Dehydration affects brain structure and function in healthy adolescents.

It was recently observed that dehydration causes shrinkage of brain tissue and an associated increase in ventricular volume. Negative effects of dehydration on cognitive performance have been shown in some but not all studies, and it has also been reported that an increased perceived effort may be required following dehydration. However, the effects of dehydration on brain function are unknown. We investigated this question using functional magnetic resonance imaging (fMRI) in 10 healthy adolescents (mean age = 16.8, five females). Each subject completed a thermal exercise protocol and nonthermal exercise control condition in a cross-over repeated measures design. Subjects lost more weight via perspiration in the thermal exercise versus the control condition (P < 0.0001), and lateral ventricle enlargement correlated with the reduction in body mass (r = 0.77, P = 0.01). Dehydration following the thermal exercise protocol led to a significantly stronger increase in fronto-parietal blood-oxygen-level-dependent (BOLD) response during an executive function task (Tower of London) than the control condition, whereas cerebral perfusion during rest was not affected. The increase in BOLD response after dehydration was not paralleled by a change in cognitive performance, suggesting an inefficient use of brain metabolic activity following dehydration. This pattern indicates that participants exerted a higher level of neuronal activity in order to achieve the same performance level. Given the limited availability of brain metabolic resources, these findings suggest that prolonged states of reduced water intake may adversely impact executive functions such as planning and visuo-spatial processing.

Multisensory representation of limb position in human premotor cortex.

Using functional magnetic resonance imaging (fMRI) in humans, we identified regions of cortex involved in the encoding of limb position. Tactile stimulation of the right hand, across the body midline, activated the right parietal cortex when the eyes were closed; activation shifted to a left parietofrontal network when the eyes were open. These data reveal important similarities between human and non-human primates in the network of brain areas involved in the multisensory representation of limb position.

Tool-use: capturing multisensory spatial attention or extending multisensory peripersonal space?

The active and skilful use of tools has been claimed to lead to the "extension" of the visual receptive fields of single neurons representing peripersonal space--the visual space immediately surrounding one's body parts. While this hypothesis provides an attractive and potentially powerful explanation for one neural basis of tool-use behaviours in human and nonhuman primates, a number of competing hypotheses for the reported behavioural effects of tool-use have not yet been subjected to empirical test. Here, we report five behavioural experiments in healthy human participants (n=120) involving the effects of tool-use on visual-tactile interactions in peripersonal space. Specifically, we address the possibility that the use of only a single tool, which is typical of many neuropsychological studies of tool-use, induces a spatial allocation of attention towards the side where the tool is held. Participants' tactile discrimination responses were more strongly affected by visual stimuli presented on the right side when they held a single tool on the right, compared to visual stimuli presented on the left. When [corrected] two tools were held, one in each hand, this spatial effect disappeared. Our results are incompatible with the hypothesis that tool-use extends peripersonal space, and suggest instead that the use and/or manipulation of [corrected] tools results in an automatic multisensory shift of spatial attention to the side of space where the tip of the tool is actively held. These results have implications for many of the cognitive neuroscientific studies of tool-use published to date.

Tool use changes multisensory interactions in seconds: evidence from the crossmodal congruency task.

Active tool use in human and non-human primates has been claimed to alter the neural representations of multisensory peripersonal space. To date, most studies suggest that a short period of tool use leads to an expansion or elongation of these spatial representations, which lasts several minutes after the last tool use action. However, the possibility that multisensory interactions also change on a much shorter time scale following or preceding individual tool use movements has not yet been investigated. We measured crossmodal (visual-tactile) congruency effects as an index of multisensory integration during two tool use tasks. In the regular tool use task, the participants used one of two tools in a spatiotemporally predictable sequence after every fourth crossmodal congruency trial. In the random tool use task, the required timing and spatial location of the tool use task varied unpredictably. Multisensory integration effects increased as a function of the number of trials since tool use in the regular tool use group, but remained relatively constant in the random tool use group. The spatial distribution of these multisensory effects, however, was unaffected by tool use predictability, with significant spatial interactions found only near the hands and at the tips of the tools. These data suggest that endogenously preparing to use a tool enhances visual-tactile interactions near the tools. Such enhancements are likely due to the increased behavioural relevance of visual stimuli as each tool use action is prepared before execution.

