Spatiotemporal maturation of the central and lateral N1 components to tones.

This paper examines maturational changes in the spatiotemporal features of central and lateral N1 components of the auditory evoked potentials (AEPs) to tone stimuli presented with a long stimulus onset asyncrony (SOA; 4200 ms) using the scalp current density (SCD) technique. A group of typically developing children ranging from 6 to 12 years of age and a group of adults were studied. Recently studies have begun to explore the topography of these components in children. These studies, however, often used rapidly presented stimuli and did not elicit observable central N1s in the younger children. Our stimuli elicited both central and lateral N1s. Peak latencies of both components decreased with age. Peak amplitude also decreased with age for the lateral N1 but not for the central N1. Consequently, the difference between the lateral N1 and the central N1 amplitudes (or the ratio of lateral N1 amplitude to central N1 amplitude) also decreased with age, dramatically altering the morphology of the elicited AEP waveforms. Topography of the lateral N1 did not change with age. The location of maximal activation for the central N1 appeared to move more medially with age but this 'apparent' movement is probably due to the decreasing impact of the partially overlapping lateral N1 component whose amplitude is significantly smaller in adults than in children.

Visual activation of frontal cortex: segregation from occipital activity.

Studies in primates have found visually responsive neurons that are distributed beyond cortical areas typically described as directly involved in vision. Among these areas are premotor cortex, supplementary motor area, dorsolateral prefrontal cortex and frontal eye fields. Given these findings, visual stimulation would be expected to result in activation of human frontal cortex. However, few human studies have described sensory activations in frontal regions in response to simple visual stimulation. Such studies have classically described event-related potential (ERP) components over occipital regions. The present study sought to further characterize the spatiotemporal dynamics of visually-evoked electrocortical responses elicited by simple visual stimuli using scalp current density measures derived from high-density ERP recordings, with particular emphasis on the distribution of stimulus-related activity over frontal cortex. Hemiretinal stimuli were viewed passively and during a simple ipsi- or contramanual (RT) task. The motor requirement was included to investigate the effects of response preparation on premovement frontal activations. The results indicate early frontocentral activation, particularly over the right hemisphere (peak magnitude 124-148 ms) that is independent of input visual field or motor response requirement, and that is clearly separate in timecourse from the posterior responses elicited by visual input. These findings are in accord with the multiplicity of visual inputs to frontal cortex and are discussed in terms of frontal lobe functions as may be required in these tasks.

Mismatch negativity in children and adults, and effects of an attended task.

Attention has been shown to modulate the amplitude of the mismatch negativity (MMN) elicited by a small deviation in auditory stimuli in adults. The present study examined the effects of attention and deviant size on MMN amplitude in children. Children and adults were presented with sequences of tones containing standards (1000 Hz) and three deviants varying in degree of deviance from the standard (1050, 1200, and 1500 Hz). Tones were presented in three conditions: (1) while participants ignored them; (2) while participants listened to them and responded to all three deviants; and (3) while participants again ignored them. We found that the MMNs elicited by the hard deviant (1050 Hz) were larger when the children were actively discriminating the stimuli than when they were ignoring them. However, the MMNs elicited by the easy and medium deviants (1500 and 1200 Hz, respectively) in the children and by all three deviants in the adults were not affected by attention.

Two cognitive systems simultaneously prepared for opposite events.

Event-related potentials (ERPs) were recorded during a go/no-go reaction time (RT) task in which subjects responded to rare target tones and withheld response to frequent tones. In a predictable condition, a rare visual stimulus signalled the impending occurrence of a rare tone, whereas a frequent visual stimulus signalled that a frequent tone would be presented. The rare visual stimuli elicited P3, associated with violations of expectations, whereas the rare tones, being predictable, did not. The rare tones elicited the mismatch negativity (MMN), a component associated with preattentive deviance detection, despite the fact that they were expected. RT was faster in the predictable condition than in another condition in which the tones were not predictable. The P3 and RT data indicate, respectively, that subjects anticipated and were ready to respond to the target tones. The MMN result indicates that immediately before target tones, the preattentive system underlying the MMN was set for frequent tones, being unaffected by the information available to the higher order system. Thus, the higher order cognitive system associated with P3 and the lower order cognitive system associated with the MMN were prepared simultaneously for opposite events.

