Competition between auditory and visual spatial cues during visual task performance

There is debate in the crossmodal cueing literature as to whether capture of visual attention by means of sound is a fully automatic process. Recent studies show that when visual attention is endogenously focused sound still captures attention. The current study investigated whether there is interaction between exogenous auditory and visual capture. Participants preformed an orthogonal cueing task, in which, the visual target was preceded by both a peripheral visual and auditory cue. When both cues were presented at chance level, visual and auditory capture was observed. However, when the validity of the visual cue was increased to 80% only visual capture and no auditory capture was observed. Furthermore, a highly predictive (80% valid) auditory cue was not able to prevent visual capture. These results demonstrate that crossmodal auditory capture does not occur when a competing predictive visual event is presented and is therefore not a fully automatic process.     

Connections between the amygdala and auditory cortical areas in the macaque monkey

Connections between the amygdala and auditory cortical areas TC, and the rostral, intermediate and caudal regions of area TA (TAr, TAi and TAc, respectively) in the macaque monkey (Macaca fuscata and Macaca nemestrina) were investigated following placements of cortical deposits of wheat germ agglutinin-conjugated horseradish peroxidase (WGA-HRP). Areas TC and TAc received weak projections and these derived only from the lateral basal nucleus. Areas TAi and TAr received projections from the lateral, lateral basal and accessory basal nuclei. In contrast, corticopetal projections to the amygdala originated in areas TAi and TAr, but never in TAc or TC. The projections from areas TAi and TAr terminated only in the lateral nucleus, and in particular at the lateral part of the middle and caudal portions of the amygdala. Thus, the amygdalofugal projections to the auditory cortices are more widespread and more complex than the amygdalopetal projections of the auditory cortices. As judged from experiments in which deposits were made at different sites along the rostrocaudal axis of the auditory cortex, there was a progressive increase seen in density of the amygdala connections with more anteriorly-placed injection sites.     

Sound-induced synchronization of neural activity between and within three auditory cortical areas

Neural synchrony within and between auditory cortical fields is evaluated with respect to its potential role in feature binding and in the coding of tone and noise sound pressure level. Simultaneous recordings were made in 24 cats with either two electrodes in primary auditory cortex (AI) and one in anterior auditory field (AAF) or one electrode each in AI, AAF, and secondary auditory cortex. Cross-correlograms (CCHs) for 1-ms binwidth were calculated for tone pips, noise bursts, and silence (i.e., poststimulus) as a function of intensity level. Across stimuli and intensity levels the total percentage of significant stimulus onset CCHs was 62% and that of significant poststimulus CCHs was 58% of 1,868 pairs calculated for each condition. The cross-correlation coefficient to stimulus onsets was higher for single-electrode pairs than for dual-electrode pairs and higher for noise bursts compared with tone pips. The onset correlation for single-electrode pairs was only marginally larger than the poststimulus correlation. For pairs from electrodes across area boundaries, the onset correlations were a factor 3-4 higher than the poststimulus correlations. The within-AI dual-electrode peak correlation was higher than that across areas, especially for spontaneous conditions. Correlation strengths for between area pairs were independent of the difference in characteristic frequency (CF), thereby providing a mechanism of feature binding for broadband sounds. For noise-burst stimulation, the onset correlation for between area pairs was independent of stimulus intensity regardless the difference in CF. In contrast, for tone-pip stimulation a significant dependence on intensity level of the peak correlation strength was found for pairs involving AI and/or AAF with CF difference less than one octave. Across all areas, driven rate, between-area peak correlation strength, or a combination of the two did not predict stimulus intensity. However, between-area peak correlation strength performs better than firing rate to decide if a stimulus is present or absent.     