Multisensory interactions follow the hands across the midline: evidence from a non-spatial visual-tactile congruency task.

Crossing the hands over, whether across the body midline or with respect to each other, leads to measurable changes in spatial compatibility, spatial attention, and frequently to a general decrement in discrimination performance for tactile stimuli. The majority of multisensory crossed hands effects, however, have been demonstrated with explicit or implicit spatial discrimination tasks, raising the question of whether non-spatial discrimination tasks also show spatial effects when the hands are crossed. We designed a novel, non-spatial tactile discrimination task to address this issue. Participants made speeded discriminations of single- versus double-pulse vibrotactile targets, while trying to ignore simultaneous visual distractor stimuli, in both hands uncrossed and hands crossed postures. Tactile discrimination performance was significantly affected by the visual distractors (demonstrating a significant crossmodal congruency effect) and was affected most by visual distractors in the same external location as the tactile target (i.e., spatial modulation), regardless of the posture (uncrossed or crossed) of the hands (i.e., spatial 'remapping' of visual-tactile interactions). Finally, crossing the hands led to a general performance decrement with visual distractors, but not in a control task with unimodal visual or tactile judgements. These results demonstrate, for the first time, significant spatial and postural modulations of crossmodal congruency effects in a non-spatial discrimination task.

Evidence for asymmetric frontal-lobe involvement in episodic memory from functional magnetic resonance imaging and patients with unilateral frontal-lobe excisions.

Recently, there has been considerable debate regarding the involvement of the left and right prefrontal cortices in the encoding and retrieval of episodic memory. In a previous PET study, we found that the use of easily verbalisable material may lead to activation predominantly in the left lateral frontal cortex whilst the use of non-easily verbalisable material may lead to activation predominantly in the right lateral frontal cortex, in both cases irrespective of encoding and retrieval processes. In order to replicate and extend these findings, the same task was modified for use with fMRI. Six healthy volunteers were scanned while encoding and then recalling stimuli that either emphasised visual or verbal processes. It was found that, in comparison to a baseline condition, the encoding of visual stimuli led to a bilateral activation of the prefrontal cortex whilst the encoding of verbal stimuli led to a preferential activation of the left prefrontal cortex. An effect of stimulus type was less evident during retrieval, with both visual and verbal stimuli leading to bilateral prefrontal cortex activation. Overall, encoding and retrieval activated similar regions of the prefrontal cortex suggesting that these areas mediate processes that are fundamental to both aspects of memory. To extend these findings further, the tasks used in the fMRI study were used to assess a group of patients with unilateral frontal lesions and a group of healthy control volunteers. The patients were significantly impaired compared to the healthy volunteers, although no significant differences were found in performance between the right- and left-sided lesioned patients. This result suggests that the memory-related asymmetries observed during functional neuroimaging studies may not be critical for task performance.

Speechreading circuits in people born deaf.

In hearing people, silent speechreading generates bilateral activation in superior temporal regions specialised for the perception of auditory speech [Science 276 (1997) 593; Neuroreport 11 (2000) 1729; Proceedings of the Royal Society London B 268 (2001) 451]. In the present study, FMRI data were collected from deaf and hearing volunteers while they speechread numbers and during a control task in which they counted nonsense mouth movements (gurns). Brain activation for silent speechreading in oral deaf participants was found primarily in posterior cingulate cortex and hippocampal/lingual gyri. In contrast to the pattern observed in the hearing group, deaf participants showed no speechreading-specific activation in left lateral temporal regions. These data suggest that acoustic experience shapes the functional circuits for analysing speech. We speculate on the functional role, the posterior cingulate gyrus may play in speechreading by profoundly congenitally deaf people.

