The contribution of human cortical area V3A to the perception of chromatic motion: a transcranial magnetic stimulation study

Area V3A was identified in five human subjects on both a functional and retinotopic basis using functional magnetic resonance imaging techniques. V3A, along with other visual areas responsive to motion, was then targeted for disruption by repetitive transcranial magnetic stimulation (rTMS) whilst the participants performed a delayed speed matching task. The stimuli used for this task included chromatic, isoluminant motion stimuli that activated either the L?M or S?(L+M) cone‐opponent mechanisms, in addition to moving stimuli that contained only luminance contrast (L+M). The speed matching task was performed for chromatic and luminance stimuli that moved at slow (2°/s) or faster (8°/s) speeds. The application of rTMS to area V3A produced a perceived slowing of all chromatic and luminance stimuli at both slow and fast speeds. Similar deficits were found when rTMS was applied to V5/MT+. No deficits in performance were found when areas V3B and V3d were targeted by rTMS. These results provide evidence of a causal link between neural activity in human area V3A and the perception of chromatic isoluminant motion. They establish area V3A, alongside V5/MT+, as a key area in a cortical network that underpins the analysis of not only luminance but also chromatically‐defined motion.     

A neural signature of colour and luminance correspondence in bistable apparent motion

The 'correspondence problem' refers to the ambiguity of apparent motion (AM) paths if several similar objects are displaced across successive displays. We investigated the effect of intrinsic object properties such as colour and luminance on AM paths, and used functional magnetic resonance imaging to localize neural correlates of correspondence matching in visual cortical regions. Human subjects looked at an AM display where two dots in diagonally opposite corners of an implicit rectangle were flashed in alternation with two dots in the other two corners, yielding spontaneous alternations between horizontal and vertical AM. The dots differed in colour or luminance, or were identical. Neural activity was analysed as a function of whether the perceived AM path matched the dots' colour or luminance, and was also compared to activity during bistable AM displays without correspondence cues. When AM paths matched colour and luminance cues, activity in early visual cortex was the same as during perception of uncued displays, whereas it was suppressed when perceived AM paths violated colour or luminance cues. Colour-sensitive extrastriate cortex (V4 complex) transiently activated whenever AM perception switched from a pattern violating colour correspondence to one consistent with colour. We propose that the neural correlate of correspondence in early visual cortex reflects regulatory mechanisms that flexibly gate early visual feature processing in accord with an overriding perceptual decision. Conversely, activation of feature-selective extrastriate regions depends on the type of cue used for correspondence matching and may reflect the salience of percepts that match in colour and motion.     

Brain activation associated with visual motion studied by functional magnetic resonance imaging in humans

Localized brain activation in response to moving visual stimuli was studied by functional magnetic resonance imaging (fMRI). Stimuli were 100 small white dots randomly arranged on a visual display. During the Motion condition, the dots moved along random, noncoherent linear trajectories at different velocities. During the Blink condition, the dots remained stationary but blinked on and off every 500 ms. The Motion and Blink conditions continuously alternated with 10 cycles per run and 6–8 runs per experiment. In half of the runs, the starting stimulus condition was Motion, while in the remaining runs it was Blink. A series of 128 gradient echo echoplanar images were acquired from 5–7 slices during each run using a 1.5 T GE Signa with an Advanced NMR echoplanar subsystem. The time series for each voxel were analyzed in the frequency domain. Voxels which demonstrated a significant spectral peak at the alternation frequency and whose phase changed in response to stimulus order were considered activated. These activated voxels were displayed upon high resolution anatomical images to determine the sites of activation and were also transformed into the coordinates of Talairach and Tournoux ([1988] Co-planar Stereotaxic Atlas of the Human Brain, New York: Thieme) for comparison to prior neuroimaging studies. Seven of ten subjects showed clusters of activation bilaterally at the junction of the temporal and occipital lobes (area 37) in response to moving stimuli. Most activated voxels were located within or adjacent to a region designated the parietal-temporal-occipital fossa, or PTOF. Five subjects also showed activation to moving stimuli in midline occipital cortex. The activated voxels in midline cortex had a significantly shorter phase delay in their MR signal change relative to voxels in PTOF. © 1995 Wiley-Liss, Inc. 1

1 This article is a US Government work and, as such, is in the public domain in the United States of America

     

Amygdala and insular responses to emotionally valenced human faces in small animal specific phobia.

