Topographic plasticity in primary visual cortex is mediated by local corticocortical connections

The placement of monocular laser lesions in the adult cat retina produces a lesion projection zone (LPZ) in primary visual cortex (V1) in which the majority of neurons have a normally located receptive field (RF) for stimulation of the intact eye and an ectopically located RF ( displaced to intact retina at the edge of the lesion) for stimulation of the lesioned eye. Animals that had such lesions for 14 - 85 d were studied under halothane and nitrous oxide anesthesia with conventional neurophysiological recording techniques and stimulation of moving light bars. Previous work suggested that a candidate source of input, which could account for the development of the ectopic RFs, was long-range horizontal connections within V1. The critical contribution of such input was examined by placing a pipette containing the neurotoxin kainic acid at a site in the normal V1 visual representation that overlapped with the ectopic RF recorded at a site within the LPZ. Continuation of well defined responses to stimulation of the intact eye served as a control against direct effects of the kainic acid at the LPZ recording site. In six of seven cases examined, kainic acid deactivation of neurons at the injection site blocked responsiveness to lesioned-eye stimulation at the ectopic RF for the LPZ recording site. We therefore conclude that long-range horizontal projections contribute to the dominant input underlying the capacity for retinal lesion-induced plasticity in V1

Comparison of level discrimination, increment detection, and comodulation masking release in the audio- and envelope-frequency domains

In general, the temporal structure of stimuli must be considered to account for certain observations made in detection and masking experiments in the audio-frequency domain. Two such phenomena are (1) a heightened sensitivity to amplitude increments with a temporal fringe compared to gated level discrimination performance and (2) lower tone-in-noise detection thresholds using a modulated masker compared to those using an unmodulated masker. In the current study, translations of these two experiments were carried out to test the hypothesis that analogous cues might be used in the envelope-frequency domain. Pure-tone carrier amplitude-modulation (AM) depth-discrimination thresholds were found to be similar using both traditional gated stimuli and using a temporally modulated fringe for a fixed standard depth (m(s)=0.25) and a range of AM frequencies (4-64 Hz). In a second experiment, masked sinusoidal AM detection thresholds were compared in conditions with and without slow and regular fluctuations imposed on the instantaneous masker AM depth. Release from masking was obtained only for very slow masker fluctuations (less than 2 Hz). A physiologically motivated model that effectively acts as a first-order envelope change detector accounted for several, but not all, of the key aspects of the data

The effects of immediate vision on implicit hand maps

Perceiving the external spatial location of the limbs using position sense requires that immediate proprioceptive afferent signals be combined with a stored body model specifying the size and shape of the body. Longo and Haggard (Proc Natl Acad Sci USA 107:11727–11732, 2010) developed a method to isolate and measure this body model in the case of the hand in which participants judge the perceived location in external space of several landmarks on their occluded hand. The spatial layout of judgments of different landmarks is used to construct implicit hand maps, which can then be compared with actual hand shape. Studies using this paradigm have revealed that the body model of the hand is massively distorted, in a highly stereotyped way across individuals, with large underestimation of finger length and overestimation of hand width. Previous studies using this paradigm have allowed participants to see the locations of their judgments on the occluding board. Several previous studies have demonstrated that immediate vision, even when wholly non-informative, can alter processing of somatosensory signals and alter the reference frame in which they are localised. The present study therefore investigated whether immediate vision contributes to the distortions of implicit hand maps described previously. Participants judged the external spatial location of the tips and knuckles of their occluded left hand either while being able to see where they were pointing (as in previous studies) or while blindfolded. The characteristic distortions of implicit hand maps reported previously were clearly apparent in both conditions, demonstrating that the distortions are not an artefact of immediate vision. However, there were significant differences in the magnitude of distortions in the two conditions, suggesting that vision may modulate representations of body size and shape, even when entirely non-informative

