Biomedical subjects

Charles E Schroeder

Publications and source records attributed to Charles E Schroeder.

At least 19 recordsLinked to original sources

PubMed2007

Neuronal oscillations and multisensory interaction in primary auditory cortex.

Recent anatomical, physiological, and neuroimaging findings indicate multisensory convergence at early, putatively unisensory stages of cortical processing. The objective of this study was to confirm somatosensory-auditory interaction in A1 and to define both its physiological mechanisms and its consequences for auditory information processing. Laminar current source density and multiunit activity sampled during multielectrode penetrations of primary auditory area A1 in awake macaques revealed clear somatosensory-auditory interactions, with a novel mechanism: somatosensory inputs appear to reset the phase of ongoing neuronal oscillations, so that accompanying auditory inputs arrive during an ideal, high-excitability phase, and produce amplified neuronal responses. In contrast, responses to auditory inputs arriving during the opposing low-excitability phase tend to be suppressed. Our findings underscore the instrumental role of neuronal oscillations in cortical operations. The timing and laminar profile of the multisensory interactions in A1 indicate that nonspecific thalamic systems may play a key role in the effect.

Afferent Pathways↗

PubMed2006

Multisensory processing and oscillatory activity: analyzing non-linear electrophysiological measures in humans and simians.

Stimulus-related oscillations are known to be closely linked to integrative processing in the brain. One research domain within which there has been tremendous interest in oscillatory mechanisms is in the integration of inputs across the widely separated sensory systems. Under the standard approach of assessing multisensory interactions in electrophysiological datasets, the event-related response to a multisensory stimulus is directly compared with the sum of the responses to its unisensory constituents when presented alone. When using methods like wavelet transformation or fast Fourier transformation to derive induced oscillatory signals, however, such linear operations are not appropriate. Here we introduce a simple bootstrapping procedure wherein the linear summation of single unisensory trials forms a distribution against which multisensory trials may be statistically compared, an approach that circumvents the issue of non-linearity when combining unisensory oscillatory responses. To test this approach we applied it to datasets from intracranial recordings in non-human primates and human scalp-recorded EEG, both derived from a simple audio-visual integration paradigm. Significant multisensory interactions were revealed in oscillatory activity centered at 15 and 20 Hz (the so-called beta band). Simulations of different levels of background noise further validated the results obtained by this method. By demonstrating super- and sub-additive effects, our analyses showed that this approach is a valuable metric for studying multisensory interactions reflected in induced oscillatory responses.

Adult↗

PubMed2006

Functional anatomy and interaction of fast and slow visual pathways in macaque monkeys.

We measured the timing, areal distribution, and laminar profile of fast, wavelength-insensitive and slower, wavelength-sensitive responses in V1 and extrastriate areas, using laminar current-source density analysis in awake macaque monkeys. There were 3 main findings. 1) We confirmed previously reported significant ventral-dorsal stream latency lags at the level of V4 (V4 mean = 38.7 ms vs. middle temporal mean = 26.9 ms) and inferotemporal cortex (IT mean = 43.4 ms vs. dorsal bank of the superior temporal sulcus mean = 33.9 ms). 2) We found that wavelength-sensitive inputs in areas V1, V4, and IT lagged the wavelength-insensitive responses by significant margins; this lag increased over successive levels of the system. 3) We found that laminar activation profiles in V4 and IT were inconsistent with "feedforward" input through the ascending ventral cortical pathway; the likely alternative input routes include both lateral inputs from the dorsal stream and direct inputs from nonspecific thalamic neurons. These findings support a "Framing" Model of ventral stream visual processing in which rapidly conducted inputs, mediated by one or more accessory pathways, modulate the processing of more slowly conducted feedforward inputs.

Animals↗

PubMed2006

Is neocortex essentially multisensory?

Although sensory perception and neurobiology are traditionally investigated one modality at a time, real world behaviour and perception are driven by the integration of information from multiple sensory sources. Mounting evidence suggests that the neural underpinnings of multisensory integration extend into early sensory processing. This article examines the notion that neocortical operations are essentially multisensory. We first review what is known about multisensory processing in higher-order association cortices and then discuss recent anatomical and physiological findings in presumptive unimodal sensory areas. The pervasiveness of multisensory influences on all levels of cortical processing compels us to reconsider thinking about neural processing in unisensory terms. Indeed, the multisensory nature of most, possibly all, of the neocortex forces us to abandon the notion that the senses ever operate independently during real-world cognition.

Animals↗

PubMed2006

Changing plans: neural correlates of executive control in monkey and human frontal cortex.