Orientation sensitivity of cat LGN neurones with and without inputs from visual cortical areas 17 and 18

Summary Orientation sensitivity was tested, using moving bars as stimuli, in 136 LGN cells in normal cats and 82 LGN cells in cats with areas 17 and 18 lesioned.The responses of most neurones showed some dependence on the orientation of the line stimulus. The orientation bias was more pronounced for long, narrow bars moving at rather slow velocities. Length-response curves revealed less end-inhibition along the optimum orientation than along the nonoptimum orientation. Thirty-two percent of the cells in the normal cats and 50% in the lesioned animals responded best to orientations within 10 ° of the vertical or horizontal. The oblique orientations were represented poorly in the lesioned group. Thus the corticogeniculate feedback may serve to confer a more uniform distribution of orientation preferences on the LGN.It is suggested that the orientation biases of LGN neurones may play a role in building orientation-selective cells in the visual cortex. Further, the preferences for horizontal and vertical orientations in the LGN may explain the preferences for these orientations reported for visual cortical cells.     

Unidirectional monosynaptic connections from auditory areas to the primary visual cortex in the marmoset monkey

Brain Structure and Function - Until the late twentieth century, it was believed that different sensory modalities were processed by largely independent pathways in the primate cortex, with...     

Mechanisms of cross-modal priming between visual and haptic modalities

We explored mechanisms of cross-modal priming between visual and haptic modalities. Specifically, we investigated a mechanism of the visual-to-haptic transfer (Experiment 1) and vice versa (Experiment 2). In Experiment 1, three experimental groups, presented visual prime stimuli using novel three-line patterns, were asked to form visual images matching only with shape, haptic images matching only with shape, or haptic images matching with both shape and texture of haptic targets. Priming occurred only when induced haptic images of prime stimuli coincided with the actual texture of haptic targets. In Experiment 2, two experimental groups, presented haptic prime stimuli, were asked to form visual images matching only with shape, or visual images matching with both shape and material (i.e., monochromatic contrast between foreground and background) of visual targets. Priming occurred regardless of experimental conditions, including the control group. Thus, both shape and material representations significantly contributed to the visual-to-haptic transfer. Contrastingly, only shape representation played a significant role in the haptic-to-visual transfer.     

Synaptic properties of thalamic and intracortical inputs to layer 4 of the first- and higher-order cortical areas in the auditory and somatosensory systems

The thalamus is an essential structure in the mammalian forebrain conveying information topographically from the sensory periphery to primary neocortical areas. Beyond this initial processing stage, "higher-order" thalamocortical connections have been presumed to serve only a modulatory role, or are otherwise functionally disregarded. Here we demonstrate that these "higher-order" thalamic nuclei share similar synaptic properties with the "first-order" thalamic nuclei. Using whole cell recordings from layer 4 neurons in thalamocortical slice preparations in the mouse somatosensory and auditory systems, we found that electrical stimulation in all thalamic nuclei elicited large, glutamatergic excitatory postsynaptic potentials (EPSPs) that depress in response to repetitive stimulation and that fail to activate a metabotropic glutamate response. In contrast, the intracortical inputs from layer 6 to layer 4 exhibit facilitating EPSPs. These data suggest that higher-order thalamocortical projections may serve a functional role similar to the first-order nuclei, whereas both are physiologically distinct from the intracortical layer 6 inputs. These results suggest an alternate route for information transfer between cortical areas via a corticothalamocortical pathway.     

Spatial and cross-modal attention alter responses to unattended sensory information in early visual and auditory human cortex

Attending to a visual or auditory stimulus often requires irrelevant information to be filtered out, both within the modality attended and in other modalities. For example, attentively listening to a phone conversation can diminish our ability to detect visual events. We used functional magnetic resonance imaging (fMRI) to examine brain responses to visual and auditory stimuli while subjects attended visual or auditory information. Although early cortical areas are traditionally considered unimodal, we found that brain responses to the same ignored information depended on the modality attended. In early visual area V1, responses to ignored visual stimuli were weaker when attending to another visual stimulus, compared with attending to an auditory stimulus. The opposite was true in more central visual area MT+, where responses to ignored visual stimuli were weaker when attending to an auditory stimulus. Furthermore, fMRI responses to the same ignored visual information depended on the location of the auditory stimulus, with stronger responses when the attended auditory stimulus shared the same side of space as the ignored visual stimulus. In early auditory cortex, responses to ignored auditory stimuli were weaker when attending a visual stimulus. A simple parameterization of our data can describe the effects of redirecting attention across space within the same modality (spatial attention) or across modalities (cross-modal attention), and the influence of spatial attention across modalities (cross-modal spatial attention). Our results suggest that the representation of unattended information depends on whether attention is directed to another stimulus in the same modality or the same region of space.     