BACKGROUND: Contemporary neurobiological models suggest that the amygdala plays an important role in the pathophysiology of anxiety disorders. However, it is not clear to what extent this concept applies across anxiety disorders. Several studies have examined brain function in specific phobias but did not demonstrate amygdala responses or use specific probes of the amygdala. METHODS: Ten subjects with specific small animal phobia and 10 matched control subjects were studied with functional magnetic resonance imaging. Subjects viewed emotionally expressive and neutral faces, and amygdala blood oxygenation level dependent responses from each group were compared. RESULTS: There was a significant response to the fearful versus neutral faces in the amygdala across both groups but no diagnosis x condition interaction. Post hoc analysis of the whole brain revealed a significantly greater response to the fearful versus neutral faces in the right insular cortex of the specific phobia group than in the control group. CONCLUSIONS: Amygdala hyperresponsivity to emotional faces was not observed in subjects with small animal specific phobia, in contrast to findings in other anxiety disorders (e.g., posttraumatic stress disorder). This suggests a restricted role for the amygdala in specific phobia. The insular hyperresponsivity to fearful versus neutral faces in the subjects with specific phobias warrants further study.     

Retinotopic distribution of chromatic responses in human primary visual cortex

In non-human primates at least three anatomically and functionally distinct channels convey signals from the retina to the primary visual cortex (V1). Two of these channels, the parvocellular and the koniocellular, are sensitive to chromatic contrasts and form the basis of color vision. In humans, common phylogenetic history with other primates and psychophysical experiments suggest identical retinocortical mechanisms but separate evaluation of the distinct anatomical channels has been difficult because signals are already combined in V1. We studied the spatial distribution of activation to chromatic stimuli along the two opponent chromatic axes in human V1 with multifocal functional magnetic resonance imaging. The signal strength was quantified from three experiments with stimuli up to 20 degrees eccentricity. The hypothesis was that, although the parvo- and koniocellular signals are mixed in V1, distinct distributions of signal strength would be evident. We found that whereas different conditions activated the same areas of cortex, indicating that they have identical magnification factors, the responses to red/green stimulation were stronger close to the fovea whereas the blue/yellow responses were much less diminished with increasing eccentricity. Both chromatic axes showed saturating contrast response functions. Our measure directly from human V1 is in line with earlier psychophysical studies suggesting relatively stronger parvocellular channel representation close to the fovea, and more uniform distribution of the koniocellular and achromatic channels. In addition, our study presents a way to rapidly quantify retinotopic signal transmission in distinct retinocortical pathways of individual subjects.     

Competing Theories of Binocular Rivalry: A Possible Resolution

The neural basis of binocular rivalry has beenthe subject of vigorous debate. Do discrepantmonocular patterns rival for awareness becauseof neural competition among patternrepresentations or monocular channels? In thisarticle, I briefly review psychophysical andneurophysiological evidence pertaining to boththeories and discuss important new neuroimagingdata which reveal that rivalry is fullyresolved in monocular visual cortex. These newfindings strongly suggest that interocularcompetition mediates binocular rivalry and thatV1 plays an important role in the selection ofconscious visual information. They furthersuggest that rivalry is not a unitaryphenomenon. Interocular competition may fullyaccount for binocular rivalry whereas aseparate mechanism involving patterncompetition likely accounts for monocular andstimulus rivalry.     

Functional decoupling of BOLD and gamma-band amplitudes in human primary visual cortex

Although functional magnetic resonance imaging is an important tool for measuring brain activity, the hemodynamic blood oxygenation level dependent (BOLD) response is only an indirect measure of neuronal activity. Converging evidence obtained from simultaneous recording of hemodynamic and electrical measures suggest that the best correlate of the BOLD response in primary visual cortex is gamma-band oscillations ( approximately 40 Hz). Here, we examined the coupling between BOLD and gamma-band amplitudes measured with magntoencephalography (MEG) in human primary visual cortex in 10 participants. In Experiment A, participants were exposed to grating stimuli at two contrast levels and two spatial frequencies and in Experiment B square and sine wave stimuli at two spatial frequencies. The amplitudes of both gamma-band oscillations and BOLD showed tuning with stimulus contrast and stimulus type; however, gamma-band oscillations showed a 300% increase across two spatial frequencies, whereas BOLD exhibited no change. This functional decoupling demonstrates that increased amplitude of gamma-band oscillations as measured with MEG is not sufficient to drive the subsequent BOLD response.     