The Illusion of Owning a Third Arm

Could it be possible that, in the not-so-distant future, we will be able to reshape the human body so as to have extra limbs? A third arm helping us out with the weekly shopping in the local grocery store, or an extra artificial limb assisting a paralysed person? Here we report a perceptual illusion in which a rubber right hand, placed beside the real hand in full view of the participant, is perceived as a supernumerary limb belonging to the participant's own body. This effect was supported by questionnaire data in conjunction with physiological evidence obtained from skin conductance responses when physically threatening either the rubber hand or the real one. In four well-controlled experiments, we demonstrate the minimal required conditions for the elicitation of this “supernumerary hand illusion”. In the fifth, and final experiment, we show that the illusion reported here is qualitatively different from the traditional rubber hand illusion as it is characterised by less disownership of the real hand and a stronger feeling of having two right hands. These results suggest that the artificial hand ‘borrows’ some of the multisensory processes that represent the real hand, leading to duplication of touch and ownership of two right arms. This work represents a major advance because it challenges the traditional view of the gross morphology of the human body as a fundamental constraint on what we can come to experience as our physical self, by showing that the body representation can easily be updated to incorporate an additional limb

Forward Masking Estimated by Signal Detection Theory Analysis of Neuronal Responses in Primary Auditory Cortex

Psychophysical forward masking is an increase in threshold of detection of a sound (probe) when it is preceded by another sound (masker). This is reminiscent of the reduction in neuronal responses to a sound following prior stimulation. Studies in the auditory nerve and cochlear nucleus using signal detection theory techniques to derive neuronal thresholds showed that in centrally projecting neurons, increases in masked thresholds were significantly smaller than the changes measured psychophysically. Larger threshold shifts have been reported in the inferior colliculus of awake marmoset. The present study investigated the magnitude of forward masking in primary auditory cortical neurons of anaesthetised guinea-pigs. Responses of cortical neurons to unmasked and forward masked tones were measured and probe detection thresholds estimated using signal detection theory methods. Threshold shifts were larger than in the auditory nerve, cochlear nucleus and inferior colliculus. The larger threshold shifts suggest that central, and probably cortical, processes contribute to forward masking. However, although methodological differences make comparisons difficult, the threshold shifts in cortical neurons were, in contrast to subcortical nuclei, actually larger than those observed psychophysically. Masking was largely attributable to a reduction in the responses to the probe, rather than either a persistence of the masker responses or an increase in the variability of probe responses

Perinatal Asphyxia Affects Rat Auditory Processing: Implications for Auditory Perceptual Impairments in Neurodevelopmental Disorders

Perinatal asphyxia, a naturally and commonly occurring risk factor in birthing, represents one of the major causes of neonatal encephalopathy with long term consequences for infants. Here, degraded spectral and temporal responses to sounds were recorded from neurons in the primary auditory cortex (A1) of adult rats exposed to asphyxia at birth. Response onset latencies and durations were increased. Response amplitudes were reduced. Tuning curves were broader. Degraded successive-stimulus masking inhibitory mechanisms were associated with a reduced capability of neurons to follow higher-rate repetitive stimuli. The architecture of peripheral inner ear sensory epithelium was preserved, suggesting that recorded abnormalities can be of central origin. Some implications of these findings for the genesis of language perception deficits or for impaired language expression recorded in developmental disorders, such as autism spectrum disorders, contributed to by perinatal asphyxia, are discussed

Spectrotemporal Processing in Spectral Tuning Modules of Cat Primary Auditory Cortex

Spectral integration properties show topographical order in cat primary auditory cortex (AI). Along the iso-frequency domain, regions with predominantly narrowly tuned (NT) neurons are segregated from regions with more broadly tuned (BT) neurons, forming distinct processing modules. Despite their prominent spatial segregation, spectrotemporal processing has not been compared for these regions. We identified these NT and BT regions with broad-band ripple stimuli and characterized processing differences between them using both spectrotemporal receptive fields (STRFs) and nonlinear stimulus/firing rate transformations. The durations of STRF excitatory and inhibitory subfields were shorter and the best temporal modulation frequencies were higher for BT neurons than for NT neurons. For NT neurons, the bandwidth of excitatory and inhibitory subfields was matched, whereas for BT neurons it was not. Phase locking and feature selectivity were higher for NT neurons. Properties of the nonlinearities showed only slight differences across the bandwidth modules. These results indicate fundamental differences in spectrotemporal preferences - and thus distinct physiological functions - for neurons in BT and NT spectral integration modules. However, some global processing aspects, such as spectrotemporal interactions and nonlinear input/output behavior, appear to be similar for both neuronal subgroups. The findings suggest that spectral integration modules in AI differ in what specific stimulus aspects are processed, but they are similar in the manner in which stimulus information is processed