Changing plans depends on executive control, the orchestration of behavior based on knowledge of both goal and context. Dorsolateral prefrontal (DLPFC) and anterior cingulate (ACC) cortices are clearly involved in these processes. Intracranial recordings in these regions were obtained from a monkey performing an executive control-challenging task that is widely used in clinic and laboratory to assess the integrity of cognitive function, the AX version of the continuous performance task (AX-CPT), and directly compared to scalp-recorded evoked potentials in humans. In this task the subject presses a button when detecting a frequent cue-target probe sequence in a stream of letters presented on a computer screen, and withholds response following incorrect sequences. Thus correct performance requires correct encoding of cue and probe instruction and inhibitory control. Intracranial recordings showed that DLPFC in the monkey was primarily activated by conditions that required inhibition of imminent action, as had been shown in human event-related potential (ERP) recordings. Different subregions of monkey ACC were activated primarily by either initiating or inhibiting action, whereas human ERP had shown ACC activation in both situations. We suggest that simultaneous activation of both types of subregions in conflict conditions may account the ubiquitous ACC activation observed with fMRI and ERP in those conditions.

Animals↗

PubMed2006

Differentially variable component analysis: Identifying multiple evoked components using trial-to-trial variability.

Electric potentials and magnetic fields generated by ensembles of synchronously active neurons, either spontaneously or in response to external stimuli, provide information essential to understanding the processes underlying cognitive and sensorimotor activity. Interpreting recordings of these potentials and fields is difficult because detectors record signals simultaneously generated by various regions throughout the brain. We introduce a novel approach to this problem, the differentially variable component analysis (dVCA) algorithm, which relies on trial-to-trial variability in response amplitude and latency to identify multiple components. Using simulations we demonstrate the importance of response variability to component identification, the robustness of dVCA to noise, and its ability to characterize single-trial data. We then compare the source-separation capabilities of dVCA with those of principal component analysis and independent component analysis. Finally, we apply dVCA to neural ensemble activity recorded from an awake, behaving macaque-demonstrating that dVCA is an important tool for identifying and characterizing multiple components in the single trial.

Animals↗

PubMed2006

Ipsilateral hand input to area 3b revealed by converging hemodynamic and electrophysiological analyses in macaque monkeys.

Functional magnetic resonance imaging (fMRI) of the hand representation in primary somatosensory cortex (area 3b) of macaque monkeys revealed an ipsilateral hand input undetected by most previous studies. Ipsilateral responses had a hemodynamic signature indistinguishable from that of contralateral hand responses. We explored the neural mechanisms of the fMRI effects using a second derivative analysis of field potentials [current source density (CSD) analysis] combined with action potential profiles, sampled from area 3b using linear array multielectrodes. In contrast to the predominantly excitatory contralateral response, the colocated ipsilateral response appeared dominated by inhibition, suggesting that ipsilateral inputs may have modulatory effects on contralateral input processing. Our findings confirm bimanual convergence at the earliest stage of cortical somatosensory processing in primates. They also illustrate the value of combined CSD and fMRI analyses in monkeys for defining hidden aspects of sensory function and for investigating the neuronal processes generating fMRI signals.

Animals↗

PubMed2005

Timing of pure tone and noise-evoked responses in macaque auditory cortex.

We compared onset latencies for characteristic frequency pure tone and broadband noise responses in AI and posterior belt regions of the auditory cortex in awake macaques. We found that (1) in AI, responses to characteristic frequency tones and broadband noise have similar latencies, (2) in belt regions, characteristic frequency tone and broadband noise latencies differ significantly; broadband noise latencies are shorter, while characteristic frequency tone latencies are longer than corresponding values in AI, (3) for both pure tone and broadband noise responses in AI, latency decreases with increasing characteristic frequency and (4) despite a similar inverse relationship of tone latency and local characteristic frequency in belt areas, broadband noise latencies are uniformly short, and appear unrelated to local characteristic frequency. Dissociation of broadband noise and pure tone latencies may reflect the use of parallel anatomical routes into belt regions.

Acoustic Stimulation↗

PubMed2005

An oscillatory hierarchy controlling neuronal excitability and stimulus processing in the auditory cortex.

EEG oscillations are hypothesized to reflect cyclical variations in the neuronal excitability, with particular frequency bands reflecting differing spatial scales of brain operation. However, despite decades of clinical and scientific investigation, there is no unifying theory of EEG organization, and the role of ongoing activity in sensory processing remains controversial. This study analyzed laminar profiles of synaptic activity [current source density CSD] and multiunit activity (MUA), both spontaneous and stimulus-driven, in primary auditory cortex of awake macaque monkeys. Our results reveal that the EEG is hierarchically organized; delta (1-4 Hz) phase modulates theta (4-10 Hz) amplitude, and theta phase modulates gamma (30-50 Hz) amplitude. This oscillatory hierarchy controls baseline excitability and thus stimulus-related responses in a neuronal ensemble. We propose that the hierarchical organization of ambient oscillatory activity allows auditory cortex to structure its temporal activity pattern so as to optimize the processing of rhythmic inputs.