Specificity of regions processing biological motion

Using functional magnetic resonance imaging and point light displays portraying six different human actions, we were able to show that several visual cortical regions, including human MT/V5 complex, posterior inferior temporal gyrus and superior temporal sulcus, are differentially active in the subtraction comparing biological motion to scrambled motion. Comparison of biological motion to three-dimensional rotation (of a human figure), articulated motion and translation suggests that human superior temporal sulcus activity reflects the action portrayed in the biological motion stimuli, whereas posterior inferior temporal gyrus responds to the figure and hMT/V5+ to the complex motion pattern present in biological motion stimuli. These results were confirmed with implied action stimuli.     

Characterizing modulatory interactions between areas V1 and V2 in human cortex: A new treatment of functional MRI data

Reversible cooling experiments in monkey visual cortex have demonstrated that visually driven neuronal activity in V2 depends on feedforward projections from V1, whereas neuronal activity in V1 is modulated by feedback, or reentrant, projections from V2. We present evidence for a homologous asymmetry in reciprocal connections between V1 and V2 in human cortex using physiological measurements obtained with functional MRI. The analysis was based on a nonlinear model of effective connectivity that partitioned the influence that one region exerted over another into an obligatory effect (an effect that depended only on the input) and a modulatory effect (an effect that represented an interaction between input and activity intrinsic to the target region). Using estimates of the modulatory effect we tested two related hypotheses: (1) that V2 would be a major source of modulatory influences on V1; and (2) that the modulatory effects of V2 on V1 would be greater than those of V1 on V2. The first constitutes a hypothesis about the regional or topographic organization of (modulatory) effective connectivity and the second hypothesis directly addresses the functional asymmetry suggested by reversible cooling experiments. The results confirmed that the origins of feedback modulatory effects on V1 were regionally specific and most pronounced in V2. In contrast, feedforward modulatory influences on V1 on V2 were negligible. This apparent asymmetry between feedforward and feedback modulatory interactions was evident in both hemispheres and appears to be a fairly robust feature of nonlinear interactions between striate and extrastriate cortex. © 1995 Wiley-Liss, Inc.     

The postprandial increase in blood triglycerides has no direct effect on the brain BOLD response

A previous study showed that ingestion of a liquid meal high in polyunsaturated lipids decreased the blood-oxygenation-level-dependent (BOLD) response measured by functional magnetic resonance imaging (fMRI) during a finger-tapping motor task, and suggested that this effect was due to a direct effect of blood lipids on the cerebral vasculature. This study compared the time course and magnitude of the BOLD response in fixed anatomic locations before and 3 h after ingestion of high versus low lipid content liquid meals (235 ml Ensure Plus [Abbot Labs] with or without 50 ml added canola oil). Blood triglyceride content peaked 3 h after the high lipid meal and was elevated by 33% compared with the low lipid meal. There was no significant effect of meal composition on the time course or magnitude of the BOLD response in fixed-location clusters of voxels which were activated during either a motor (finger-tapping), a visual (flashing checkerboard), or an integrative/cognitive (number addition) block-design task paradigm. The results indicate that increased blood total triglyceride content after a meal with relatively high polyunsaturated fat does not directly alter the hemodynamic BOLD response to neural activity. However, the postprandial effect on BOLD response of other meals with varying fat types and amounts, as well as other nutrients and phytochemicals, remains to be determined.     

Processing of sound location in human cortex

This functional magnetic resonance imaging study was focused on the neural substrates underlying human auditory space perception. In order to present natural-like sound locations to the subjects, acoustic stimuli convolved with individual head-related transfer functions were used. Activation foci, as revealed by analyses of contrasts and interactions between sound locations, formed a complex network, including anterior and posterior regions of temporal lobe, posterior parietal cortex, dorsolateral prefrontal cortex and inferior frontal cortex. The distinct topography of this network was the result of different patterns of activation and deactivation, depending on sound location, in the respective voxels. These patterns suggested different levels of complexity in processing of auditory spatial information, starting with simple left/right discrimination in the regions surrounding the primary auditory cortex, while the integration of information on hemispace and eccentricity of sound may take place at later stages. Activations were identified as being located in regions assigned to both the dorsal and ventral auditory cortical streams, that are assumed to be preferably concerned with analysis of spatial and non-spatial sound features, respectively. The finding of activations also in the ventral stream could, on the one hand, reflect the well-known functional duality of auditory spectral analysis, that is, the concurrent extraction of information based on location (due to the spectrotemporal distortions caused by head and pinnae) and spectral characteristics of a sound source. On the other hand, this result may suggest the existence of shared neural networks, performing analyses of auditory 'higher-order' cues for both localization and identification of sound sources.     

