Left Hemisphere Mechanisms Research Articles

Interhemispheric Parietal-Frontal Connectivity Predicts the Ability to Acquire a Nondominant Hand Skill.

Introduction: After chronic impairment of the right dominant hand, some individuals are able to compensate with increased performance with the intact left nondominant hand. This process may depend on the nondominant (right) hemisphere's ability to access dominant (left) hemisphere mechanisms. To predict or modulate patients' ability to compensate with the left hand, we must understand the neural mechanisms and connections that underpin this process. Methods: We studied 17 right-handed healthy adults who underwent resting-state functional connectivity (FC) magnetic resonance imaging scans before 10 days of training on a left-hand precision drawing task. We sought to identify right-hemisphere areas where FC from left-hemisphere seeds (primary motor cortex, intraparietal sulcus [IPS], inferior parietal lobule) would predict left-hand skill learning or magnitude. Results: Left-hand skill learning was predicted by convergent FC from left primary motor cortex and left IPS onto the same small region (0.31 cm3) in the right superior parietal lobule (SPL). Discussion: For patients who must compensate with the left hand, the right SPL may play a key role in integrating left-hemisphere mechanisms that typically control the right hand. Our study provides the first model of how interhemispheric functional connections in the human brain may support compensation after chronic injury to the right hand.

Left hemisphere damage produces deficits in predictive control of bilateral coordination.

Previous research has demonstrated hemisphere-specific motor deficits in ipsilesional and contralesional unimanual movements in patients with hemiparetic stroke due to MCA infarct. Due to the importance of bilateral motor actions on activities of daily living, we now examine how bilateral coordination may be differentially affected by right or left hemisphere stroke. To avoid the caveat of simply adding unimanual deficits in assessing bimanual coordination, we designed a unique task that requires spatiotemporal coordination features that do not exist in unimanual movements. Participants with unilateral left (LHD) or right hemisphere damage (RHD) and age-matched controls moved a virtual rectangle (bar) from a midline start position to a midline target. Movement along the long axis of the bar was redundant to the task, such that the bar remained in the center of and parallel to an imaginary line connecting each hand. Thus, to maintain midline position of the bar, movements of one hand closer to or further away from the bar midline required simultaneous, but oppositely directed displacements with the other hand. Our findings indicate that left (LHD), but not right (RHD) hemisphere-damaged patients showed poor interlimb coordination, reflected by significantly lower correlations between displacements of each hand along the bar axis. These left hemisphere-specific deficits were only apparent prior to peak velocity, likely reflecting predictive control of interlimb coordination. In contrast, the RHD group bilateral coordination was not significantly different than that of the control group. We conclude that predictive mechanisms that govern bilateral coordination are dependent on left hemisphere mechanisms. These findings indicate that assessment and training in cooperative bimanual tasks should be considered as part of an intervention framework for post-stroke physical rehabilitation.

What's “left”? Hemispheric sensitivity to predictability and congruity during sentence reading by older adults

A number of studies have found that older adults' sentence processing tends not to be characterized by the prediction-related effects attested for young adults. Here, we further probed older adults’ sensitivity to predictability and congruity by recording event-related brain potentials (ERPs) as adults over age 60 read pairs of sentences, which ended with either the expected word, an unexpected word from the same semantic category, or an unexpected word from a different category. Half of the contexts were highly constraining. Consistent with patterns attested when older adults listened to these same materials (Federmeier et al., 2002), N400s, on average, were smaller to expected than to unexpected words, but did not show constraint-related reductions for unexpected words that shared features with the most predictable completion (an effect well-attested in young adults). This pattern resembles that seen in young adults for right-hemisphere-biased processing. To assess whether older adults retain young-like hemispheric asymmetries but recruit right hemisphere mechanisms more, we examined responses to the target words using visual half-field presentation. Whereas young adults show an asymmetric pattern, with prediction-related N400 amplitude reductions for left- but not right-hemisphere-initiated processing (Federmeier and Kutas, 1999b), older adults showed no reliable processing asymmetries and no evidence for prediction with left hemisphere-initiated presentation. The results suggest that left hemisphere mechanisms important for prediction during language processing are less efficacious in older adulthood.

Brain activity during spoken word recognition in subacute aphasia

• In subacute aphasia, left hemisphere mechanisms support spoken word recognition. • Left aMTG and IFG are recruited differently in subacute aphasia compared to controls. • An auditory fMRI task can be used reliably by people with subacute aphasia.