Acoustic Stimulation↗

PubMed2005

The case for feedforward multisensory convergence during early cortical processing.

The prevailing hierarchical model of sensory processing in the brain holds that different modalities of sensory information emanating from a single object are analyzed extensively during passage through their respective unisensory processing streams before they are combined in higher-order 'multisensory' regions of the cortex. Because of this view, multisensory interactions that have been found at early, putatively 'unisensory' cortical processing stages during hemodynamic imaging studies have been assumed to reflect feedback modulations that occur subsequent to multisensory processing in the higher-order multisensory areas. In this paper, we consider findings that challenge an exclusively feedback interpretation of early multisensory integration effects. First, high-density electrical mapping studies in humans have shown that multisensory convergence and integration effects can occur so early in the time course of sensory processing that purely feedback mediation becomes extremely unlikely. Second, direct neural recordings in monkeys show that, in some cases, convergent inputs at early cortical stages have physiological profiles characteristic of feedforward rather than feedback inputs. Third, damage to higher-order integrative regions in humans often spares the ability to integrate across sensory modalities. Finally, recent anatomic tracer studies have reported direct anatomical connections between primary visual and auditory cortex. These findings make it clear that multisensory convergence at early stages of cortical processing results from feedforward as well as feedback and lateral connections, thus using the full range of anatomical connections available in brain circuitry.

Animals↗

PubMed2005

Can whole brain nerve conduction velocity be derived from surface-recorded visual evoked potentials? A re-examination of Reed, Vernon, and Johnson (2004).

Reed, Vernon, and Johnson [Reed, T. E., Vernon, P. A., & Johnson, A. M. (2004). Sex difference in brain nerve conduction velocity in normal humans. Neuropsychologia, 42, 1709-1714] reported that "nerve conduction velocity" (NCV) of visual transmission from retina to the primary visual area (V1) is significantly faster in males than females. The authors estimated the NCV by dividing head length (nasion-to-inion distance) by the latency of the well-known P100 component of the visual evoked potential (VEP). Here, we critically examine these metrics and we contend that knowledge of the underlying physiology of neural transmission across the initial stages of the visual processing hierarchy dictates that a number of their assumptions cannot be reasonably upheld. Alternative, and we believe, more parsimonious interpretations of the data are also proposed.

Evoked Potentials, Visual↗

PubMed2005

Neurosurgical access to cortical areas in the lateral fissure of primates.

In this report, a method is presented for gaining direct access to cortical areas within the lateral fissure of primates for neuroanatomical tracer injections and electrode array implantation. Compared to areas on the surface of the brain, the anatomical and physiological properties of areas within the fissure are poorly understood. Typically, access to these areas is indirectly achieved by ablating or passing through intervening areas. To enable direct experimental access, a neurosurgical technique was developed in primates whereby the banks of the lateral fissure were retracted with sparing of the vascular network and intervening areas. In some animals, anatomical tracers were directly injected into target fields without contamination of other areas. In others, multichannel electrode arrays were implanted into target areas for chronic recording of neural activity. Since, these techniques could be adapted for exploration of areas within other sulci, the approach represents an important advance in efforts to elucidate the functional organization of the primate cerebral cortex.

Animals↗

PubMed2005

Multisensory contributions to low-level, 'unisensory' processing.

Neurobiologists have traditionally assumed that multisensory integration is a higher order process that occurs after sensory signals have undergone extensive processing through a hierarchy of unisensory subcortical and cortical regions. Recent findings, however, question this assumption. Studies in humans, nonhuman primates and other species demonstrate multisensory convergence in low level cortical structures that were generally believed to be unisensory in function. In addition to enriching current models of multisensory processing and perceptual functions, these new findings require a revision in our thinking about unisensory processing in low level cortical areas.

Animals↗

PubMed2004

Grabbing your ear: rapid auditory-somatosensory multisensory interactions in low-level sensory cortices are not constrained by stimulus alignment.

Multisensory interactions are observed in species from single-cell organisms to humans. Important early work was primarily carried out in the cat superior colliculus and a set of critical parameters for their occurrence were defined. Primary among these were temporal synchrony and spatial alignment of bisensory inputs. Here, we assessed whether spatial alignment was also a critical parameter for the temporally earliest multisensory interactions that are observed in lower-level sensory cortices of the human. While multisensory interactions in humans have been shown behaviorally for spatially disparate stimuli (e.g. the ventriloquist effect), it is not clear if such effects are due to early sensory level integration or later perceptual level processing. In the present study, we used psychophysical and electrophysiological indices to show that auditory-somatosensory interactions in humans occur via the same early sensory mechanism both when stimuli are in and out of spatial register. Subjects more rapidly detected multisensory than unisensory events. At just 50 ms post-stimulus, neural responses to the multisensory 'whole' were greater than the summed responses from the constituent unisensory 'parts'. For all spatial configurations, this effect followed from a modulation of the strength of brain responses, rather than the activation of regions specifically responsive to multisensory pairs. Using the local auto-regressive average source estimation, we localized the initial auditory-somatosensory interactions to auditory association areas contralateral to the side of somatosensory stimulation. Thus, multisensory interactions can occur across wide peripersonal spatial separations remarkably early in sensory processing and in cortical regions traditionally considered unisensory.