Functional dissociation of transient and sustained fMRI BOLD components in human auditory cortex revealed with a streaming paradigm based on interaural time differences

A number of physiological studies suggest that feature-selective adaptation is relevant to the pre-processing for auditory streaming, the perceptual separation of overlapping sound sources. Most of these studies are focused on spectral differences between streams, which are considered most important for streaming. However, spatial cues also support streaming, alone or in combination with spectral cues, but physiological studies of spatial cues for streaming remain scarce. Here, we investigate whether the tuning of selective adaptation for interaural time differences (ITD) coincides with the range where streaming perception is observed. FMRI activation that has been shown to adapt depending on the repetition rate was studied with a streaming paradigm where two tones were differently lateralized by ITD. Listeners were presented with five different ΔITD conditions (62.5, 125, 187.5, 343.75, or 687.5 μs) out of an active baseline with no ΔITD during fMRI. The results showed reduced adaptation for conditions with ΔITD ≥ 125 μs, reflected by enhanced sustained BOLD activity. The percentage of streaming perception for these stimuli increased from approximately 20% for ΔITD = 62.5 μs to > 60% for ΔITD = 125 μs. No further sustained BOLD enhancement was observed when the ΔITD was increased beyond ΔITD = 125 μs, whereas the streaming probability continued to increase up to 90% for ΔITD = 687.5 μs. Conversely, the transient BOLD response, at the transition from baseline to ΔITD blocks, increased most prominently as ΔITD was increased from 187.5 to 343.75 μs. These results demonstrate a clear dissociation of transient and sustained components of the BOLD activity in auditory cortex.     

Cortical activity to vibrotactile stimulation: an fMRI study in blind and sighted individuals

Blind individuals show visual cortex activity during Braille reading. We examined whether such cross-modal activations reflect processing somatosensory stimuli independent of language by identifying cortical activity during a one-back vibrotactile matching task. Three groups (sighted, early-onset, and late-onset [>12 years] blind) detected whether paired vibrations (25 and 100 Hz), delivered to the right index finger, differed in frequency. Three successive paired vibrations, followed by a no-stimulation interval, were presented in a long event-related design. A fixed effects average z-score analysis showed increased activity throughout the visuotopic visual cortex, where it was mostly restricted to foveal and parafoveal eccentricities. Early blind showed the most extensive distribution of activity. Late blind exhibited activity mostly in similar regions but with declining response magnitudes with age of blindness onset. Three sighted individuals had suprathreshold activity in V1 but negative responses elsewhere in visual cortex. Mixed effects ANOVA confirmed group distinctions in defined regions (V1, V3, V4v, V7, LOC, and MT). These results suggest cross-modal adaptation to tactile stimulation in visual cortex independent of language processes. All groups showed increased activity in left primary (S1) and bilateral second somatosensory areas, but without response magnitude differences between groups throughout sensorimotor cortex. Early blind showed the greatest spatial extent of S1 activity. Blind participants had more extensive bilateral activity in anterior intraparietal sulcus and supramarginal gyrus. Extensive usage of touch in Braille reading may underlie observed S1 expansions in the reading finger representation. In addition, learned attentiveness to touch may explain similar expansion of parietal tactile attention regions.     

Electrophysiological correlates of the BOLD signal for EEG‐informed fMRI

Electroencephalography (EEG) and functional magnetic resonance imaging (fMRI) are important tools in cognitive and clinical neuroscience. Combined EEG–fMRI has been shown to help to characterise brain networks involved in epileptic activity, as well as in different sensory, motor and cognitive functions. A good understanding of the electrophysiological correlates of the blood oxygen level‐dependent (BOLD) signal is necessary to interpret fMRI maps, particularly when obtained in combination with EEG. We review the current understanding of electrophysiological–haemodynamic correlates, during different types of brain activity. We start by describing the basic mechanisms underlying EEG and BOLD signals and proceed by reviewing EEG‐informed fMRI studies using fMRI to map specific EEG phenomena over the entire brain (EEG–fMRI mapping), or exploring a range of EEG‐derived quantities to determine which best explain colocalised BOLD fluctuations (local EEG–fMRI coupling). While reviewing studies of different forms of brain activity (epileptic and nonepileptic spontaneous activity; cognitive, sensory and motor functions), a significant attention is given to epilepsy because the investigation of its haemodynamic correlates is the most common application of EEG‐informed fMRI. Our review is focused on EEG‐informed fMRI, an asymmetric approach of data integration. We give special attention to the invasiveness of electrophysiological measurements and the simultaneity of multimodal acquisitions because these methodological aspects determine the nature of the conclusions that can be drawn from EEG‐informed fMRI studies. We emphasise the advantages of, and need for, simultaneous intracranial EEG–fMRI studies in humans, which recently became available and hold great potential to improve our understanding of the electrophysiological correlates of BOLD fluctuations. Hum Brain Mapp, 36:391–414, 2015. © 2014 The Authors Human Brain Mapping Published by Wiley Periodicals, Inc.     

