; each area contains neurons selective for the nonpreferred effector, and many neurons in both areas respond for both effectors. We used functional magnetic resonance imaging to assess the degree of effector preference at the population level, focusing on topographically organized regions in the human PPC [visual area V7, intraparietal sulcus 1 (IPS1), and IPS2]. An event-related design adapted from monkey experiments was used. In each trial, an effector cue preceded the appearance of a spatial target, after which a Go signal instructed subjects to produce the specified movement with the specified effector. Our results show that the degree of effector specificity is limited in many cortical areas and transitions gradually from saccade to reach preference as one moves through the hierarchy of areas in the occipital, parietal, and frontal cortices. Saccade preference was observed in visual cortex, including early areas and V7. IPS1 exhibited balanced activation to saccades and reaches, whereas IPS2 showed a weak but significant preference for reaches. In frontal cortex, areas near the central sulcus showed a clear and absolute preference for reaches, whereas the frontal eye field showed little or no effector selectivity. Although these results contradict many theoretical conclusions about effector specificity, they are compatible with the complex picture arising from electrophysiological studies and also with previous imaging studies that reported mostly overlapping saccade-and arm-related activation. The results are also compatible with theories of efficient coding in cortex.