Spatial distribution of responses to simple and complex sounds in the primary auditory cortex

The basic functional organization of the cat primary auditory cortex is discussed as it is revealed by electrophysiological studies of the distribution of elementary receptive field (RF) parameters. RFs of cortical neurons have been shown to vary considerably from neuron to neuron; additionally, specific RF properties vary independently. Furthermore, some of the RF properties are nonhomogeneously distributed across the auditory cortex and can be interpreted as forming "maps" that represent specific stimulus information in a topographic way. Accordingly, the functional organization of the primary auditory cortex is interpreted as a series of superimposed independent parameter maps. The consequences of such a layout for the spatial and temporal coding of pure tones and speech sounds is illustrated and ramifications for the interpretation of far-field event-related potentials are discussed.     

Physiological memory in primary auditory cortex: characteristics and mechanisms

"Physiological memory" is enduring neuronal change sufficiently specific to represent learned information. It transcends both sensory traces that are detailed but transient and long-term physiological plasticities that are insufficiently specific to actually represent cardinal details of an experience. The specificity of most physiological plasticities has not been comprehensively studied. We adopted receptive field analysis from sensory physiology to seek physiological memory in the primary auditory cortex of adult guinea pigs. Receptive fields for acoustic frequency were determined before and at various retention intervals after a learning experience, typified by single-tone delay classical conditioning, e.g., 30 trials of tone-shock pairing. Subjects rapidly (5-10 trials) acquire behavioral fear conditioned responses, indexing acquisition of an association between the conditioned and the unconditioned stimuli. Such stimulus-stimulus association produces receptive field plasticity in which responses to the conditioned stimulus frequency are increased in contrast to responses to other frequencies which are decreased, resulting in a shift of tuning toward or to the frequency of the conditioned stimulus. This receptive field plasticity is associative, highly specific, acquired within a few trials, and retained indefinitely (tested to 8 weeks). It thus meets criteria for "physiological memory." The acquired importance of the conditioned stimulus is thought to be represented by the increase in tuning to this stimulus during learning, both within cells and across the primary auditory cortex. Further, receptive field plasticity develops in several tasks, one-tone and two-tone discriminative classical and instrumental conditioning (habituation produces a frequency-specific decrease in the receptive field), suggesting it as a general process for representing the acquired meaning of a signal stimulus. We have proposed a two-stage model involving convergence of the conditioned and unconditioned stimuli in the magnocellular medial geniculate of the thalamus followed by activation of the nucleus basalis, which in turn releases acetylcholine that engages muscarinic receptors in the auditory cortex. This model is supported by several recent findings. For example, tone paired with NB stimulation induces associative, specific receptive field plasticity of at least a 24-h duration. We propose that physiological memory in auditory cortex is not "procedural" memory, i.e., is not tied to any behavioral conditioned response, but can be used flexibly.     

Auditory spatial tuning of cortical neurons is sharpened in cats with early blindness

1. The specificity for the location of a sound source in azimuth was measured in single neurons of the anterior ectosylvian (AE) region of the cat's cortex, which includes the anterior auditory field (AAF) and the anterior ectosylvian auditory field (AEA). 2. The influence of visual experience on auditory spatial tuning of these neurons was determined by comparing responses in cats with binocular deprivation from birth with those in normal control cats. 3. Spatial tuning was measured under near free-field conditions by presenting broadband sounds through a speaker in seven different azimuthal locations, from -60 to +60 degree at 20 degree intervals. Elevation was constant at the cats' ears. 4. In normal cats, a little over one-half of the neurons in the AE region (82/146 = 56%) showed some degree of azimuthal spatial tuning, as defined by at least a 2:1 ratio of responses between best and worst location. The rest (44%) were omnidirectional. 5. In binocularly deprived cats, a significantly higher proportion (70/82 = 86%) of the neurons in the AE region were spatially tuned. Only 14% were omnidirectional. Median spatial tuning width was significantly sharper than in normal cats. 6. We conclude that visual deprivation from birth induces intermodal changes that enhance the response specificity of neurons in the auditory cortex. These modifications may constitute the neural basis of behavioral compensation for early blindness.     