Multisensory integration: methodological approaches and emerging principles in the human brain.

Understanding the conditions under which the brain integrates the different sensory streams and the mechanisms supporting this phenomenon is now a question at the forefront of neuroscience. In this paper, we discuss the opportunities for investigating these multisensory processes using modern imaging techniques, the nature of the information obtainable from each method and their benefits and limitations. Despite considerable variability in terms of paradigm design and analysis, some consistent findings are beginning to emerge. The detection of brain activity in human neuroimaging studies that resembles multisensory integration responses at the cellular level in other species, suggests similar crossmodal binding mechanisms may be operational in the human brain. These mechanisms appear to be distributed across distinct neuronal networks that vary depending on the nature of the shared information between different sensory cues. For example, differing extents of correspondence in time, space or content seem to reliably bias the involvement of different integrative networks which code for these cues. A combination of data obtained from haemodynamic and electromagnetic methods, which offer high spatial or temporal resolution respectively, are providing converging evidence of multisensory interactions at both "early" and "late" stages of processing--suggesting a cascade of synergistic processes operating in parallel at different levels of the cortex.

Extending or projecting peripersonal space with tools? Multisensory interactions highlight only the distal and proximal ends of tools.

The effects of tool-use on the brain's representation of the body and of the space surrounding the body ('peripersonal space') has recently been studied within a number of disciplines in cognitive neuroscience, and is also of great interest to philosophers and behavioural ecologists. To date, most experimental findings suggest that tool-use extends the boundary of peripersonal space-visual stimuli presented at the tips of tools interact more with simultaneous tactile stimuli presented at the hands than visual stimuli presented at the same distance, but not associated with the tools. We studied the proposed extension of peripersonal space by tool-use by measuring the effects of three different tool-use tasks on the integration of visual and tactile stimuli at three distances from participants' hands along two hand-held tools. When the tool-use task required using the shafts or the tips of the tools, visuotactile interactions were stronger at the tips of the tools than in the middle of the shaft. When the handles of the tools were used, however, visuotactile interactions were strongest near the hands and decreased with distance along the tools. These results suggest that tools do not simply 'extend' peripersonal space, but that just the tips of tools actively manipulated in extrapersonal space are incorporated into the brain's visuotactile representations of the body and of peripersonal space.

Predicting consumer behavior: using novel mind-reading approaches.

Advances in machine learning as applied to functional magnetic resonance imaging (fMRI) data offer the possibility of pretesting and classifying marketing communications using unbiased pattern recognition algorithms. By using these algorithms to analyze brain responses to brands, products, or existing marketing communications that either failed or succeeded in the marketplace and identifying the patterns of brain activity that characterize success or failure, future planned campaigns or new products can now be pretested to determine how well the resulting brain responses match the desired (successful) pattern of brain activity without the need for verbal feedback. This major advance in signal processing is poised to revolutionize the application of these brain-imaging techniques in the marketing sector by offering greater accuracy of prediction in terms of consumer acceptance of new brands, products, and campaigns at a speed that makes them accessible as routine pretesting tools that will clearly demonstrate return on investment.

The multisensory attentional consequences of tool use: a functional magnetic resonance imaging study.