Lateralized motor control processes determine asymmetry of interlimb transfer

This experiment tested the hypothesis that interlimb transfer of motor performance depends on recruitment of motor control processes that are specialized to the hemisphere contralateral to the arm that is initially trained. Right-handed participants performed a single-joint task, in which reaches were targeted to 4 different distances. While the speed and accuracy was similar for both hands, the underlying control mechanisms used to vary movement speed with distance were systematically different between the arms: the amplitude of the initial acceleration profiles scaled greater with movement speed for the right-dominant arm, while the duration of the initial acceleration profile scaled greater with movement speed for the left-non-dominant arm. These two processes were previously shown to be differentially disrupted by left and right hemisphere damage, respectively. We now hypothesize that task practice with the right arm might reinforce left-hemisphere mechanisms that vary acceleration amplitude with distance, while practice with the left arm might reinforce right-hemisphere mechanisms that vary acceleration duration with distance. We thus predict that following right arm practice, the left arm should show increased contributions of acceleration amplitude to peak velocities, and following left arm practice, the right arm should show increased contributions of acceleration duration to peak velocities. Our findings support these predictions, indicating that asymmetry in interlimb transfer of motor performance, at least in the task used here, depends on recruitment of lateralized motor control processes.

Increased functional connectivity between cortical hand areas and praxis network associated with training-related improvements in non-dominant hand precision drawing

Chronic forced use of the non-dominant left hand yields substantial improvements in the precision and quality of writing and drawing. These changes may arise from increased access by the non-dominant (right) hemisphere to dominant (left) hemisphere mechanisms specialized for end-point precision control. To evaluate this prediction, 22 healthy right-handed adults underwent resting state functional connectivity (FC) MRI scans before and after 10 days of training on a left hand precision drawing task. 89% of participants significantly improved left hand speed, accuracy, and smoothness. Smoothness gains were specific to the trained left hand and persistent: 6 months after training, 71% of participants exhibited above-baseline movement smoothness. Contrary to expectations, we found no evidence of increased FC between right and left hemisphere hand areas. Instead, training-related improvements in left hand movement smoothness were associated with increased FC between both sensorimotor hand areas and a left-lateralized parieto-prefrontal network implicated in manual praxis. By contrast, skill retention at 6 months was predicted by changes including decreased FC between the representation of the trained left hand and bilateral sensorimotor, parietal, and premotor cortices, possibly reflecting consolidation and a disengagement of early learning processes. These data indicate that modest amounts of training (<200min total) can induce substantial, persistent improvements the precision and quality of non-dominant hand control in healthy adults, supported by strengthened connectivity between bilateral sensorimotor hand areas and a left-lateralized parieto-prefrontal praxis network.

Left-hemisphere activation is associated with enhanced vocal pitch error detection in musicians with absolute pitch

The ability to process auditory feedback for vocal pitch control is crucial during speaking and singing. Previous studies have suggested that musicians with absolute pitch (AP) develop specialized left-hemisphere mechanisms for pitch processing. The present study adopted an auditory feedback pitch perturbation paradigm combined with ERP recordings to test the hypothesis whether the neural mechanisms of the left-hemisphere enhance vocal pitch error detection and control in AP musicians compared with relative pitch (RP) musicians and non-musicians (NM). Results showed a stronger N1 response to pitch-shifted voice feedback in the right-hemisphere for both AP and RP musicians compared with the NM group. However, the left-hemisphere P2 component activation was greater in AP and RP musicians compared with NMs and also for the AP compared with RP musicians. The NM group was slower in generating compensatory vocal reactions to feedback pitch perturbation compared with musicians, and they failed to re-adjust their vocal pitch after the feedback perturbation was removed. These findings suggest that in the earlier stages of cortical neural processing, the right hemisphere is more active in musicians for detecting pitch changes in voice feedback. In the later stages, the left-hemisphere is more active during the processing of auditory feedback for vocal motor control and seems to involve specialized mechanisms that facilitate pitch processing in the AP compared with RP musicians. These findings indicate that the left hemisphere mechanisms of AP ability are associated with improved auditory feedback pitch processing during vocal pitch control in tasks such as speaking or singing.