Cortex controls multisensory depression in superior colliculus

Multisensory depression is a fundamental index of multisensory integration in superior colliculus (SC) neurons. It is initiated when one sensory stimulus (auditory) located outside its modality-specific receptive field degrades or eliminates the neuron's responses to another sensory stimulus (visual) presented within its modality-specific receptive field. The present experiments demonstrate that the capacity of SC neurons to engage in multisensory depression is strongly dependent on influences from two cortical areas (the anterior ectosylvian and rostral lateral suprasylvian sulci). When these cortices are deactivated, the ability of SC neurons to synthesize visual-auditory inputs in this way is compromised; multisensory responses are disinhibited, becoming more vigorous and in some cases indistinguishable from responses to the visual stimulus alone. Although obtaining a more robust multisensory SC response when cortex is nonfunctional than when it is functional may seem paradoxical, these data may help explain previous observations that the loss of these cortical influences permits visual orientation behavior in the presence of a normally disruptive auditory stimulus.     

Multimodal integration for the representation of space in the posterior parietal cortex

The posterior parietal cortex has long been considered an 'association' area that combines information from different sensory modalities to form a cognitive representation of space. However, until recently little has been known about the neural mechanisms responsible for this important cognitive process. Recent experiments from the author's laboratory indicate that visual, somatosensory, auditory and vestibular signals are combined in areas LIP and 7a of the posterior parietal cortex. The integration of these signals can represent the locations of stimuli with respect to the observer and within the environment. Area MSTd combines visual motion signals, similar to those generated during an observer's movement through the environment, with eye-movement and vestibular signals. This integration appears to play a role in specifying the path on which the observer is moving. All three cortical areas combine different modalities into common spatial frames by using a gain-field mechanism. The spatial representations in areas LIP and 7a appear to be important for specifying the locations of targets for actions such as eye movements or reaching; the spatial representation within area MSTd appears to be important for navigation and the perceptual stability of motion signals.     

Involvement of cyclic GMP in the mode of action of a new antithrombotic agent PCA-4230; inhibition of the platelet cyclic GMP dependent phosphodiesterase

The division of the auditory cortex into various fields, functional aspects of these fields, and neuronal coding in the primary auditory cortical field (AI) are reviewed with stress on features that may be common to mammals. On the basis of 14 topographies and clustered distributions of neuronal response characteristics in the primary auditory cortical field, a hypothesis is developed of how a certain complex acoustic pattern may be encoded in an equivalent spatial activity pattern in AI, generated by time-coordinated firing of groups of neurons. The auditory cortex, demonstrated specifically for AI, appears to perform sound analysis by synthesis, i.e. by combining spatially distributed coincident or time-coordinated neuronal responses. The dynamics of sounds and the plasticity of cortical responses are considered as a topic for research.     

Auditory motion induces directionally dependent receptive field shifts in inferior colliculus neurons

This research focused on the response of neurons in the inferior colliculus of the unanesthetized mustached bat, Pteronotus parnelli, to apparent auditory motion. We produced the apparent motion stimulus by broadcasting pure-tone bursts sequentially from an array of loudspeakers along horizontal, vertical, or oblique trajectories in the frontal hemifield. Motion direction had an effect on the response of 65% of the units sampled. In these cells, motion in opposite directions produced shifts in receptive field locations, differences in response magnitude, or a combination of the two effects. Receptive fields typically were shifted opposite the direction of motion (i.e., units showed a greater response to moving sounds entering the receptive field than exiting) and shifts were obtained to horizontal, vertical, and oblique motion orientations. Response latency also shifted as a function of motion direction, and stimulus locations eliciting greater spike counts also exhibited the shortest neural latency. Motion crossing the receptive field boundaries appeared to be both necessary and sufficient to produce receptive field shifts. Decreasing the silent interval between successive stimuli in the apparent motion sequence increased both the probability of obtaining a directional effect and the magnitude of receptive field shifts. We suggest that the observed directional effects might be explained by "spatial masking," where the response of auditory neurons after stimulation from particularly effective locations in space would be diminished. The shift in auditory receptive fields would be expected to shift the perceived location of a moving sound and may explain shifts in localization of moving sources observed in psychophysical studies. Shifts in perceived target location caused by auditory motion might be exploited by auditory predators such as Pteronotus in a predictive tracking strategy to capture moving insect prey.     