BACKGROUND: Tool use in humans requires that multisensory information is integrated across different locations, from objects seen to be distant from the hand, but felt indirectly at the hand via the tool. We tested the hypothesis that using a simple tool to perceive vibrotactile stimuli results in the enhanced processing of visual stimuli presented at the distal, functional part of the tool. Such a finding would be consistent with a shift of spatial attention to the location where the tool is used. METHODOLOGY/PRINCIPAL FINDINGS: We tested this hypothesis by scanning healthy human participants' brains using functional magnetic resonance imaging, while they used a simple tool to discriminate between target vibrations, accompanied by congruent or incongruent visual distractors, on the same or opposite side to the tool. The attentional hypothesis was supported: BOLD response in occipital cortex, particularly in the right hemisphere lingual gyrus, varied significantly as a function of tool position, increasing contralaterally, and decreasing ipsilaterally to the tool. Furthermore, these modulations occurred despite the fact that participants were repeatedly instructed to ignore the visual stimuli, to respond only to the vibrotactile stimuli, and to maintain visual fixation centrally. In addition, the magnitude of multisensory (visual-vibrotactile) interactions in participants' behavioural responses significantly predicted the BOLD response in occipital cortical areas that were also modulated as a function of both visual stimulus position and tool position. CONCLUSIONS/SIGNIFICANCE: These results show that using a simple tool to locate and to perceive vibrotactile stimuli is accompanied by a shift of spatial attention to the location where the functional part of the tool is used, resulting in enhanced processing of visual stimuli at that location, and decreased processing at other locations. This was most clearly observed in the right hemisphere lingual gyrus. Such modulations of visual processing may reflect the functional importance of visuospatial information during human tool use.

Physical and physiological consequences of passive intra-oral shimming.

Imaging the human orbitofrontal cortex (OFC) with fMRI is problematic due to the proximity of this region to the air-filled sinuses, which causes susceptibility artifacts. Placing a strongly diamagnetic material into the mouth ('mouthshim') of a human volunteer can significantly reduce the artifacts in this region. Using the same combined olfactory and visual fMRI paradigm, we compared brain activation and static B0 field maps of participants being scanned both with and without the 'mouthshim'. Results demonstrate that the device improves the B0 field homogeneity within OFC, resulting in significantly stronger BOLD activation in this region. However, the device also caused both increased head motion and reduced activation in insular cortices due to more frequent swallowing and tactile stimulation of the tongue. The 'mouthshim' should only, therefore, be used where sensitivity in OFC regions is paramount.

Lexical and sentential processing in British Sign Language.

Studies of spoken and written language suggest that the perception of sentences engages the left anterior and posterior temporal cortex and the left inferior frontal gyrus to a greater extent than non-sententially structured material, such as word lists. This study sought to determine whether the same is true when the language is gestural and perceived visually. Regional neural activity was measured using functional MRI while Deaf and hearing native signers of British Sign Language (BSL) detected semantic anomalies in well-formed BSL sentences and when they detected nonsense signs in lists of unconnected BSL signs. Processing BSL sentences, when contrasted with signed lists, was reliably associated with greater activation in the posterior portions of the left middle and superior temporal gyri and in the left inferior frontal cortex, but not in the anterior temporal cortex, which was activated to a similar extent whether lists or sentences were processed. Further support for the specificity of these areas for processing the linguistic-rather than visuospatial-features of signed sentences came from a contrast of hearing native signers and hearing sign-naïve participants. Hearing signers recruited the left posterior temporal and inferior frontal regions during BSL sentence processing to a greater extent than hearing non-signers. These data suggest that these left perisylvian regions are differentially associated with sentence processing, whatever the modality of the linguistic input.

Perceiving identical sounds as speech or non-speech modulates activity in the left posterior superior temporal sulcus.

The left superior temporal cortex shows greater responsiveness to speech than to non-speech sounds according to previous neuroimaging studies, suggesting that this brain region has a special role in speech processing. However, since speech sounds differ acoustically from the non-speech sounds, it is possible that this region is not involved in speech perception per se, but rather in processing of some complex acoustic features. "Sine wave speech" (SWS) provides a tool to study neural speech specificity using identical acoustic stimuli, which can be perceived either as speech or non-speech, depending on previous experience of the stimuli. We scanned 21 subjects using 3T functional MRI in two sessions, both including SWS and control stimuli. In the pre-training session, all subjects perceived the SWS stimuli as non-speech. In the post-training session, the identical stimuli were perceived as speech by 16 subjects. In these subjects, SWS stimuli elicited significantly stronger activity within the left posterior superior temporal sulcus (STSp) in the post- vs. pre-training session. In contrast, activity in this region was not enhanced after training in 5 subjects who did not perceive SWS stimuli as speech. Moreover, the control stimuli, which were always perceived as non-speech, elicited similar activity in this region in both sessions. Altogether, the present findings suggest that activation of the neural speech representations in the left STSp might be a pre-requisite for hearing sounds as speech.