Cone inputs in macaque primary visual cortex

To understand the role of primary visual cortex (V1) in color vision, we measured directly the input from the 3 cone types in macaque V1 neurons. Cells were classified as luminance-preferring, color-luminance, or color-preferring from the ratio of the peak amplitudes of spatial frequency responses to red/green equiluminant and to black/white (luminance) grating patterns, respectively. In this study we used L-, M-, and S-cone-isolating gratings to measure spatial frequency response functions for each cone type separately. From peak responses to cone-isolating stimuli we estimated relative cone weights and whether cone inputs were the same or opposite sign. For most V1 cells the relative S-cone weight was <0.1. All color-preferring cells were cone opponent and their L/M cone weight ratio was clustered around a value of -1, which is roughly equal and opposite L and M cone signals. Almost all cells (88%) classified as luminance cells were cone nonopponent, with a broad distribution of cone weights. Most cells (73%) classified as color-luminance cells were cone opponent. This result supports our conclusion that V1 color-luminance cells are double-opponent. Such neurons are more sensitive to color boundaries than to areas of color and thereby could play an important role in color perception. The color-luminance population had a broad distribution of L/M cone weight ratios, implying a broad distribution of preferred colors for the double-opponent cells.     

Functional topography of converging visual and auditory inputs to neurons in the rat superior colliculus

We have used a slice preparation of the infant rat midbrain to examine converging inputs onto neurons in the deeper multisensory layers of the superior colliculus (dSC). Electrical stimulation of the superficial visual layers (sSC) and of the auditory nucleus of the brachium of the inferior colliculus (nBIC) evoked robust monosynaptic responses in dSC cells. Furthermore, the inputs from the sSC were found to be topographically organized as early as the second postnatal week and thus before opening of the eyes and ear canals. This precocious topography was found to be sculpted by GABAA-mediated inhibition of a more widespread set of connections. Tracer injections in the nBIC, both in coronal slices as well as in hemisected brains, confirmed a robust projection originating in the nBIC with distinct terminals in the proximity of the cell bodies of dSC neurons. Combined stimulation of the sSC and nBIC sites revealed that the presumptive visual and auditory inputs are summed linearly. Finally, whereas either input on its own could manifest a significant degree of paired-pulse facilitation, temporally offset stimulation of the two sites revealed no synaptic interactions, indicating again that the two inputs function independently. Taken together, these data provide the first detailed intracellular analysis of convergent sensory inputs onto dSC neurons and form the basis for further exploration of multisensory integration and developmental plasticity.     

Differential dependence of LTD on glutamate receptors in the auditory cortical synapses of cortical and thalamic inputs

Pyramidal neurons in the auditory cortex (AC) receive glutamatergic inputs from the medial geniculate body (MGB inputs) and other pyramidal neurons (pyramidal inputs). We found that the induction of long-term depression (LTD) in supragranular layers was only partially suppressed by 50 microM D-(-)-2-amino-5-phosphonovalerate (APV), an antagonist of N-methyl-D-aspartate (NMDA) receptors (NMDARs), and 500 microM (+)-alpha-methyl-4-carboxyphenylglycine (MCPG), an antagonist of metabotropic glutamate receptors (mGluRs). However, LTD was not observed in the mixture of APV and MCPG. We hypothesized that the mixed dependence of LTD on glutamate receptors could be attributed to the heterogeneity of MGB inputs and pyramidal inputs. To test this hypothesis, the angle of slicing and other recording conditions were adjusted so that postsynaptic potentials were recorded in normal slices, but not in the slices prepared from the rats with MGB lesion. In these experiments, LTD was suppressed by MCPG alone. The conditions were adjusted to minimize the contribution of MGB inputs in field potentials. In these experiments, the induction of LTD was suppressed by APV alone. Interestingly, the induction of LTD was partially suppressed by 20 microM nifedipine, a blocker of L-type Ca(2+) channels, in the slices prepared from the rats with MGB lesions, but not in normal slices. These findings suggest that the induction of LTD requires activation of mGluRs in the synapses of MGB inputs and of NMDARs in the synapses of pyramidal inputs.     