Two visual areas located in the middle suprasylvian gyrus (cytoarchitectonic field 7) of the cat's cortex

Neuronal properties and topographic organization of the middle suprasylvian gyrus (cortical cytoarchitectonic field 7) were studied in three behaving cats with painlessly fixed heads. Two main neuronal types were found within this field. Type 1 neurons occupied the lateral part of the field and bordered representation of directionally selective neurons of the lateral suprasylvian visual area by vertical retinal meridian. Type 1 neurons had elongated and radially oriented receptive fields located in the lower part of contralateral visual field. Type 1 neurons preferred stimuli moving out or to the centre of gaze at a low or moderate speed, and many of them were depth selective. The responses were enhanced by attention, oriented to the presented stimulus. Medial part of the field 7 along the border with the area V3 was occupied by neurons with not elongated receptive fields (type 2). These neurons preferred moderate and high speeds of motion, and gratings of proper spatial frequency and orientation were effective stimuli for them. Border between representations of type 2 and type 1 neurons coincided with projection of horizontal retinal meridian. At the rostral and caudal borders of the field 7 abrupt changes of neuronal properties took place. Neurons which abutted field 7 anteriorly and posteriorly resembled hypercomplex cells and their small receptive fields were located in the central part of the visual field. Topographical considerations and receptive field properties allowed us to conclude that the medial part of the field 7 (included type 2 neurons) is functionally equivalent to the area V4 in the cortex of primates, while the lateral part (type 1 neurons) may correspond to the area V4T.     

Cortical control of reaching movements

Recent studies provide further support for the hypothesis that spatial representations of limb position, target locations, and potential motor actions are expressed in the neuronal activity in parietal cortex. In contrast, precentral cortical activity more strongly expresses processes involved in the selection and execution of motor actions. As a general conceptual framework, these processes may be interpreted in terms of such formalisms as sensorimotor transformations and 'internal models'.     

A neuronal representation of the location of nearby sounds

Humans can accurately perceive the location of a sound source-not only the direction, but also the distance. Sounds near the head, within ducking or reaching distance, have a special saliency. However, little is known about this perception of auditory distance. The direction to a sound source can be determined by interaural differences, and the mechanisms of direction perception have been studied intensively; but except for studies on echolocation in the bat, little is known about how neurons encode information on auditory distance. Here we describe neurons in the brain of macaque monkeys (Macaca fascicularis) that represent the auditory space surrounding the head, within roughly 30 cm. These neurons, which are located in the ventral premotor cortex, have spatial receptive fields that extend a limited distance outward from the head.     

Image processing in the primary visual cortex

INTRODUCTION: Area 17 or the primary visual area forms the first link in the chain of cerebral analysis of a visual image. The neurones forming the primary visual cortex are characterized by the extreme precision of their connections, functional specialization and hierarchic organization. The spatial precision of the connections within the system for vision permit retinotopic representation in the visual cortex, so that each point of the retina is projected into a specific area of the cortex. The cortical neurones which analyze the characteristics of the image situated in a precise zone of the visual field are themselves organized into a basic functional unit known as a hypercolumn. Within each hypercolumn there are various columnar cell systems with receptive fields having similar characteristics. Thus, each hypercolumn is made up of multiple orientation columns, two ocular dominance columns and 'blob' regions. All these systems permit the analysis of different aspects of the image. The neurones belonging to the orientation columns are sensitive to the orientation, spatial frequency and movement of a visual stimulus; those of the 'blob' regions to colour, and the binocular neurones of the ocular dominance columns to depth. Within each column, the hierarchical pattern of neurone interconnections determines the successive appearance of cells with receptive fields having new properties.     