Color of scents: chromatic stimuli modulate odor responses in the human brain.

Color has a profound effect on the perception of odors. For example, strawberry-flavored drinks smell more pleasant when colored red than green and descriptions of the "nose" of a wine are dramatically influenced by its color. Using functional magnetic resonance imaging, we demonstrate a neurophysiological correlate of these cross-modal visual influences on olfactory perception. Subjects were scanned while exposed either to odors or colors in isolation or to color-odor combinations that were rated on the basis of how well they were perceived to match. Activity in caudal regions of the orbitofrontal cortex and in the insular cortex increased progressively with the perceived congruency of the odor-color pairs. These findings demonstrate the neuronal correlates of olfactory response modulation by color cues in brain areas previously identified as encoding the hedonic value of smells.

Reading speech from still and moving faces: the neural substrates of visible speech.

Speech is perceived both by ear and by eye. Unlike heard speech, some seen speech gestures can be captured in stilled image sequences. Previous studies have shown that in hearing people, natural time-varying silent seen speech can access the auditory cortex (left superior temporal regions). Using functional magnetic resonance imaging (fMRI), the present study explored the extent to which this circuitry was activated when seen speech was deprived of its time-varying characteristics. In the scanner, hearing participants were instructed to look for a prespecified visible speech target sequence ("voo" or "ahv") among other monosyllables. In one condition, the image sequence comprised a series of stilled key frames showing apical gestures (e.g., separate frames for "v" and "oo" [from the target] or "ee" and "m" [i.e., from nontarget syllables]). In the other condition, natural speech movement of the same overall segment duration was seen. In contrast to a baseline condition in which the letter "V" was superimposed on a resting face, stilled speech face images generated activation in posterior cortical regions associated with the perception of biological movement, despite the lack of apparent movement in the speech image sequence. Activation was also detected in traditional speech-processing regions including the left inferior frontal (Broca's) area, left superior temporal sulcus (STS), and left supramarginal gyrus (the dorsal aspect of Wernicke's area). Stilled speech sequences also generated activation in the ventral premotor cortex and anterior inferior parietal sulcus bilaterally. Moving faces generated significantly greater cortical activation than stilled face sequences, and in similar regions. However, a number of differences between stilled and moving speech were also observed. In the visual cortex, stilled faces generated relatively more activation in primary visual regions (V1/V2), while visual movement areas (V5/MT+) were activated to a greater extent by moving faces. Cortical regions activated more by naturally moving speaking faces included the auditory cortex (Brodmann's Areas 41/42; lateral parts of Heschl's gyrus) and the left STS and inferior frontal gyrus. Seen speech with normal time-varying characteristics appears to have preferential access to "purely" auditory processing regions specialized for language, possibly via acquired dynamic audiovisual integration mechanisms in STS. When seen speech lacks natural time-varying characteristics, access to speech-processing systems in the left temporal lobe may be achieved predominantly via action-based speech representations, realized in the ventral premotor cortex.

Neural systems underlying British Sign Language and audio-visual English processing in native users.