Ascending projections to the medial geniculate-body of the cat - evidence for multiple, parallel auditory pathways through thalamus

The sources of ascending input to the medial geniculate body (MGB) of the cat were studied using the retrograde transport of horseradish peroxidase (HRP). HRP injections were made iontophoretically through micropipettes which were also used to record physiological properties at the injection sites. This technique produced small injections which appeared to be restricted to single subnuclei. The tectothalamic projection of the auditory system was found to consist of at least four distinct and separate pathways. The ventral division of the MGB receives a topographical projection from the central nucleus of the inferior colliculus (ICC) which preserves tonotopicity and provides short latency, sharply frequency-tuned responses. The medial part of the ICC projects to the deep dorsal nucleus, which contains only units tuned to high frequencies. The major inputs to the caudodorsal nucleus (DC) stem from nucleus sagulum and the pericentral nucleus of the inferior colliculus (ICP). Units in DC and the ventrolateral nucleus, which also receive input from ICP, have very broad tuning properties and late, habituating responses. Injections of HRP into the medial division (MGM) produced labeled cells scattered throughout the external nucleus of the inferior colliculus and the ventral part of ICC. This widespread input is reflected in the wide range of auditory responses found in MGM. Auditory responses in the suprageniculate nucleus were poorly defined and many units did not respond to tonal stimuli; following HRP injections no filled cells were found in the inferior colliculus, but labeled cells were found in the deeper layers of the superior colliculus and in the interstitial nucleus of the brachium of the inferior colliculus. Together with recent findings on the auditory thalamocortical projection, these results provide evidence for multiple parallel auditory pathways through the thalamus

Dynamic representational plasticity in sensory cortex

Studies of the effects of peripheral and central lesions, perceptual learning and neurochemical modification on the sensory representations in cortex have had a dramatic effect in alerting neuroscientists and therapists to the reorganizational capacity of the adult brain. An intriguing aspect of some of these investigations, such as partial peripheral denervation, is the short-term expression of these changes. Indeed, in visual cortex, auditory cortex and somatosensory cortex loss of input from a region of the peripheral receptor epithelium (retinal, basilar and cutaneous, respectively) induces rapid expression of ectopic, or expanded, receptive fields of affected neurons and reorganization of topographic maps to fill in the representation of the denervated area. The extent of these changes can, in some cases, match the maximal extents demonstrated with chronic manipulations. The rapidity, and reversibility, of the effects rules out many possible explanations which involve synaptic plasticity and points to a capacity for representational plasticity being inherent in the circuitry of a topographic pathway. Consequently, topographic representations must be considered as manifestations of physiological interaction rather than as anatomical constructs. Interference with this interaction can produce an unmasking of previously inhibited responsiveness. Consideration of the nature of masking inhibition which is consistent with the precision and order of a topographic representation and which has a capacity for rapid plasticity requires, in addition to stimulus-driven inhibition, a source of tonic input from the periphery. Such input, acting locally to provide tonic inhibition, has been directly demonstrated in the somatosensory system and is consistent with results obtained in auditory and visual systems

Limits on short-term plasticity in somatosensory cortex

It is now clear, from a large number of studies, that disruption of a subset of the inputs to the sensory representations in cortex of adult manunals induces a rapid unmasking of larger neuronal receptive fields and an increased neuronal responsiveness. Within the somatosensory cortex, such demonstrations indicate that a capacity for short-term plasticity results from the unmasking of normally unexpressed inputs to a cortical locus. Despite the clear demonstration of such unmasking, an explanation for how this occurs is not immediately apparent. Aside from the studies directly demonstrating short-term plasticity, there have been anatomical (Landry and Deschenes, 1981; Garraghty and Sur, 1990), physiological (Snow et aI., 1988) and pharmacological (e.g. Dykes et al., 1984) demonstrations of cortical afferents that are not normally expressed as part of neural receptive fields. The rapid unmasking phenomena indicate that these inputs are inherently viable and thus must normally be actively suppressed. The only candidate source for an inhibition which can account for this suppression is that produced by the GABAergic interneurons located within a given cortical area. This short review will examine the implications that studies of short-term plasticity require for the nature of that inhibition, and the limitations of the capacity for plasticity that is dependent upon these normally unexpressed inputs