Hemispheric asymmetry for auditory processing in the human auditory brain stem, thalamus, and cortex

We report evidence for a context- and not stimulus-dependent functional asymmetry in the left and right human auditory midbrain, thalamus, and cortex in response to monaural sounds. Neural activity elicited by left- and right-ear stimulation was measured simultaneously in the cochlear nuclei, inferior colliculi (ICs), medial geniculate bodies (MGBs), and auditory cortices (ACs) in 2 functional magnetic resonance imaging experiments. In experiment 1, pulsed noise was presented monaurally to either ear, or binaurally, simulating a moving sound source. In experiment 2, only monaural sounds were presented. The results show a modulation of the neural responses to monaural sounds by the presence of binaural sounds at a time scale of tens of seconds: In the absence of binaural stimulation, the left and right ICs, MGBs, and ACs responded stronger to stimulation of the contralateral ear. When blocks of binaural stimuli were interspersed in the sound sequence, the contralateral preference vanished in those structures in the right hemisphere. The resulting hemispheric asymmetry was similar to the asymmetry demonstrated for spatial sound processing. Taken together, the data demonstrate that functional asymmetries in auditory processing are modulated by context. The observed long time constant suggests that this effect results from a "top-down" mechanism.     

Hemispheric specialization in human prefrontal cortex for resolving certain and uncertain inferences

Uncertainty is a fact of life that must be accommodated in real-world decision making. Although it has been suggested that the right prefrontal cortex (PFC) has a special role to play in decision making under uncertainty, there is very little hard data to support this hypothesis. To better understand the roles of left and right PFCs in reasoning and decision making in situations with complete and incomplete information, we administered simple inference problems to 18 patients with lateralized focal lesions to PFC (9 right hemisphere, 9 left hemisphere) and 22 age- and education-matched normal controls. The stimuli were systematically manipulated for completeness of information regarding the status of the conclusion. Our results demonstrated a 2-way interaction such that patients with left PFC lesions were selectively impaired in trials with complete information, whereas patients with right PFC lesions were selectively impaired in trials with incomplete information. These results provide compelling evidence for hemispheric specialization for reasoning in PFC and suggest that the right PFC has a critical role to play in reasoning about incompletely specified situations. We postulate this role involves the maintenance of ambiguous mental representations that temper premature overinterpretation by the left hemisphere.     

Lateralized anterior cingulate function during error processing and conflict monitoring as revealed by high-resolution fMRI

Recent studies have reported that functional subdivisions of anterior cingulate cortex (ACC) may be selectively responsible for conflict and error-related processing. We examined this claim by imaging ACC activation to correct and erroneous response inhibitions in a GoNogo task. After localizing the ACC cluster in individual subjects using functional magnetic resonance imaging (fMRI) at standard resolution (2 x 2 x 4 mm(3)), high-resolution fMRI (1.5 x 1.5 x 1.5 mm(3)) of the ACC was performed in a second session to investigate its precise functional anatomy. At standard resolution, and in agreement with previous studies, ACC was activated for correct and incorrect responses, albeit more so for errors. High-resolution maps of activated ACC clusters revealed localized and reproducible foci in 9 out of 10 volunteers. Multisubject analysis suggested a bilateral distribution of error-related processes in ACC, whereas correct inhibitions only seemed to activate ACC in the right hemisphere. Subsequent region of interest analysis largely confirmed the activation maps. Our results contribute toward a better understanding of the microanatomy of ACC and demonstrate the potential of fMRI for mapping the functional architecture of brain regions involved in cognitive tasks at a previously unaccomplished spatial scale.     

Cortical specialization for processing first- and second-order motion

Distinct mechanisms underlying the visual perception of luminance-(first-order) and contrast-defined (second-order) motion havebeen proposed from electrophysiological, human psychophysicaland neurological studies; however a cortical specializationfor these mechanisms has proven elusive. Here human brain imaging combined with psychophysical methods was used to assess corticalspecializations for processing these two kinds of motion. Acommon stimulus construction was employed, controlling for differencesin spatial and temporal properties, psychophysical performanceand attention. Distinct cortical regions have been found preferentiallyprocessing either first- or second-order motion, both in occipitaland parietal lobes, producing the first physiological evidencein humans to support evidence from psychophysical studies, brainlesion sites and computational models. These results provideevidence for the idea that first-order motion is computed inV1 and second-order motion in later occipital visual areas,and additionally suggest a functional dissociation between thesetwo kinds of motion beyond the occipital lobe.     