Noninvasive brain stimulation in the treatment of aphasia: Exploring interhemispheric relationships and their implications for neurorehabilitation

Aphasia is a common consequence of unilateral stroke, typically involving perisylvian regions of the left hemisphere. The course of recovery from aphasia after stroke is variable, and relies on the emergence of neuroplastic changes in language networks. Recent evidence suggests that rehabilitation interventions may facilitate these changes. Functional reorganization of language networks following left-hemisphere stroke and aphasia has been proposed to involve multiple mechanisms, including intrahemispheric recruitment of perilesional left-hemisphere regions and transcallosal interhemispheric interactions between lesioned left-hemisphere language areas and homologous regions in the right hemisphere. Moreover, it is debated whether interhemispheric interactions are beneficial or deleterious to recovering language networks. Transcranial magnetic stimulation (TMS) and transcranial direct current stimulation (tDCS) are two safe and noninvasive procedures that can be applied clinically to modulate cortical excitability during poststroke language recovery. Intervention with these noninvasive brain stimulation techniques also allows for inferences to be made regarding mechanisms of recovery, including the role of intrahemispheric and interhemispheric interactions. Here we review recent evidence that suggests that TMS and tDCS are promising tools for facilitating language recovery in aphasic patients, and examine evidence that indicates that both right and left hemisphere mechanisms of plasticity are instrumental in aphasia recovery.

Functional MRI of Language in Aphasia: A Review of the Literature and the Methodological Challenges

Animal analogue studies show that damaged adult brains reorganize to accommodate compromised functions. In the human arena, functional magnetic resonance imaging (fMRI) and other functional neuroimaging techniques have been used to study reorganization of language substrates in aphasia. The resulting controversy regarding whether the right or the left hemisphere supports language recovery and treatment progress must be reframed. A more appropriate question is when left-hemisphere mechanisms and when right-hemisphere mechanisms support recovery of language functions. Small lesions generally lead to good recoveries supported by left-hemisphere mechanisms. However, when too much language eloquent cortex is damaged, right-hemisphere structures may provide the better substrate for recovery of language. Some studies suggest that recovery is particularly supported by homologues of damaged left-hemisphere structures. Evidence also suggests that under some circumstances, activity in both the left and right hemispheres can interfere with recovery of function. Further research will be needed to address these issues. However, daunting methodological problems must be managed to maximize the yield of future fMRI research in aphasia, especially in the area of language production. In this review, we cover six challenges for imaging language functions in aphasia with fMRI, with an emphasis on language production: (1) selection of a baseline task, (2) structure of language production trials, (3) mitigation of motion-related artifacts, (4) the use of stimulus onset versus response onset in fMRI analyses, (5) use of trials with correct responses and errors in analyses, and (6) reliability and stability of fMRI images across sessions. However, this list of methodological challenges is not exhaustive. Once methodology is advanced, knowledge from conceptually driven fMRI studies can be used to develop theoretically driven, mechanism-based treatments that will result in more effective therapy and to identify the best patient candidates for specific treatments. While the promise of fMRI in the study of aphasia is great, there is much work to be done before this technique will be a useful clinical tool.

Part 2: Abstracts of posters

Previous research has shown that individuals with traumatic brain injury (TBI) are significantly impaired on emotion recognition tasks. In this study, the relationship between neuropsychological functioning and emotion recognition was examined in 11 individuals with moderate to severe TBI and a control group of 13 individuals matched for age,sex, and education. Emotion recognition stimuli were from Ekman and Friesen's pictures of facial affect. The group with TBI showed neuropsychological deficits consistent with those commonly found following moderate to severe TBI. The group with TBI also identified significantly fewer emotion recognition stimuli than the control group. The number of correctly identified emotion recognition stimuli was significantly correlated with measures of verbal cognitive processing in the group with TBI. These findings suggest that the role of left hemisphere brain mechanisms in the recognition of facial (nonverbal)emotion may be more important than previously recognized.