Capacity for plasticity in the adult owl auditory system expanded by juvenile experience

In the process of creating a multimodal map of space, auditory-visual neurons in the optic tectum establish associations between particular values of auditory spatial cues and locations in the visual field. In the barn owl, tectal neurons reveal these associations in the match between their tuning for interaural time differences (ITDs) and the locations of their visual receptive fields (VRFs). In young owls ITD-VRF associations can be adjusted by experience over a wide range, but the range of adjustment normally becomes quite restricted in adults. This normal range of adjustment in adults was greatly expanded in owls that had previously learned abnormal ITD-VRF associations as juveniles. Thus, the act of learning abnormal associations early in life leaves an enduring trace in this pathway that enables unusual functional connections to be reestablished, as needed, in adulthood, even when the associations represented by these connections have not been used for an extended period of time.     

Heterosynaptic long-term facilitation of sensory-evoked responses in the auditory cortex by stimulation of the magnocellular medial geniculate body in guinea pigs.

The magnocellular nucleus of the medial geniculate body (MGm) develops physiological plasticity during classical conditioning and may be involved in learning-induced receptive field plasticity in the auditory cortex. To determine the ability of the MGm to produce long-term modification of evoked activity in the auditory cortex, the experimenters paired electrical stimulation of the MGm with preceding clicks in adult guinea pigs under barbiturate anesthesia. The amplitudes of average click-evoked potentials were significantly facilitated in all Ss. Facilitation endured for 2 hrs, the maximum duration of recording. Sham-stimulated control guinea pigs did not develop facilitation. Thus, a nonlemniscal thalamic sensory nucleus can produce enduring facilitation of sensory-evoked activity in primary sensory cortex, suggesting that long-term physiological plasticity in the sensory cortex during learning may involve nonlemniscal thalamic mechanisms. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Functional organization of the auditory cortex in a native Chilean rodent (Octodon degus)

The tonotopic organization of primary auditory cortex (AI) and surrounding secondary regions has been studied in the Octodon degus using standard microelectrode mapping techniques. The results confirm and extend previous observations made in other species. The tonotopic organization of the largest field (AI) apparently covered the hearing range of O. degus. Low tonal frequencies were represented rostroventrally and high frequencies caudally, with isofrequency contours orientated dorsoventrally in a ventrocaudal slant. There were additional tonotopic representations adjacent to AI. Rostral to AI, a small field with a tonotopic gradient reversed with respect to that in AI (mirror image representation) was mapped and termed rostral auditory field (R). Best frequencies (BF's) in a range from 0.1-30.0 kHz were found in AI and R, with higher spatial resolution for the representation of lower BF's up to 10.0 kHz. Responses obtained in AI as well as in R were strong, with narrow tuning and short latencies. Caudal to AI, two small additional, tonotopically organized fields, the dorsoposterior field (DP) and the ventroposterior field (VP), could be distinguished. In fields VP and DP, high BF's were situated rostrally, adjacent to the high frequency representation in AI. Low frequency representations were found in caudal part of DP and VP fields. Responses to tone burst within DP and VP were mostly weak, with longer latencies and broader tuning compared to those found in AI and R.     

Prefrontal connections of the parabelt auditory cortex in macaque monkeys

In the present study, we determined connections of three newly defined regions of auditory cortex with regions of the frontal lobe, and how two of these regions in the frontal lobe interconnect and connect to other portions of frontal cortex and the temporal lobe in macaque monkeys. We conceptualize auditory cortex as including a core of primary areas, a surrounding belt of auditory areas, a lateral parabelt of two divisions, and adjoining regions of temporal cortex with parabelt connections. Injections of several different fluorescent tracers and wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP) were placed in caudal (CPB) and rostral (RPB) divisions of the parabelt, and in cortex of the superior temporal gyrus rostral to the parabelt with parabelt connections (STGr). Injections were also placed in two regions of the frontal lobe that were labeled by a parabelt injection in the same case. The results lead to several major conclusions. First, CPB injections label many neurons in dorsal prearcuate cortex in the region of the frontal eye field and neurons in dorsal prefrontal cortex of the principal sulcus, but few or no neurons in orbitofrontal cortex. Fine-grain label in these same regions as a result of a WGA-HRP injection suggests that the connections are reciprocal. Second, RPB injections label overlapping prearcuate and principal sulcus locations, as well as more rostral cortex of the principal sulcus, and several locations in orbitofrontal cortex. Third, STGr injections label locations in orbitofrontal cortex, some of which overlap those of RPB injections, but not prearcuate or principal sulcus locations. Fourth, injections in prearcuate and principal sulcus locations labeled by a CPB injection labeled neurons in CPB and RPB, with little involvement of the auditory belt and no involvement of the core. In addition, the results indicated that the two frontal lobe regions are densely interconnected. They also connect with largely separate regions of the frontal pole and more medial premotor and dorsal prefrontal cortex, but not with the extensive orbitofrontal region which has RPB and STGr connections. The results suggest that both RPB and CPB provide the major auditory connections with the region related to directing eye movements towards stimuli of interest, and the dorsal prefrontal cortex for working memory. Other auditory connections to these regions of the frontal lobe appear to be minor. RPB has connections with orbitofrontal cortex, important in psychosocial and emotional functions, while STGr primarily connects with orbital and polar prefrontal cortex.     