Hemispheric asymmetries in cortical thickness

Using magnetic resonance imaging and computational cortical pattern matching methods, we analyzed hemispheric differences in regional gray matter thickness across the lateral and medial cortices in young, healthy adults (n = 60). In addition, we investigated the influence of gender on the degree of thickness asymmetry. Results revealed global and regionally specific differences between the two hemispheres, with generally thicker cortex in the left hemisphere. Regions with significant leftward asymmetry were identified in the precentral gyrus, middle frontal, anterior temporal and superior parietal lobes, while rightward asymmetry was prominent in the inferior posterior temporal lobe and inferior frontal lobe. On the medial surface, significant rightward asymmetries were observed in posterior regions, while significant leftward asymmetries were evident from the vicinity of the paracentral gyrus extending anteriorly. Asymmetry profiles were similar in both sexes, but hemispheric differences appeared slightly pronounced in males compared with females, albeit a few regions also indicated greater asymmetry in females compared with males. Hemispheric differences in the thickness of the cortex might be related to hemisphere-specific functional specializations that are possibly related to behavioral asymmetries.     

Neural substrates of phonemic perception

The temporal lobe in the left hemisphere has long been implicated in the perception of speech sounds. Little is known, however, regarding the specific function of different temporal regions in the analysis of the speech signal. Here we show that an area extending along the left middle and anterior superior temporal sulcus (STS) is more responsive to familiar consonant-vowel syllables during an auditory discrimination task than to comparably complex auditory patterns that cannot be associated with learned phonemic categories. In contrast, areas in the dorsal superior temporal gyrus bilaterally, closer to primary auditory cortex, are activated to the same extent by the phonemic and nonphonemic sounds. Thus, the left middle/anterior STS appears to play a role in phonemic perception. It may represent an intermediate stage of processing in a functional pathway linking areas in the bilateral dorsal superior temporal gyrus, presumably involved in the analysis of physical features of speech and other complex non-speech sounds, to areas in the left anterior STS and middle temporal gyrus that are engaged in higher-level linguistic processes.     

Hemispheric specialization for movement control produces dissociable differences in online corrections after stroke

In this study, we examine whether corrections made during an ongoing movement are differentially affected by left hemisphere damage (LHD) and right hemisphere damage (RHD). Our hypothesis of motor lateralization proposes that control mechanisms specialized to the right hemisphere rely largely on online processes, while the left hemisphere primarily utilizes predictive mechanisms to specify optimal coordination patterns. We therefore predict that RHD, but not LHD, should impair online correction when task goals are unexpectedly changed. Fourteen stroke subjects (7 LHD, 7 RHD) and 14 healthy controls reached to 1 of the 3 targets that unexpectedly "jumped" during movement onset. RHD subjects showed a considerable delay in initiating the corrective response relative to controls and LHD subjects. However, both stroke groups made large final position errors on the target jump trials. Position deficits following LHD were associated with poor intersegmental coordination, while RHD subjects had difficulty terminating their movements appropriately. These findings confirm that RHD, but not LHD, produces a deficit in the timing of online corrections and also indicate that both stroke groups show position deficits that are related to the specialization of their damaged hemisphere. Further research is needed to identify specific neural circuits within each hemisphere critical for these processes.     

Hemispheric Lateralization Moderates the Life Events–Distress Relationship

Past studies show that life events (LE) predict mental distress. This research tested whether hemispheric lateralization (HL) moderated the relationship between LE and mental distress. In studies 1 and 2, different instruments for assessing HL were used (questionnaire and neuropsychological test). In both studies, LE or daily hassles were positively correlated with distress (study 1) and with anxiety and depression (study 2), only in people with right but not left HL, controlling for effects of gender. In study 3, experimentally induced stress led to increased perceived stress, again only in participants with right but not left HL. These results show consistently that left HL may protect against adverse effects of LE, hassles or acute stress on well‐being. We propose possible mechanisms and future research directions. Copyright © 2014 John Wiley & Sons, Ltd.     

Temporal envelope processing in the human left and right auditory cortices

The goal of this study was to determine the temporal response properties of different auditory cortical areas in humans. This is achieved by recording the phase-locked neural activity to white noises modulated sinusoidally in amplitude (AM) at frequencies between 4 and 128 Hz, in the left and right cortices of 20 subjects. Phase-locked neural responses are recorded in four auditory cortical areas with intracerebral electrodes, and modulation transfer functions (MTFs) are computed from these responses. A number of MTFs are bandpass in shape, demonstrating a selective encoding of AM frequencies below 64 Hz in the auditory cortex. This result provides strong physiological support to the idea that the human auditory system decomposes the temporal envelope of sounds (such as speech) into its constituting AM components. Moreover, the results show a predominant response of cortical auditory areas to the lowest AM frequencies (4-16 Hz). This range matches the range of AM frequencies crucial for speech intelligibility, emphasizing therefore the role played by these initial stations of cortical processing in the analysis of speech. Finally, the results show differences in AM sensitivity across cortical areas and hemispheres, and provide a physiological foundation for claims of functional specialization of auditory areas based on previous population measures.     