Perceptional and executive deficits of chronic schizophrenic patients in attentional and intentional tasks

The present study investigated whether schizophrenic patients could develop appropriate visual orientation and motor set under precuing conditions which contrasted attentional (input selective) and intentional (output selective) information. The aim was to evaluate perceptual performance in processing visuospatial information, and executive performance in response preparation. Stimuli and/or elicited responses were controlled for selective hemispheric engagement. Age, sex and handedness matched groups of 33 chronic schizophrenic patients and 33 normal subjects were tested on choice reaction time (RT) tasks in which warning signals were manipulated regarding either where a target stimulus would occur (selective attention) or which hand to use for responding (response preparation). All subjects benefited from precued information regarding subsequent responses. However, schizophrenic patients were not able to use intentional cues as effectively as control subjects did. Interhemispheric asymmetry of spatial attention was found in patients with schizophrenia, with slowing of responses to uncued targets presented in the right visual field. There was also a decreased advantage of within-hemisphere stimulus-response conditions in the schizophrenic group. Our results support the notion that a dysfunction involving parietal and premotor areas has potential importance in the schizophrenic illness. We replicated findings which indicate that deficits of information processing in schizophrenia may affect left hemispheric mechanisms to a larger extent. The results also point toward a possible abnormal connectivity between frontal and parietal circuits in schizophrenia.

Right hemisphere reading mechanisms in a global alexic patient

We investigated the implicit, or covert, reading ability of a global alexic patient (EA) to help determine the contribution of the right hemisphere to reading. Previous studies of alexic patients with left hemisphere damage have suggested that the ability to derive meaning from printed words that cannot be read out loud may reflect right hemisphere reading mechanisms. Other investigators have argued that residual left hemisphere abilities are sufficient to account for implicit reading and moreover do not require the postulation of a right hemisphere system that has no role in normal reading processes. However, few studies have assessed covert reading in patients with lesions as extensive as the one in EA, which affected left medial, inferior temporal–occipital cortex, hippocampus, splenium, and dorsal white matter. EA was presented with lexical decision, semantic categorization, phonemic categorization, and letter matching tasks. Although EA was unable to access phonology and could not overtly name words or letters, she was nevertheless capable of making lexical and semantic decisions at above chance levels, with an advantage for concrete versus abstract words. Her oral and written spelling were relatively intact, suggesting that orthographic knowledge is retained, although inaccessible through the visual modality. Based on her ability to access lexical and semantic information without contacting phonological representations, we propose that EA’s implicit reading emerges from, and is supported, by the right hemisphere. Finally, we conclude that her spelling and writing abilities are supported by left hemisphere mechanisms.

Prosody in apraxia of speech.

Prosody is a complex process that involves modulation of pitch, loudness, duration, and linearity in the acoustic stream to serve linguistic and affective communication goals. It arises from the interaction of distributed neural networks that may be anatomically and functionally lateralized. Intrinsic prosody is mediated largely through left hemisphere mechanisms and encompasses those elements of linguistic microstructure (e.g., syllabic magnitudes and durations, basic consonantal and vocalic gesture specifications, and so) that yield the segmental aspects of speech. Extrinsic prosody is processed primarily by right hemisphere (RH) mechanisms and involves manipulation of intonation across longer perceptual groupings. Intrinsic prosody deficits can lead to several core symptoms of speech apraxia such as difficulty with utterance initiation and syllable transitionalization and may lead to the establishment of inappropriate syllable boundaries. The intrinsic prosody profiles associated with acquired apraxia of speech, developmental speech apraxia, and ataxic dysarthria may aid in the clinical differentiation of these disorders.

Role of the Frontal Cortical Areas in the Analysis of Visual Stimuli at Conscious and Unconscious Levels

Event-related potentials (ERP) in response to complex target stimuli, which consisted of a central recognizable picture and a lateral masked image (analyzed at the unconscious level) were recorded in adult subjects and seven-year-old children. ERP components N200, N300, and P400/N400 had different topography and were differently pronounced in adults and children. In adult subjects, the N200 component that reflects the processing of a sensory stimulus was recorded in the temporo-parieto-occipital and occipital areas. In children, N200 was recorded in the caudal regions and the frontal areas of the cortex. Analysis of different waveforms obtained by subtraction of the ERP to the central stimulus from the ERP to the complex stimulus showed that unconscious stimulus processing in adult subjects is not reflected in the ERP structure. In children, an unconsciously processed image incorporated into a complex stimulus evokes processing negativity in the occipital and frontal cortical areas. Comparison of ERP in groups of children divided by their reflectivity/impulsivity showed that, predominantly, the left frontal area is involved in image analysis at the unconscious level in reflective children and, predominantly, the right frontal area participates in unconscious image analysis in impulsive children. It is suggested that the perfection of the visual recognition of a target stimulus, which contains additional unconsciously processed information, consists in growth of the involvement of the left-hemispheric mechanisms (with respective growth of significance of the left-hemispheric mechanisms) and in a decrease in the role of the frontal areas in analysis of sensory information.