An investigation of the auditory streaming effect using event-related brain potentials.

There is uncertainty concerning the extent to which the auditory streaming effect is a function of attentive or preattentive mechanisms. The mismatch negativity (MMN), which indexes preattentive acoustic processing, was used to probe whether the segregation associated with the streaming effect occurs preattentively. In Experiment 1, alternating high and low those were presented at fast and slow paces while subjects ignored the stimuli. At the slow pace, tones were heard as alternating high and low pitches, and no MMN was elicited. At the fast pace a streaming effect was induced and an MMN was observed for the low stream, indicating a preattentive locus for the streaming effect. The high deviant did not elicit an MMN. MMNs were obtained to both the high and low deviants when the interval between the across-stream deviance was lengthened to more than 250 ms in Experiment 2, indicating that the MMN system is susceptible to processing constraints.

Electrophysiological evidence of developmental changes in the duration of auditory sensory memory.

In behavioral studies, children's memory for tonal frequency has been found to persist for less time than adults' (T. A. Keller & N. Cowan, 1994). The present study was done to evaluate the argument that this effect is due to changes in auditory sensory memory and not to attentional mechanisms. This question was investigated using mismatch negativity (MMN), an auditory event-related potential considered to be insensitive to attention. Participants were 6-7-, 8-10-, and 11-12-year-old children and adults. They were presented with trains of stimuli, beginning with either a standard (1000 Hz) or a deviant (1200 Hz) tone with trains separated by either 1 s or 8 s. All 4 groups exhibited MMNs after delays of 1 s, but only the adults and oldest children exhibited MMNs after 8 s, indicating that there are maturational changes in the duration of auditory sensory memory.

Reactivation of a dormant representation of an auditory stimulus feature.

Research with the mismatch negativity component of event-related potentials has uncovered a system that detects change in the acoustic environment on an automatic basis. The system is considered to compare incoming stimuli to representations of the past and to emit an MMN if change is detected. Previous investigations have shown that the relevant memory of the past can become dormant and then be reactivated by a reminder stimulus. It is unclear, however, whether what is reactivated is an holistic representation of stimuli or separate representations of features of stimuli. The present study provides data that supports the latter possibility but leaves open the former one.

Feature conjunctions and auditory sensory memory.

This study sought to obtain additional evidence that transient auditory memory stores information about conjunctions of features on an automatic basis. The mismatch negativity of event-related potentials was employed because its operations are based on information that is stored in transient auditory memory. The mismatch negativity was found to be elicited by a tone that differed from standard tones in a combination of its perceived location and frequency. The result lends further support to the hypothesis that the system upon which the mismatch negativity relies processes stimuli in an holistic manner.

Attention affects the organization of auditory input associated with the mismatch negativity system.

The mismatch negativity (MMN), a component of event-related potentials (ERP), was used to investigate the effect of attention on auditory stream segregation. Subjects were presented with sequences of alternating high and low tones that occurred at a constant rate, which they ignored. When subjects ignored the stimuli, the three-tone standard and deviant sequences contained within the high- and low-pitched tones did not emerge and no MMNs were obtained. Subjects were then instructed to attend to the high-pitched tones of the stimulus sequences and detect the within-stream deviants. When subjects selectively attended the high-pitched tones, MMNs were obtained to the deviants within both the attended and unattended streams. These results indicate that attention can produce segregation such that the sequences of low- and high-pitched tones are available to the automatic deviance detection system that underlies the generation of the MMN. Selective attention can alter the organization of sensory input in the early stages of acoustic processing.

Predictability of stimulus deviance and the mismatch negativity.

The purpose of this study was to test the previous report that generation of the mismatch negativity (MMN) component of event-related brain potentials (ERPs) is indifferent to the predictable occurrence of stimulus deviance. A pattern of standards (S) and deviants (D) were delivered in a predictable fashion (SSSSD) at two different speeds (1.3 s and 100 ms). An MMN was obtained to the D position tone at the slow but not the fast pace. These results demonstrate that, unlike the P3 component, the MMN is sensitive to the predictable occurrence of stimulus deviance when the predictability can be detected by the brain within the estimated limits of sensory memory.