Attachment avoidance modulates neural response to masked facial emotion

According to recent models of individual differences in attachment organization, a basic dimension of adult attachment is avoidance. Attachment‐related avoidance corresponds to tendencies to withdraw from close relationships and to an unwillingness to rely on others. In the formation of attachment orientation during infancy facial emotional interaction plays a central role. There exists an inborn very rapid decoding capacity for facial emotional expression. In this study, functional magnetic resonance imaging was used to examine differences in automatic brain reactivity to facial emotions as a function of attachment avoidance in a sample of 51 healthy adults. Pictures of sad and happy faces (which are approach‐related interpersonal signals) were presented masked by neutral faces. The Relationship Scales Questionnaire (RSQ) was used to assess the attachment avoidance. Masked sad faces activated the amygdala, the insula, occipito‐temporal areas, and the somatosensory cortices. Independently from trait anxiety, depressivity, and detection performance, attachment avoidance was found to be inversely related to responses of the primary somatosensory cortex (BA 3) to masked sad faces. A low spontaneous responsivity of the primary somatosensory cortex to negative faces could be a correlate of the habitual unwillingness to deal with partners' distress and needs for proximity. The somatosensory cortices are known to be critically involved in the processes of emotional mimicry and simulation which have the potential to increase social affiliation. Our data are consistent with the idea that people who withdraw from close relationships respond spontaneously to a lesser extent to negative interpersonal emotional signals than securely attached individuals. Hum Brain Mapp, 2009. © 2009 Wiley‐Liss, Inc.     

Age-dependent and -independent changes in attention-deficit/hyperactivity disorder (ADHD) during spatial working memory performance

Objectives: Attention-deficit/hyperactivity disorder (ADHD) has been associated with spatial working memory as well as frontostriatal core deficits. However, it is still unclear how the link between these frontostriatal deficits and working memory function in ADHD differs in children and adults. This study examined spatial working memory in adults and children with ADHD, focussing on identifying regions demonstrating age-invariant or age-dependent abnormalities. Methods: We used functional magnetic resonance imaging to examine a group of 26 children and 35 adults to study load manipulated spatial working memory in patients and controls. Results: In comparison to healthy controls, patients demonstrated reduced positive parietal and frontostriatal load effects, i.e., less increase in brain activity from low to high load, despite similar task performance. In addition, younger patients showed negative load effects, i.e., a decrease in brain activity from low to high load, in medial prefrontal regions. Load effect differences between ADHD and controls that differed between age groups were found predominantly in prefrontal regions. Age-invariant load effect differences occurred predominantly in frontostriatal regions. Conclusions: The age-dependent deviations support the role of prefrontal maturation and compensation in ADHD, while the age-invariant alterations observed in frontostriatal regions provide further evidence that these regions reflect a core pathophysiology in ADHD.     

Multi-modality mapping of motor cortex: comparing echoplanar BOLD fMRI and transcranial magnetic stimulation

Summary Multiple non-invasive methods of imaging brain function are now available for presurgical planning and neurobiological research. As these new methods become available, it is important to understand their relative advantages and liabilities, as well as how the information gained compares across different methods. A current and future trend in neurobiological studies as well as presurgical planning is to combine information from different imaging techniques. Multi-modal integration may perhaps give more powerful information than each modality alone, especially when one of the methods is transcranial magnetic stimulation (TMS), with its ability to non-invasively activate the brain. As an initial venture in cross comparing new imaging methods, we performed the following 2 studies, locating motor cortex with echoplanar BOLD fMRI and TMS. The two methods can be readily integrated, with concurring results, although each have important limitations.     