Interactions between P300 and passive probe responses differ in different visual cortical areas

The regulation of firing thresholds of cortical neurons was suggested as one of the mechanisms underlying the generation of the P300 component in the human event-related potential. According to this hypothesis, the detection of an important stimulus produced the widespread inhibition of "irrelevant" networks, interrupting their ongoing activity and facilitating the analysis of selected information. In the present experiment, the responsiveness of visual cortex was evaluated during the P300 potential by using additional, probing stimuli. Large separation of the cortical visual fields permitted separate analysis of the input and more advanced stages of processing. Responses were recorded from Fz, Cz, Pz and Oz scalp sites. P300 waves were evoked by visual, mentally counted stimuli in a standard "odd-ball" procedure. Visual probes were delivered 200, 300, 400, 500, 700 and 1000 ms later. No responses to the probes were required. Significant suppression of responses to the probes delivered less than 400 ms after target stimuli was found in Oz and Pz but not in Cz or Fz. The suppression was not proportional to the voltage levels from which probe responses started. In Fz and Cz, latencies of probe responses were elongated if probes were delivered less than 400 ms after target stimuli. The results suggest that probe responses suppressed by the P300 potential in occipital and parietal cortex may be restored in frontal areas. In these areas the P300 potential could delay probe responses instead of suppressing them.     

Synaptic interactions between thalamic and cortical inputs onto cortical neurons in vivo

To study the interactions between thalamic and cortical inputs onto neocortical neurons, we used paired-pulse stimulation (PPS) of thalamic and cortical inputs as well as PPS of two cortical or two thalamic inputs that converged, at different time intervals, onto intracellularly recorded cortical and thalamocortical neurons in anesthetized cats. PPS of homosynaptic cortico-cortical pathways produced facilitation, depression, or no significant effects in cortical pathways, whereas cortical responses to thalamocortical inputs were mostly facilitated at both short and long intervals. By contrast, heterosynaptic interactions between either cortical and thalamic, or thalamic and cortical, inputs generally produced decreases in the peak amplitudes and depolarization area of evoked excitatory postsynaptic potentials (EPSPs), with maximal effect at approximately 10 ms and lasting from 60 to 100 ms. All neurons tested with thalamic followed by cortical stimuli showed a decrease in the apparent input resistance (R(in)), the time course of which paralleled that of decreased responses, suggesting that shunting is the factor accounting for EPSP's decrease. Only half of neurons tested with cortical followed by thalamic stimuli displayed changes in R(in). Spike shunting in the thalamus may account for those cases in which decreased synaptic responsiveness of cortical neurons was not associated with decreased R(in) because thalamocortical neurons showed decreased firing probability during cortical stimulation. These results suggest a short-lasting but strong shunting between thalamocortical and cortical inputs onto cortical neurons.     

The cortical visual areas of the sheep.

1. A stereotaxic method for the sheep brain is described. 2. At its widest part the primary visual area (Visual I) of each hemisphere extends approximately 20 mm anteroposteriorly and, when unfolded, approximately 35 mm from side to side. It occupies both walls of the lateral sulcus, and extends medially to the medial wall of the hemisphere and to the depth of the ectolateral sulcus laterally. 3. The most lateral part of the primary visual area includes 10-15 degrees of the ipsilateral field; the contralateral field is represented to 135 degrees from the mid line. 4. Visual II also includes a strip of ipsilateral representation on its medial edge and extends to the supra-sylvian sulcus on the lateral surface of the brain. The furthest lateral representation recorded was 130 degrees lateral. 5. Most of both visual areas is concerned with the area centralis and the visual streak. The remainder of the retina has very little cortical representation. 6. Most cells in Visual I are simple with orientational and sometimes directional sensitivity. Some complex and hypercomplex cells have been seen in Visual I, and these predominate in Visual II. Receptive field sizes from 0-25 to 10 degree were found. Within 15 degrees of the vertical meridian, binocular cells are common in both Visual I and II.     