Storage of feature conjunctions in transient auditory memory.

The purpose of this study was to determine whether feature conjunctions are stored in transient auditory memory. The mismatch negativity (MMN), an event-related potential that is elicited by stimuli that differ from a series of preceding stimuli, was used in this endeavour. A tone that differed from the preceding series of stimuli in the conjunction of two of its features, both present in preceding stimuli but in different combinations, was found to elicit the MMN. The data are interpreted to indicate that information about the conjunction of features is stored in the memory.

N-methyl-D-aspartate enhancement of phasic responses in primate neocortex.

In area 17 of the awake macaque, disinhibition by blockade of GABA(A) receptors results in a marked elevation in neuronal excitability, with a particular focus in the supragranular laminae. We examined the possibility that the excitatory supragranular response is N-methyl-D-aspartate (NMDA)-mediated. Laminar activity profiles consisting of flash-evoked field potential, current source density (CSD) and multiunit activity (MUA) measures were obtained during striate cortex penetrations using multicontact electrodes that incorporated single or double microinjection cannulae. Profiles were recorded before and at successive time points after bicuculline induction of disinhibition. Both the noncompetitive NMDA antagonist MK-801 and the competitive antagonist APV reversed bicuculline effects, producing a normal laminar activity profile. NMDA-mediated enhancement of excitatory responses in the supragranular laminae of neocortex is believed to play a role in normal signal processing, as well as in epileptic manifestations.

Lexical processing of visually and auditorily presented nouns and verbs: evidence from reaction time and N400 priming data.

We investigated two aspects of lexical organization in normal adults employing behavioral and electrophysiological indices of semantic priming, namely: (1) Is there evidence for differential processing of nouns and verbs? (2) Is there evidence for separate systems for processing of orthographic and phonologic representations of words? Reaction time (RT), N400 amplitude and latency were used to examine the effect of semantic priming on lexical access of auditorily and visually presented nouns and verbs. We found that the temporal patterns of primed RTs and N400 latencies differed for nouns and verbs, indicating a functional difference in processing. However, the absence of topographic differences in N400 between nouns and verbs did not support anatomically distinct representations of these word classes. By contrast, a modality-specific topography at N400, in addition to RT and N400 amplitude differences between auditory and visual conditions, supported the proposed separation of the orthographic and phonologic representations of words. The implications of the findings for general theories of lexical organization are discussed.

Cognitive and neurologic development of the premature, small for gestational age infant through age 6: comparison by birth weight and gestational age.

OBJECTIVE: To compare the neurologic and cognitive outcomes of 129 premature small for gestational age (SGA) infants with 300 premature appropriate for gestational age (AGA) infants through 6 years of age. DESIGN: Infants born at < or = 37 weeks gestational age and < or = 2500 g with birth weight 2 standard deviations or more below the mean birth weight for gestational age were categorized as SGA. Cognitive and neurologic outcomes of SGA and AGA prematures at 1, 2, 3, and 5 and/or 6 years of age were compared when the infants were stratified by gestational age in 2-week intervals or by birth weight in 500-g intervals. The association between SGA/AGA and neurologic status on cognitive outcomes at each age was also examined. RESULTS: SGA infants had significantly poorer cognitive scores at each age when compared with AGA infants of similar gestational ages. Normal neurologic status was more likely at all assessments for the AGA than for SGA infants of comparable gestational age. There were no differences between SGA and AGA children in cognitive or neurologic outcomes at any age when grouped by birth weight. Cognitive impairment was closely associated with neurologic abnormality in both SGA and AGA groups. There was, nevertheless, a significant effect of SGA on cognitive outcome independent of neurologic status at all ages except 3 years. CONCLUSIONS: Irrespective of degree of prematurity, SGA infants are at greater risk for neurodevelopmental impairment than are equally premature AGA infants. The cognitive impairment can be largely, but not entirely, attributed to a higher incidence of neurologic abnormalities in the SGA infants at each gestational age.

Storage of information in transient auditory memory.