Adult↗

PubMed2004

Timing and laminar profile of eye-position effects on auditory responses in primate auditory cortex.

We examined effects of eye position on auditory cortical responses in macaques. Laminar current-source density (CSD) and multiunit activity (MUA) profiles were sampled with linear array multielectrodes. Eye position significantly modulated auditory-evoked CSD amplitude in 24/29 penetrations (83%), across A1 and belt regions; 4/24 cases also showed significant MUA AM. Eye-position effects occurred mainly in the supragranular laminae and lagged the co-located auditory response by, on average, 38 ms. Effects in A1 and belt regions were indistinguishable in amplitude, laminar profile, and latency. The timing and laminar profile of the eye-position effects suggest that they are not combined with auditory signals at a subcortical stage of the lemniscal auditory pathways and simply "fed-forward" into cortex. Rather, these effects may be conveyed to auditory cortex by feedback projections from parietal or frontal cortices, or alternatively, they may be conveyed by nonclassical feedforward projections through auditory koniocellular (calbindin positive) neurons.

Acoustic Stimulation↗

PubMed2004

Neural dynamics and the fundamental mechanisms of event-related brain potentials.

Event-related potentials (ERPs) provide a critical link between the hemodynamic response, as measured by functional magnetic resonance imaging, and the dynamics of the underlying neuronal activity. Single-trial ERP recordings capture the oscillatory activity that are hypothesized to underlie both communication between brain regions and amplified processing of behaviorally relevant stimuli. However, precise interpretations of ERPs are precluded by uncertainty about their neural mechanisms. One influential theory holds that averaged sensory ERPs are generated by partial phase resetting of ongoing electroencephalographic oscillations, while another states that ERPs result from stimulus-evoked neural responses. We formulated critical predictions of each theory and tested these using direct, intracortical analyses of neural activity in monkeys. Our findings support a predominant role for stimulus-evoked activity in sensory ERP generation, and they outline both logic and methodology necessary for differentiating evoked and phase resetting contributions to cognitive and motor ERPs in future studies.

Animals↗

PubMed2003

Estimation of single-trial multicomponent ERPs: differentially variable component analysis (dVCA).

A Bayesian inference framework for estimating the parameters of single-trial, multicomponent, event-related potentials is presented. Single-trial recordings are modeled as the linear combination of ongoing activity and multicomponent waveforms that are relatively phase-locked to certain sensory or motor events. Each component is assumed to have a trial-invariant waveform with trial-dependent amplitude scaling factors and latency shifts. A Maximum a Posteriori solution of this model is implemented via an iterative algorithm from which the component's waveform, single-trial amplitude scaling factors and latency shifts are estimated. Multiple components can be derived from a single-channel recording based on their differential variability, an aspect in contrast with other component analysis techniques (e.g., independent component analysis) where the number of components estimated is equal to or smaller than the number of recording channels. Furthermore, we show that, by subtracting out the estimated single-trial components from each of the single-trial recordings, one can estimate the ongoing activity, thus providing additional information concerning task-related brain dynamics. We test this approach, which we name differentially variable component analysis (dVCA), on simulated data and apply it to an experimental dataset consisting of intracortically recorded local field potentials from monkeys performing a visuomotor pattern discrimination task.

Algorithms↗

PubMed2003

Auditory cortical neurons respond to somatosensory stimulation.

The prevailing hierarchical model of cortical sensory processing holds that early processing is specific to individual modalities and that combination of information from different modalities is deferred until higher-order stages of processing. In this paper, we present physiological evidence of multisensory convergence at an early stage of cortical auditory processing. We used multi-neuron cluster recordings, along with a limited sample of single-unit recordings, to determine whether neurons in the macaque auditory cortex respond to cutaneous stimulation. We found coextensive cutaneous and auditory responses in caudomedial auditory cortex, an area lying adjacent to A1, and at the second stage of the auditory cortical hierarchy. Somatosensory-auditory convergence in auditory cortex may underlie effects observed in human studies. Convergence of inputs from different sensory modalities at very early stages of cortical sensory processing has important implications for both our developing understanding of multisensory processing and established views of unisensory processing.

Acoustic Stimulation↗