Within‐limb somatotopic representation of acute muscle pain in the human contralateral dorsal posterior insula

It is well established that the insular cortex processes noxious information. We have previously shown that noxious inputs from the arm and leg are coarsely organized somatotopically within the dorsal posterior insula. The same has been shown for inputs from C tactile afferents, which mediate affective touch, and it has been suggested that the insula may be responsible for the localization of some somatosensory stimuli. Knowing the degree of spatial detail may have significant implications for the potential role of the dorsal posterior insula in the processing of noxious stimuli. Using high‐resolution functional magnetic resonance imaging (fMRI), we compared insula activation patterns in 13 subjects during muscle pain induced by injection of hypertonic saline (5%) into three muscles within the same limb: shoulder (deltoid), forearm (flexor carpi radialis), and hand (first dorsal interosseous). Mapping the maximally activated voxels within the contralateral dorsal posterior insula in each individual subject during each pain stimulus revealed a clear somatotopy of activation within the contralateral dorsal posterior insula. Shoulder pain was represented anterior to forearm pain and medial to hand pain. This fine somatotopic organization may be crucial for pain localization or other aspects of the pain experience that differ depending on stimulation site. Hum Brain Mapp, 2010. © 2010 Wiley‐Liss, Inc.     

Functional connectivity corresponding to the tonotopic differentiation of the human auditory cortex

Recent research has demonstrated that resting‐state functional connectivity (RS‐FC) within the human auditory cortex (HAC) is frequency‐selective, but whether RS‐FC between the HAC and other brain areas is differentiated by frequency remains unclear. Three types of data were collected in this study, including resting‐state functional magnetic resonance imaging (fMRI) data, task‐based fMRI data using six pure tone stimuli (200, 400, 800, 1,600, 3,200, and 6,400 Hz), and structural imaging data. We first used task‐based fMRI to identify frequency‐selective cortical regions in the HAC. Six regions of interest (ROIs) were defined based on the responses of 50 participants to the six pure tone stimuli. Then, these ROIs were used as seeds to determine RS‐FC between the HAC and other brain regions. The results showed that there was RS‐FC between the HAC and brain regions that included the superior temporal gyrus, dorsolateral prefrontal cortex (DL‐PFC), parietal cortex, occipital lobe, and subcortical structures. Importantly, significant differences in FC were observed among most of the brain regions that showed RS‐FC with the HAC. Specifically, there was stronger RS‐FC between (1) low‐frequency (200 and 400 Hz) regions and brain regions including the premotor cortex, somatosensory/‐association cortex, and DL‐PFC; (2) intermediate‐frequency (800 and 1,600 Hz) regions and brain regions including the anterior/posterior superior temporal sulcus, supramarginal gyrus, and inferior frontal cortex; (3) intermediate/low‐frequency regions and vision‐related regions; (4) high‐frequency (3,200 and 6,400 Hz) regions and the anterior cingulate cortex or left DL‐PFC. These findings demonstrate that RS‐FC between the HAC and other brain areas is frequency selective.     

Hemispheric asymmetry for spectral and temporal processing in the human antero-lateral auditory belt cortex

The present study investigates the acoustic basis of the hemispheric asymmetry for the processing of speech and music. Experiments on this question ideally involve stimuli that are perceptually unrelated to speech and music, but contain acoustic characteristics of both. Stimuli in previous studies were derived from speech samples or tonal sequences. Here we introduce a new class of noise-like sound stimuli with no resemblance of speech or music that permit independent parametric variation of spectral and temporal acoustic complexity. Using these stimuli in a functional MRI experiment, we test the hypothesis of a hemispheric asymmetry for the processing of spectral and temporal sound structure by seeking cortical areas in which the blood oxygen level dependent (BOLD) signal covaries with the number of simultaneous spectral components (spectral complexity) or the temporal modulation rate (temporal complexity) of the stimuli. BOLD-responses from the left and right Heschl's gyrus (HG) and part of the right superior temporal gyrus covaried with the spectral parameter, whereas covariation analysis for the temporal parameter highlighted an area on the left superior temporal gyrus. The portion of superior temporal gyrus in which asymmetrical responses are apparent corresponds to the antero-lateral auditory belt cortex, which has been implicated with spectral integration in animal studies. Our results support a similar function of the anterior auditory belt in humans. The findings indicate that asymmetrical processing of complex sounds in the cerebral hemispheres does not depend on semantic, but rather on acoustic stimulus characteristics.