Working memory specific activity in auditory cortex: potential correlates of sequential processing and maintenance

Working memory (WM) tasks involve several interrelated processes during which past information must be transiently maintained, recalled, and compared with test items according to previously instructed rules. It is not clear whether the rule-specific comparisons of perceptual with memorized items are only performed in previously identified frontal and parietal WM areas or whether these areas orchestrate such comparisons by feedback to sensory cortex. We tested the latter hypothesis by focusing on auditory cortex (AC) areas with low-noise functional magnetic resonance imaging in a 2-back WM task involving frequency-modulated (FM) tones. The control condition was a 0-back task on the same stimuli. Analysis of the group data identified an area on right planum temporale equally activated by both tasks and an area on the left planum temporale specifically involved in the 2-back task. A region of interest analysis in each individual revealed that activation on the left planum temporale in the 2-back task positively correlated with the task performance of the subjects. This strongly suggests a prominent role of the AC in 2-back WM tasks. In conjunction with previous findings on FM processing, the left lateralized effect presumably reflects the complex sequential processing demand of the 2-back matching to sample task.     

Spectral and temporal processing in rat posterior auditory cortex

The rat auditory cortex is divided anatomically into several areas, but little is known about the functional differences in information processing between these areas. To determine the filter properties of rat posterior auditory field (PAF) neurons, we compared neurophysiological responses to simple tones, frequency modulated (FM) sweeps, and amplitude modulated noise and tones with responses of primary auditory cortex (A1) neurons. PAF neurons have excitatory receptive fields that are on average 65% broader than A1 neurons. The broader receptive fields of PAF neurons result in responses to narrow and broadband inputs that are stronger than A1. In contrast to A1, we found little evidence for an orderly topographic gradient in PAF based on frequency. These neurons exhibit latencies that are twice as long as A1. In response to modulated tones and noise, PAF neurons adapt to repeated stimuli at significantly slower rates. Unlike A1, neurons in PAF rarely exhibit facilitation to rapidly repeated sounds. Neurons in PAF do not exhibit strong selectivity for rate or direction of narrowband one octave FM sweeps. These results indicate that PAF, like nonprimary visual fields, processes sensory information on larger spectral and longer temporal scales than primary cortex.     

Cortical processing of complex auditory stimuli during alterations of consciousness with the general anesthetic propofol

BACKGROUND: The extent to which complex auditory stimuli are processed and differentiated during general anesthesia is unknown. The authors used blood oxygenation level-dependent functional magnetic resonance imaging to examine the processing words (10 per period; compared with scrambled words) and nonspeech human vocal sounds (10 per period; compared with environmental sounds) during propofol anesthesia. METHODS: Seven healthy subjects were tested. Propofol was given by a computer-controlled pump to obtain stable plasma concentrations. Data were acquired during awake baseline, sedation (propofol concentration in arterial plasma: 0.64 +/- 0.13 microg/ml; mean +/- SD), general anesthesia (4.62 +/- 0.57 microg/ml), and recovery. Subjects were asked to memorize the words. RESULTS: During all periods including anesthesia, the sounds conditions combined elicited significantly greater activations than silence bilaterally in primary auditory cortices (Heschl gyrus) and adjacent regions within the planum temporale. During sedation and anesthesia, however, the magnitude of the activations was reduced by 40-50% (P < 0.05). Furthermore, anesthesia abolished voice-specific activations seen bilaterally in the superior temporal sulcus during the other periods as well as word-specific activations bilaterally in the Heschl gyrus, planum temporale, and superior temporal gyrus. However, scrambled words paradoxically elicited significantly more activation than normal words bilaterally in planum temporale during anesthesia. Recognition the next day occurred only for words presented during baseline plus recovery and was correlated (P < 0.01) with activity in right and left planum temporale. CONCLUSIONS: The authors conclude that during anesthesia, the primary and association auditory cortices remain responsive to complex auditory stimuli, but in a nonspecific way such that the ability for higher-level analysis is lost.     