Pure word deafness and the bilateral processing of the speech code

The analysis of pure word deafness (PWD) suggests that speech perception, construed as the integration of acoustic information to yield representations that enter into the linguistic computational system, (i) is separable in a modular sense from other aspects of auditory cognition and (ii) is mediated by the posterior superior temporal cortex in both hemispheres. PWD data are consistent with neuropsychological and neuroimaging evidence in a manner that suggests that the speech code is analyzed bilaterally. The typical lateralization associated with language processing is a property of the computational system that acts beyond the analysis of the input signal. The hypothesis of the bilateral mediation of the speech code does not imply that both sides execute the same computation. It is proposed that the speech signal is asymmetrically analyzed in the time domain, with left-hemisphere mechanisms preferentially extracting information over shorter (25–50 ms) temporal integration windows and right mechanisms over longer (150–250 ms) windows.

Functional Heterogeneity of Inferior Frontal Gyrus Is Shaped by Linguistic Experience

A crosslinguistic, positron emission tomography (PET) study was conducted to determine the influence of linguistic experience on the perception of segmental (consonants and vowels) and suprasegmental (tones) information. Chinese and English subjects (10 per group) were presented binaurally with lists consisting of five Chinese monosyllabic morphemes (speech) or low-pass-filtered versions of the same stimuli (nonspeech). The first and last items were targeted for comparison; the time interval between target tones was filled with irrelevant distractor tones. A speeded-response, selective attention paradigm required subjects to make discrimination judgments of the target items while ignoring intervening distractor tones. PET scans were acquired for five tasks presented twice: one passive listening to pitch (nonspeech) and four active (speech = consonant, vowel, and tone; nonspeech = pitch). Significant regional changes in blood flow were identified from comparisons of group-averaged images of active tasks relative to passive listening. Chinese subjects show increased activity in left premotor cortex, pars opercularis, and pars triangularis across the four tasks. English subjects, on the other hand, show increased activity in left inferior frontal gyrus regions only in the vowel task and in right inferior frontal gyrus regions in the pitch task. Findings suggest that functional circuits engaged in speech perception depend on linguistic experience. All linguistic information signaled by prosodic cues engages left-hemisphere mechanisms. Storage and executive processes of working memory that are implicated in phonological processing are mediated in discrete regions of the left frontal lobe.

Cortical activation patterns of affective speech processing depend on concurrent demands on the subvocal rehearsal system A DC-potential study

In order to delineate brain regions specifically involved in the processing of affective components of spoken language (affective or emotive prosody), we conducted two event-related potential experiments. Cortical activation patterns were assessed by recordings of direct current components of the EEG signal from the scalp. Right-handed subjects discriminated pairs of declarative sentences with either happy, sad or neutral intonation. Each stimulus pair was derived from two identical original utterances that, due to digital signal manipulations, slightly differed in fundamental frequency (F0) range or in duration of stressed syllables. In the first experiment, subjects were asked: (i) to denote the original emotional category of each sentence pair and (ii) to decide which of the two items displayed stronger emotional expressiveness. Participants in the second experiment were asked to repeat the utterances using inner speech during stimulus presentation in addition to the discrimination task. In the absence of inner speech, a predominant activation of right frontal regions was observed, irrespective of emotional category. In the second experiment, a bilateral activation with left frontal preponderance emerged from discrimination during additional performance of inner speech. Compared with the first experiment, a new pattern of acoustic signal processing arose. A relative decrease of brain activity during processing of F0 stimulus variants was observed together with increased activation during discrimination of duration-manipulated sentence pairs. Analysis of behavioural data revealed no significant differences in evaluation of expressiveness between the two experiments. We conclude that the topographical shift of cortical activity originates from left hemisphere (LH) mechanisms of speech processing that centre around the subvocal rehearsal system as an articulatory control component of the phonological loop. A strong coupling of acoustic input and (planned) verbal output channel in the LH is initiated by subvocal articulatory activity like inner speech. These neural networks may provide interpretations of verbal acoustic signals in terms of motor programs and facilitate continuous control of speech output by comparing the signal produced with that intended. Most likely, information on motor aspects of suprasegmental signal characteristics contributes to the evaluation of affective components of spoken language. In consequence, the right hemisphere (RH) holds a merely relative dominance, both for processing of F0 and for evaluation of emotional significance of sensory input. Psychophysically, an important determinant on expression of lateralization patterns seems to be given by the degree of communicative demands such as solely perceptive (RH) or perceptive and verbal-expressive (RH and LH).