Rapid development of learning-induced receptive field plasticity in the auditory cortex.

Classical conditioning induces frequency-specific receptive field (RF) plasticity in the auditory cortex after relatively brief training (30 trials), characterized by increased response to the frequency of the CS and decreased responses to other frequencies, including the pretraining best frequency (BF). This experiment determined the development of this CS-specific RF plasticity. Guinea pigs underwent classical conditioning to a tonal frequency, and receptive fields of neurons in the auditory cortex were determined before and after 5, 15, and 30 CS–UCS (unconditioned stimulus) pairings, as well as 1 hr posttraining. Highly selective RF changes were observed as early as the first 5 training trials. They culminated after 15 trials, then stabilized after 30 trials and 1 hr posttraining. The rapid development of RF plasticity satisfies a criterion for its involvement in the neural bases of a specific associative memory. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Tonotopic organization of auditory cortical fields delineated by parvalbumin immunoreactivity in macaque monkeys

Tonotopic maps, obtained from single and multi-unit recordings in the primary and surrounding areas of the auditory cortex, were related to chemoarchitecture of the supratemporal plane, as delineated by immunoreactivity for parvalbumin. Neurons in the central core were sharply tuned and formed two complete tonotopic representations corresponding to the primary auditory area (AI) and the rostral (R) area. High frequencies were represented posteriorly in AI and anteriorly in R, the representation reversing in the anterior part of the core. Neurons in regions of less dense immunostaining previously described as lateral (L) and posteromedial (P-m) fields, showed broader frequency tuning. Two tonotopic representations were found in L: in an anterolateral (AL) field, corresponding to a field previously reported by others, high frequencies were represented anteriorly and low frequencies posteriorly; in a posterolateral field (PL) the trend reversed. There was a further reversal on entering P-m from the high frequency representation in PL and progressively lower frequencies tended to be represented more medially in P-m, but P-m may contain two representations reported by others. Neurons in the previously described anteromedial (A-m) and medial (M) fields of weaker immunostaining, were even more broadly tuned. A tonotopic progression from low frequency representation posteriorly to high frequency representation anteriorly was observed in the medial field. Frequency representation in A-m remains uncertain. No tonotopic representation could be demonstrated with the stimuli used in the zones of very weak parvalbumin immunostaining outside AL, PL, P-m, A-m, and M. The properties of neurons in the core and surrounding zones are likely to reflect inputs from the ventral and dorsal medial geniculate nuclei, respectively. The fields outside the core seem to be the starting points for separate streams of auditory corticocortical connections passing into association cortex.     

Anatomic organization of evoked potentials in rat parietotemporal cortex: somatosensory and auditory responses