In order to understand the evolution of human language, it is necessary to explore the neural systems that support language processing in its many forms. In particular, it is informative to separate those mechanisms that may have evolved for sensory processing (hearing) from those that have evolved to represent events and actions symbolically (language). To what extent are the brain systems that support language processing shaped by auditory experience and to what extent by exposure to language, which may not necessarily be acoustically structured? In this first neuroimaging study of the perception of British Sign Language (BSL), we explored these questions by measuring brain activation using functional MRI in nine hearing and nine congenitally deaf native users of BSL while they performed a BSL sentence-acceptability task. Eight hearing, non-signing subjects performed an analogous task that involved audio-visual English sentences. The data support the argument that there are both modality-independent and modality-dependent language localization patterns in native users. In relation to modality-independent patterns, regions activated by both BSL in deaf signers and by spoken English in hearing non-signers included inferior prefrontal regions bilaterally (including Broca's area) and superior temporal regions bilaterally (including Wernicke's area). Lateralization patterns were similar for the two languages. There was no evidence of enhanced right-hemisphere recruitment for BSL processing in comparison with audio-visual English. In relation to modality-specific patterns, audio-visual speech in hearing subjects generated greater activation in the primary and secondary auditory cortices than BSL in deaf signers, whereas BSL generated enhanced activation in the posterior occipito-temporal regions (V5), reflecting the greater movement component of BSL. The influence of hearing status on the recruitment of sign language processing systems was explored by comparing deaf and hearing adults who had BSL as their first language (native signers). Deaf native signers demonstrated greater activation in the left superior temporal gyrus in response to BSL than hearing native signers. This important finding suggests that left- temporal auditory regions may be privileged for processing heard speech even in hearing native signers. However, in the absence of auditory input this region can be recruited for visual processing.

Neural correlates of British sign language comprehension: spatial processing demands of topographic language.

In all signed languages used by deaf people, signs are executed in "sign space" in front of the body. Some signed sentences use this space to map detailed "real-world" spatial relationships directly. Such sentences can be considered to exploit sign space "topographically." Using functional magnetic resonance imaging, we explored the extent to which increasing the topographic processing demands of signed sentences was reflected in the differential recruitment of brain regions in deaf and hearing native signers of the British Sign Language. When BSL signers performed a sentence anomaly judgement task, the occipito-temporal junction was activated bilaterally to a greater extent for topographic than nontopographic processing. The differential role of movement in the processing of the two sentence types may account for this finding. In addition, enhanced activation was observed in the left inferior and superior parietal lobules during processing of topographic BSL sentences. We argue that the left parietal lobe is specifically involved in processing the precise configuration and location of hands in space to represent objects, agents, and actions. Importantly, no differences in these regions were observed when hearing people heard and saw English translations of these sentences. Despite the high degree of similarity in the neural systems underlying signed and spoken languages, exploring the linguistic features which are unique to each of these broadens our understanding of the systems involved in language comprehension.

Dissociating linguistic and nonlinguistic gestural communication in the brain.

Gestures of the face, arms, and hands are components of signed languages used by Deaf people. Signaling codes, such as the racecourse betting code known as Tic Tac, are also made up of such gestures. Tic Tac lacks the phonological structure of British Sign Language (BSL) but is similar in terms of its visual and articulatory components. Using fMRI, we compared the neural correlates of viewing a gestural language (BSL) and a manual-brachial code (Tic Tac) relative to a low-level baseline task. We compared three groups: Deaf native signers, hearing native signers, and hearing nonsigners. None of the participants had any knowledge of Tic Tac. All three groups activated an extensive frontal-posterior network in response to both types of stimuli. Superior temporal cortex, including the planum temporale, was activated bilaterally in response to both types of gesture in all groups, irrespective of hearing status. The engagement of these traditionally auditory processing regions was greater in Deaf than hearing participants. These data suggest that the planum temporale may be responsive to visual movement in both deaf and hearing people, yet when hearing is absent early in development, the visual processing role of this region is enhanced. Greater activation for BSL than Tic Tac was observed in signers, but not in nonsigners, in the left posterior superior temporal sulcus and gyrus, extending into the supramarginal gyrus. This suggests that the left posterior perisylvian cortex is of fundamental importance to language processing, regardless of the modality in which it is conveyed.