189 EEG Coherence as a measure of functional connectivity: From experiment to human study

The EEG inter- and intrahemispheric coherences (ICoh and HCoh) in the θ, α and β bands were studied in an acallosal group (ACCG) of five children and a normal group of 30 sex- and age-matched children (NG) during resting and tapping conditions. Being functionally deficient, tapping in the ACCG was characterized by increased intertap intervals and variability (in right-hand tapping) and by variability together with decreased synchronization (in bimanual tapping). In the ACCG, frontal, central and parietal ICohs were shown to be smaller, while temporal ICohs were larger under all conditions (see also Koeda, T., Knyazeva, M., Jonkman, J., Njiokiktjien, C., De Sonneville, L., Vildavsky, V., 1995. The resting EEG in acallosal children: compensatory left hemisphere mechanisms? Electroencephalogr. Clin. Neurophysiol. 95, 397–407). The effect was most pronounced in the EEG β band. The sagittal HCohs, including fronto-central, fronto-parietal, and centro-parietal HCohs within both hemispheres, were larger in the ACCG, whereas temporal HCoh (fronto-temporal, centro-temporal, parieto-temporal and occipito-temporal) were smaller, suggesting rearrangement of intracortical activity associated with callosal agenesis. Tapping induced an increase in ICoh and HCoh between frontal, central and parietal areas in the NG, and weak enhancement only in the left temporal HCoh in the ACCG. The β band, the most reactive band in the NG, was `silent' in the ACCG, suggesting deviant cortical function during motor activity as well.

EEG coherence changes during finger tapping in acallosal and normal children: a study of inter- and intrahemispheric connectivity

The EEG inter- and intrahemispheric coherences (ICoh and HCoh) in the θ, α and β bands were studied in an acallosal group (ACCG) of five children and a normal group of 30 sex- and age-matched children (NG) during resting and tapping conditions. Being functionally deficient, tapping in the ACCG was characterized by increased intertap intervals and variability (in right-hand tapping) and by variability together with decreased synchronization (in bimanual tapping). In the ACCG, frontal, central and parietal ICohs were shown to be smaller, while temporal ICohs were larger under all conditions (see also Koeda, T., Knyazeva, M., Jonkman, J., Njiokiktjien, C., De Sonneville, L., Vildavsky, V., 1995. The resting EEG in acallosal children: compensatory left hemisphere mechanisms? Electroencephalogr. Clin. Neurophysiol. 95, 397–407). The effect was most pronounced in the EEG β band. The sagittal HCohs, including fronto-central, fronto-parietal, and centro-parietal HCohs within both hemispheres, were larger in the ACCG, whereas temporal HCoh (fronto-temporal, centro-temporal, parieto-temporal and occipito-temporal) were smaller, suggesting rearrangement of intracortical activity associated with callosal agenesis. Tapping induced an increase in ICoh and HCoh between frontal, central and parietal areas in the NG, and weak enhancement only in the left temporal HCoh in the ACCG. The β band, the most reactive band in the NG, was `silent' in the ACCG, suggesting deviant cortical function during motor activity as well.

Left-hemisphere specialization for the processing of acoustic transients.

Previous work suggests that speech sounds incorporating short-duration spectral changes (such as the formation transitions of stop consonants) rely on left hemisphere mechanisms for their adequate processing to a greater degree than do speech sounds incorporating spectral changes of a longer duration (such as vowel sounds). Ten normal subjects were scanned using positron emission tomography while discriminating pure tone stimuli incorporating frequency glides of either short or long duration. A comparison of these two conditions yielded significant activation foci in left orbitofrontal cortex, left fusiform gyrus, and right cerebellum. Because non-linguistic stimuli were used, these foci must reflect some basic low level aspect of neural processing that may be relevant to speech but cannot be a consequence of accessing the speech system itself.