Representation of interaural temporal information from left and right auditory space in the human planum temporale and inferior parietal lobe

The localization of low-frequency sounds mainly relies on the processing of microsecond temporal disparities between the ears, since low frequencies produce little or no interaural energy differences. The overall auditory cortical response to low-frequency sounds is largely symmetrical between the two hemispheres, even when the sounds are lateralized. However, the effects of unilateral lesions in the superior temporal cortex suggest that the spatial information mediated by lateralized sounds is distributed asymmetrically across the hemispheres. This paper describes a functional magnetic resonance imaging experiment, which shows that the interaural temporal processing of lateralized sounds produces an enhanced response in the contralateral planum temporale (PT). The response is stronger and extends further into adjacent regions of the inferior parietal lobe (IPL) when the sound is moving than when it is stationary. This suggests that the interaural temporal information mediated by lateralized sounds is projected along a posterior pathway comprising the PT and IPL of the respective contralateral hemisphere. The differential responses to moving sounds further revealed that the left hemisphere responded predominantly to sound movement within the right hemifield, whereas the right hemisphere responded to sound movement in both hemifields. This rightward asymmetry parallels the asymmetry associated with the allocation of visuo-spatial attention and may underlie unilateral auditory neglect phenomena.     

Hemispheric asymmetries for different components of global/local attention occur in distinct temporo-parietal loci

Data from brain-damaged and neurologically intact populations indicate hemispheric asymmetries in the temporo-parietal cortex for discriminating an object's global form (e.g. the overall shape of a bicycle) versus its local parts (e.g. the spokes in a bicycle tire). However, it is not yet clear whether such asymmetries reflect processes that (i) bias attention toward upcoming global versus local stimuli and/or (ii) attend/identify global versus local stimuli after they are presented. To investigate these possibilities, we asked sixteen healthy participants to perform a cued global/local attention task while their brain activity was recorded using event-related functional magnetic resonance imaging (fMRI). The results indicated a novel double dissociation. Hemispheric asymmetries for deploying attention toward expected global versus local object features were specific to the intraparietal sulcus (iPs). However, hemispheric asymmetries for identifying global versus local features after they were presented were specific to the inferior parietal lobe/superior temporal gyrus (IPL/STG). This double dissociation provides the first direct evidence that hemispheric asymmetries associated with different components of global/local attention occur in distinct temporo-parietal loci. Furthermore, it parallels an analogous dissociation reported in a recent fMRI study of spatial orienting, suggesting that global/local attention and spatial attention might rely on similar cognitive/neural mechanisms.     

Brain mechanisms mediating auditory attentional capture in humans

The ability to detect and preferentially process salient auditory stimuli, even when irrelevant to a current task, is often critical for adaptive behavior. This stimulus-driven allocation of processing resources is known as "attentional capture." Here we used functional magnetic resonance imaging in humans to investigate brain activity and behavioral effects related to such auditory attentional capture. Participants searched a sequence of tones for a target tone that was shorter or longer than the nontarget tones. An irrelevant singleton feature in the tone sequence resulted in behavioral interference (attentional capture) and activation of parietal and prefrontal cortices only when the singleton was associated with a nontarget tone (nontarget singleton) and not when associated with a target tone (target singleton). In contrast, the presence (vs. absence) of a singleton feature in the sequence was associated with activation of frontal and temporal loci previously associated with auditory change detection. These results suggest that a ventral network involving superior temporal and inferior frontal cortices responds to acoustic variability, regardless of attentional significance, but a dorsal frontoparietal network responds only when a feature singleton captures attention.     

Selectivity for animal vocalizations in the human auditory cortex

We aimed at testing the cortical representation of complex natural sounds within auditory cortex using human functional magnetic resonance imaging (fMRI). To this end, we employed 2 different paradigms in the same subjects: a block-design experiment was to provide a localization of areas involved in the processing of animal vocalizations, whereas an event-related fMRI adaptation experiment was to characterize the representation of animal vocalizations in the auditory cortex. During the first experiment, we presented subjects with recognizable and degraded animal vocalizations. We observed significantly stronger fMRI responses for animal vocalizations compared with the degraded stimuli along the bilateral superior temporal gyrus (STG). In the second experiment, we employed an event-related fMRI adaptation paradigm in which pairs of auditory stimuli were presented in 4 different conditions: 1) 2 identical animal vocalizations, 2) 2 different animal vocalizations, 3) an animal vocalization and its degraded control, and 4) an animal vocalization and a degraded control of a different sound. We observed significant fMRI adaptation effects within the left STG. Our data thus suggest that complex sounds such as animal vocalizations are represented in putatively nonprimary auditory cortex in the left STG. Their representation is probably based on their spectrotemporal dynamics rather than simple spectral features.     