Absence of cross-modal reorganization in the primary auditory cortex of congenitally deaf cats

To investigate possible cross-modal reorganization of the primary auditory cortex (field A1) in congenitally deaf cats, after years of auditory deprivation, multiunit activity and local field potentials were recorded in lightly anesthetized animals and compared with responses obtained in hearing cats. Local field potentials were also used for current source-density analyses. For visual stimulation, phase-reversal gratings of three to five different spatial frequencies and three to five different orientations were presented at the point of central vision. Peripheral visual field was tested using hand-held stimuli (light bar-shaped stimulus of different orientations, moved in different directions and flashed) typically used for neurophysiological characterization of visual fields. From 200 multiunit recordings, no response to visual stimuli could be found in A1 of any of the investigated animals. Using the current source-density analysis of local field potentials, no local generators of field potentials could be found within A1, despite of the presence of small local field potentials. No multiunit responses to somatosensory stimulation (whiskers, face, pinna, head, neck, all paws, back, tail) could be obtained. In conclusion, there were no indications for a cross-modal reorganization (visual, somatosensory) of area A1 in congenitally deaf cats.     

Retinotopy of cortical connections between the striate cortex and extrastriate visual areas in the rat

Previous studies have determined that the striate cortex of the rat is reciprocally connected with multiple extrastriate cortical areas that are retinotopically organized. The objective of this study was to investigate the retinotopy of the striate-extrastriate connections in the rat, by placing triple or double injections of fluorescent tracers (fluorogold, fast blue, rhodamine dextran, or rhodamine-labeled microspheres) in different regions of the striate cortex (Oc1) and mapping the distribution of cells and fibers labeled with the different tracers in the lateral (Oc2L) and medial (Oc2M) extrastriate cortex. The tracer injection sites were visualized in tangential sections of the flattened cortex and correlated with the myelin layout of the striate cortex and with an electrophysiological map from previous studies. The results showed retinotopically organized Oc1 connections with ten different extrastriate cortical areas. The location of these extrastriate areas and the retinotopy of their striate connections remained mostly invariant despite changes of the injection sites in Oc1. Thus, the quadrantic retinotopy was obtained for striate connections to areas posterior, posterolateral, lateromedial, laterointermediate, laterolateral, anterolateral and rostrolateral in Oc2L; and to areas posteromedial, anteromedial, and anterior in Oc2M. The present anatomical map correlates well with electrophysiological maps of the rat extrastriate cortex from previous studies. Furthermore, they provide a definition of the retinotopy of some areas that have not been completely mapped before. These results reaffirm the existence of multiple extrastriate visual areas in the rat.     

Effects of auditory noise on the psychophysical detection of visual signals: cross-modal stochastic resonance

Harper [D.W. Harper, Signal detection analysis of effect of white noise intensity on sensitivity to visual flicker, Percept. Mot. Skills 48 (1979) 791-798] demonstrated that the visual flicker sensitivity was an inverted U-like function of the intensity of different levels of auditory noise from 50 to 90dB (SPL), without concomitant changes in the response bias. The aim of the present study was to extend these observations in the context of the stochastic resonance, a counterintuitive phenomenon in which a particular level of noise enhances the response of a nonlinear system to a weak input signal. We show psychophysical evidence in a yes-no paradigm for the existence of a stochastic resonance-like phenomenon in the auditory-visual interactions. We show that the detection of a weak visual signal was an inverted U-like function of the intensity of different levels of auditory noise. Nevertheless, for a strong visual signal the auditory noise acts in detriment of the ability of visual detection. Our results suggest that auditory noise could be employed in vision rehabilitation interventions in order to improve the detection of weak visual signals.     