This study concerns the manner in which features of auditory stimuli are stored in acoustic memory. Event-related potentials (ERPs) were recorded to sequences of tones in which sequential, infrequent deviant tones were presented in a row, each of which differed from the frequent standard tones along a different stimulus dimension. The object was to determine whether a change in a single feature of a stimulus would have an effect on the entire representation of the standard tone in memory, or only on the representation of the stimulus dimension by which the first deviant differed from the standards. It was found that the amplitude of the mismatch negativity elicited by subsequent deviants was not reduced by the presence of the first deviant, supporting independent storage of features.

Electrophysiologic manifestations of impaired temporal lobe auditory processing in verbal auditory agnosia.

The present study examined the extent to which verbal auditory agnosia (VAA) is primarily a phonemic decoding disorder, as contrasted to a more global defect in acoustic processing. Subjects were six young adults who presented with VAA in childhood and who, at the time of testing, showed varying degrees of residual auditory discrimination impairment. They were compared to a group of young adults with normal language development matched for age and gender. Cortical event-related potentials (ERPs) were recorded to tones and to consonant-vowel stimuli presented in an "oddball" discrimination paradigm. In addition to cortical ERPs, auditory brainstem responses (ABRs) and middle latency responses (MLRs) were recorded. Cognitive and language assessments were obtained for the VAA subjects. ABRs and MLRs were normal. In comparison with the control group, the cortical ERPs of the VAA subjects showed a delay in the N1 component recorded over lateral temporal cortex both to tones and to speech sounds, despite an N1 of normal latency overlying the frontocentral region of the scalp. These electrophysiologic findings indicate a slowing of processing of both speech and nonspeech auditory stimuli and suggest that the locus of this abnormality is within the secondary auditory cortex in the lateral surface of the temporal lobes.

Relations between infant neurobehavioral performance and cognitive outcome in very low birth weight preterm infants.

The predictive utility of three aspects of neonatal neurobehavioral performance was examined in 144 very low birth weight (< 1500 g) preterms who were followed until 6 years of age. Visual-following and auditory-orienting composites derived from the Einstein Neonatal Neurobehavioral Assessment Scale were modestly related to the Mental Developmental Index (MDI) and IQ scores at several ages, whereas the active motility composite was only related to MDI scores at 1 year of age (corrected). Infants who showed deviant performance on both visual following and auditory orienting composites had significantly lower cognitive test scores at 1 and 6 years of age and were more likely to be classified as subaverage at 6 years of age (IQ < 85). Group differences were independent of both neonatal health status and motor scores and were not due to the performance of children with severe sensory impairments. These findings suggest that visual following and auditory orienting measured in the neonatal period can offer a useful way of indexing initial capacities.

Dynamic neuroimaging of brain function.

To fully characterize the brain processes underlying sensorimotor and cognitive function, the spatial distribution of active regions, their interconnected regions must be measured. We describe methods for imaging brain sources from surface-recorded EEG and magnetoencephalographic data, called electromagnetic source imaging (EMSI). EMSI provides brain source locations within the common framework of magnetic resonance (MR) images of brain anatomy. This allows integration of data from other functional brain imaging methods, like positron emission tomography and functional MR imaging, which can improve the accuracy of EMSI localization. EMSI also provides submillisecond temporal resolution of the dynamic processes within brain systems. Examples are given of applications to visual perceptual and attentional studies.

Physiologic correlates of the voice onset time boundary in primary auditory cortex (A1) of the awake monkey: temporal response patterns.

Behavioral studies in animals support the view that categorical, phonetic phenomena are based upon specific response properties of the auditory system. This study investigated physiologic responses reflecting the phonetic parameter of voice onset time (VOT). We examined multiunit activity (MUA) in the primary auditory cortex (A1) of awake monkeys elicited by the consonant-vowel syllables /da/ and /ta/ that varied in VOT from 0 to 60 msec. Two temporal response patterns encode VOT. The first pattern contains responses time-locked to stimulus onset and to the onset of voicing. In 10 of 17 electrode penetrations that display this pattern, MUA reflects the VOT perceptual boundary by containing a prominent response to voicing onset only for /ta/ stimuli. The second pattern contains responses phase-locked to the periodic portion of the syllables. MUA exhibiting this temporal pattern does not display categorical-like properties. We conclude that specific temporal response patterns in A1 reflect the perceptual boundary for VOT and may represent a physiologic correlate for categorical perception of this phonetic parameter.