White matter pathway asymmetry underlies functional lateralization

Structural and functional asymmetry of the human brain has been well documented using techniques such as magnetic resonance imaging (MRI). However, asymmetry of underlying white matter connections is less well understood. We applied an MRI technique known as diffusion tensor tractography to reveal the morphology of the white matter in vivo by mapping directions of maximum water diffusion in brain tissue. White matter pathway asymmetry was investigated in a normalized image data set of 30 right-handed young healthy individuals. We identified, for the first time, a rightwardly asymmetric pathway connecting the posterior temporal lobe to the superior parietal lobule. This pathway may be related to auditory spatial attention and working memory for which there is evidence for a rightward laterality from functional imaging studies. Additional leftward asymmetries connecting the parietal and frontal lobes to the temporal lobe may be more closely related to laterality of language.     

Functional cerebral reorganization for auditory spatial processing and auditory substitution of vision in early blind subjects

Early blind (EB) individuals can recognize bidimensional shapes using a prosthesis substituting vision with audition (PSVA) and activate right dorsal extrastriate visual cortex during the execution of this task. The present study used repetitive transcranial magnetic stimulation (rTMS) to further examine the functional role of this structure in the successful use of the PSVA. Moreover, we investigated which auditory parameter used in the prosthesis (pitch, intensity, or spatial location) might contribute to this occipital activation. Results revealed that rTMS applied to right dorsal extrastriate cortex in EB subjects interferes with both the PSVA use and the auditory spatial location task but not with pitch and intensity discriminations. By contrast, rTMS targeting the same cortical areas in sighted subjects did not affect performance on any auditory tasks. Early visual deprivation thus leads to functional cerebral cross-modal reorganization in the processing of auditory information and auditory-to-visual sensory substitution. The findings also point to the specific involvement of the dorsal visual stream for auditory spatial processing in blind subjects. Moreover, this suggests that sensory substitution prostheses can be developed using these additional neural resources to perform tasks that partially compensate for the loss of vision.     

Cortical responses to auditory stimuli during isoflurane burst suppression anaesthesia

The cortical responses to auditory stimuli were studied in 12 patients during isoflurane anaesthesia producing burst suppression (ETisof (SD) 1.4 (0.2) vol.%). Earphones were used to give 3-s trains of auditory click stimuli (60 clicks, 20 clicks per second, 80 dB, 0.1 ms) at irregular intervals. In 10 patients, the electroencephalography (EEG) showed a burst suppression pattern consisting of high-amplitude activity intermingled with suppressed background activity. In eight patients with burst suppression patterns, there was a strong cortical reactivity to the termination, not to the beginning, of auditory stimuli: 80 (20)% of all stimuli presented during EEG suppression evoked high amplitude cortical response, offset-burst. The latency of these auditory offset evoked bursts was 540 (60) ms. Auditory offset evoked bursts suggest that in spite of cortical suppression during deep anaesthesia the brain retains its ability to respond to changes in the acoustic environment.     

Concurrent tonotopic processing streams in auditory cortex

The basis for multiple representations of equivalent frequency ranges in auditory cortex was studied with physiological and anatomical methods. Our goal was to trace the convergence of thalamic, commissural, and corticocortical information upon two tonotopic fields in the cat, the primary auditory cortex (AI) and the anterior auditory field (AAF). Both fields are among the first cortical levels of processing. After neurophysiological mapping of characteristic frequency, we injected different retrograde tracers at separate, frequency-matched loci in AI and AAF. We found differences in their projections that support the notion of largely segregated parallel processing streams in the auditory thalamus and cerebral cortex. In each field, ipsilateral cortical input amounts to approximately 70% of the number of cells projecting to an isofrequency domain, while commissural and thalamic sources are each approximately 15%. Labeled thalamic and cortical neurons were concentrated in tonotopically predicted regions and in smaller loci far from their spectrally predicted positions. The few double-labeled thalamic neurons (<2%) are consistent with the hypothesis that information to AI and AAF travels along independent processing streams despite widespread regional overlap of thalamic input sources. Double labeling is also sparse in both the corticocortical and commissural systems ( approximately 1%), confirming their independence. The segregation of frequency-specific channels within thalamic and cortical systems is consistent with a model of parallel processing in auditory cortex. The global convergence of cells outside the targeted frequency domain in AI and AAF could contribute to context-dependent processing and to intracortical plasticity and reorganization.