Sound localization during homotopic and heterotopic bilateral cooling deactivation of primary and nonprimary auditory cortical areas in the cat

Although the contributions of primary auditory cortex (AI) to sound localization have been extensively studied in a large number of mammals, little is known of the contributions of nonprimary auditory cortex to sound localization. Therefore the purpose of this study was to examine the contributions of both primary and all the recognized regions of acoustically responsive nonprimary auditory cortex to sound localization during both bilateral and unilateral reversible deactivation. The cats learned to make an orienting response (head movement and approach) to a 100-ms broad-band noise stimulus emitted from a central speaker or one of 12 peripheral sites (located in front of the animal, from left 90 degrees to right 90 degrees , at 15 degrees intervals) along the horizontal plane after attending to a central visual stimulus. Twenty-one cats had one or two bilateral pairs of cryoloops chronically implanted over one of ten regions of auditory cortex. We examined AI [which included the dorsal zone (DZ)], the three other tonotopic fields [anterior auditory field (AAF), posterior auditory field (PAF), ventral posterior auditory field (VPAF)], as well as six nontonotopic regions that included second auditory cortex (AII), the anterior ectosylvian sulcus (AES), the insular (IN) region, the temporal (T) region [which included the ventral auditory field (VAF)], the dorsal posterior ectosylvian (dPE) gyrus [which included the intermediate posterior ectosylvian (iPE) gyrus], and the ventral posterior ectosylvian (vPE) gyrus. In accord with earlier studies, unilateral deactivation of AI/DZ caused sound localization deficits in the contralateral field. Bilateral deactivation of AI/DZ resulted in bilateral sound localization deficits throughout the 180 degrees field examined. Of the three other tonotopically organized fields, only deactivation of PAF resulted in sound localization deficits. These deficits were virtually identical to the unilateral and bilateral deactivation results obtained during AI/DZ deactivation. Of the six nontonotopic regions examined, only deactivation of AES resulted in sound localization deficits in the contralateral hemifield during unilateral deactivation. Although bilateral deactivation of AI/DZ, PAF, or AES resulted in profound sound localization deficits throughout the entire field, the cats were generally able to orient toward the hemifield that contained the acoustic stimulus, but not accurately identify the location of the stimulus. Neither unilateral nor bilateral deactivation of areas AAF, VPAF, AII, IN, T, dPE, nor vPE had any effect on the sound localization task. Finally, bilateral heterotopic deactivations of AI/DZ, PAF, or AES yielded deficits that were as profound as bilateral homotopic cooling of any of these sites. The fact that deactivation of any one region (AI/DZ, PAF, or AES) was sufficient to produce a deficit indicated that normal function of all three regions was necessary for normal sound localization. Neither unilateral nor bilateral deactivation of AI/DZ, PAF, or AES affected the accurate localization of a visual target. The results suggest that hemispheric deactivations contribute independently to sound localization deficits.     

Subcortical auditory input to the primary visual cortex in anophthalmic mice

Anatomical and imaging studies show ample evidence for auditory activation of the visual cortex following early onset of blindness in both humans and animal models. Anatomical studies in animal models of early blindness clearly show intermodal pathways through which auditory information can reach the primary visual cortex. There is clear evidence for intermodal corticocortical pathways linking auditory and visual cortex and also novel connections between the inferior colliculus and the visual thalamus. A recent publication [L.K. Laemle, N.L. Strominger, D.O. Carpenter, Cross-modal innervation of primary visual cortex by auditory fibers in congenitally anophthalmic mice, Neurosci. Lett. 396 (2006) 108–112] suggested the presence of a direct reciprocal connection between the inferior colliculus and the primary visual cortex (V1) in congenitally anophthalmic ZRDCT/An mice. This implies that this mutant mouse would be the only known vertebrate having a direct tectal connection with a primary sensory cortex. The presence of this peculiar pathway was reinvestigated in the ZRDCT/An mouse with highly sensitive neuronal tracers. We found the connections normally described in the ZRDCT/An mouse between: (i) the inferior colliculus and the dorsal lateral geniculate nucleus, (ii) V1 and the superior colliculus, (iii) the lateral posterior nucleus and V1 and between (iv) the inferior colliculus and the medial geniculate nucleus. We also show unambiguously that the auditory subcortical structures do not connect the primary visual cortex in the anophthalmic mouse. In particular, we find no evidence of a direct projection from the auditory mesencephalon to the cortex in this animal model of blindness.