1. Two 8 x 8-channel microelectrode arrays were used to map epicortical field potentials from a 3.5 x 3.5-mm2 area in homologous regions of right and left parietotemporal cortex of four rats. Potentials were evoked with bilaterally presented click stimuli and with bilateral tactile stimulation of the 25 major vibrissae. The spatial distribution of temporal components of the somatosensory evoked potential (SEP) and auditory evoked potential (AEP) complex were compared directly with cytochrome oxidase-stained sections of the recorded region. 2. Epicortical responses in both hemispheres to bilateral vibrissal stimuli consisted of a biphasic sharp wave (P1a-N1) constrained to the vibrissa/barrel granular region of primary somatosensory cortex (SmI). A slightly later sharp positive wave (P1b) was localized to secondary somatosensory cortex (SmII) and to perigranular cortex medial to the vibrissa/barrel field. The SEP complex ended with a biphasic slow wave (P2-N2). The P2 was centered on SmI and spread to dysgranular lateral cortex, caudal to but excluding SmII. The N2 was centered on SmII and spread to dysgranular cortex caudal to but excluding SmI. 3. The anatomic organization of the AEP in many ways approximated that of the SEP in the same animals. The timing and morphology of the AEP were nearly identical to the SEP. The AEP consisted of a P1a-N1 sharp wave constrained to the estimated region of primary auditory cortex (AI) in the lateral parietotemporal region, a later P1b localized to secondary auditory cortex (AII), and subsequent slow waves (P2 and N2) that were centered on AI and AII, respectively, and spread to dysgranular regions overlapping the distributions of the P2 and N2 of the SEP complex. 4. These data suggest that the basic neural generators for the SEP and AEP in parietotemporal cortex are quite similar, and provide evidence for the functional anatomy of each temporal component of the sensory evoked potential complex. It is concluded that the early fast waves of the SEP and AEP are modality specific and may represent the parallel activation of primary and secondary sensory cortex through established parallel afferent projections from lateral and medial thalamic nuclei. The later slow waves of the SEP and AEP appear to selectively involve primary and secondary sensory cortex but are more widely distributed, possibly reflecting a less modality-specific level of information processing in dysgranular cortex.     

Signals from the superficial layers of the superior colliculus enable the development of the auditory space map in the deeper layers

We have examined whether the superficial layers of the superior colliculus (SC) provide the source of visual signals that guide the development of the auditory space map in the deeper layers. Anatomical tracing experiments with fluorescent microspheres revealed that a retinotopic map is present in the newborn ferret SC. Aspiration of the caudal region of the superficial layers of the right SC on postnatal day 0 did not cause a reorganization of this projection. Consequently, recordings made when the animals were mature showed that visual units in the remaining superficial layers in rostral SC had receptive fields that spanned a restricted region of anterior space. Auditory units recorded beneath the remaining superficial layers were tuned to corresponding anterior locations. Both the superficial layer visual map and the deeper layer auditory map were normal in the left, unoperated SC. The majority of auditory units recorded throughout the deeper layers ventral to the superficial layer lesion were also tuned to single sound directions. In this region of the SC, however, we observed much greater scatter in the distribution of preferred sound directions and a significant increase in the proportion of units with spatially ambiguous responses. The auditory representation was degraded, although many of these units were also visually responsive. Equivalent lesions of the superficial layers made in adult ferrets did not alter the topographic order in the auditory representation, suggesting that visual activity in these layers may be involved in aligning the different sensory maps in the developing SC.     

Antisaccade performance predicted by neuronal activity in the supplementary eye field

The voluntary control of gaze implies the ability to make saccadic eye movements specified by abstract instructions, as well as the ability to repress unwanted orientating to sudden stimuli. Both of these abilities are challenged in the antisaccade task, because it requires subjects to look at an unmarked location opposite to a flashed stimulus, without glancing at it. Performance on this task depends on the frontal/prefrontal cortex and related structures, but the neuronal operations underlying antisaccades are not understood. It is not known, for example, how excited visual neurons that normally trigger a saccade to a target (a prosaccade) can activate oculomotor neurons directing gaze in the opposite direction. Visual neurons might, perhaps, alter their receptive fields depending on whether they receive a pro- or antisaccade instruction. If the receptive field is not altered, the antisaccade goal must be computed and imposed from the top down to the appropriate oculomotor neurons. Here we show, using recordings from the supplementary eye field (a frontal cortex oculomotor centre) in monkeys, that visual and movement neurons retain the same spatial selectivity across randomly mixed pro- and antisaccade trials. However, these neurons consistently fire more before antisaccades than prosaccades with the same trajectories, suggesting a mechanism through which voluntary antisaccade commands can override reflexive glances.