Group and individual variability in speech production networks during delayed auditory feedback.

Altering reafferent sensory information can have a profound effect on motor output. Introducing a short delay [delayed auditory feedback (DAF)] during speech production results in modulations of voice and loudness, and produces a range of speech dysfluencies. The ability of speakers to resist the effects of delayed feedback is variable yet it is unclear what neural processes underlie differences in susceptibility to DAF. Here, susceptibility to DAF is investigated by looking at the neural basis of within and between subject changes in speech fluency under 50 and 200 ms delay conditions. Using functional magnetic resonance imaging, networks involved in producing speech under two levels of DAF were identified, lying largely within networks active during normal speech production. Independent of condition, fluency ratings were associated with midbrain activity corresponding to periaqueductal grey matter. Across subject variability in ability to produce normal sounding speech under a 200 ms delay was associated with activity in ventral sensorimotor cortices, whereas ability to produce normal sounding speech under a 50 ms delay was associated with left inferior frontal gyrus activity. These data indicate whilst overlapping cortical mechanisms are engaged for speaking under different delay conditions, susceptibility to different temporal delays in speech feedback may involve different processes.

Individual differences in laughter perception reveal roles for mentalizing and sensorimotor systems in the evaluation of emotional authenticity.

Humans express laughter differently depending on the context: polite titters of agreement are very different from explosions of mirth. Using functional MRI, we explored the neural responses during passive listening to authentic amusement laughter and controlled, voluntary laughter. We found greater activity in anterior medial prefrontal cortex (amPFC) to the deliberate, Emitted Laughs, suggesting an obligatory attempt to determine others' mental states when laughter is perceived as less genuine. In contrast, passive perception of authentic Evoked Laughs was associated with greater activity in bilateral superior temporal gyri. An individual differences analysis found that greater accuracy on a post hoc test of authenticity judgments of laughter predicted the magnitude of passive listening responses to laughter in amPFC, as well as several regions in sensorimotor cortex (in line with simulation accounts of emotion perception). These medial prefrontal and sensorimotor sites showed enhanced positive connectivity with cortical and subcortical regions during listening to involuntary laughter, indicating a complex set of interacting systems supporting the automatic emotional evaluation of heard vocalizations.

Do sentences with unaccusative verbs involve syntactic movement? Evidence from neuroimaging.

This study focuses on the neural processing of English sentences containing unergative, unaccusative and transitive verbs. We demonstrate common responses in bilateral superior temporal gyri in response to listening to sentences containing unaccusative and transitive verbs compared to unergative verbs; we did not detect any activation that was specific to unaccusatives. Our findings indicate that the neural processing of unaccusative and transitive verbs is highly similar, and very different from the processing of unergative verbs. We discuss the consequences of these results for the linguistic analysis of movement phenomena.

The pathways for intelligible speech: multivariate and univariate perspectives.

An anterior pathway, concerned with extracting meaning from sound, has been identified in nonhuman primates. An analogous pathway has been suggested in humans, but controversy exists concerning the degree of lateralization and the precise location where responses to intelligible speech emerge. We have demonstrated that the left anterior superior temporal sulcus (STS) responds preferentially to intelligible speech (Scott SK, Blank CC, Rosen S, Wise RJS. 2000. Identification of a pathway for intelligible speech in the left temporal lobe. Brain. 123:2400-2406.). A functional magnetic resonance imaging study in Cerebral Cortex used equivalent stimuli and univariate and multivariate analyses to argue for the greater importance of bilateral posterior when compared with the left anterior STS in responding to intelligible speech (Okada K, Rong F, Venezia J, Matchin W, Hsieh IH, Saberi K, Serences JT,Hickok G. 2010. Hierarchical organization of human auditory cortex: evidence from acoustic invariance in the response to intelligible speech. 20: 2486-2495.). Here, we also replicate our original study, demonstrating that the left anterior STS exhibits the strongest univariate response and, in decoding using the bilateral temporal cortex, contains the most informative voxels showing an increased response to intelligible speech. In contrast, in classifications using local "searchlights" and a whole brain analysis, we find greater classification accuracy in posterior rather than anterior temporal regions. Thus, we show that the precise nature of the multivariate analysis used will emphasize different response profiles associated with complex sound to speech processing.

Articulatory movements modulate auditory responses to speech.

Production of actions is highly dependent on concurrent sensory information. In speech production, for example, movement of the articulators is guided by both auditory and somatosensory input. It has been demonstrated in non-human primates that self-produced vocalizations and those of others are differentially processed in the temporal cortex. The aim of the current study was to investigate how auditory and motor responses differ for self-produced and externally produced speech. Using functional neuroimaging, subjects were asked to produce sentences aloud, to silently mouth while listening to a different speaker producing the same sentence, to passively listen to sentences being read aloud, or to read sentences silently. We show that that separate regions of the superior temporal cortex display distinct response profiles to speaking aloud, mouthing while listening, and passive listening. Responses in anterior superior temporal cortices in both hemispheres are greater for passive listening compared with both mouthing while listening, and speaking aloud. This is the first demonstration that articulation, whether or not it has auditory consequences, modulates responses of the dorsolateral temporal cortex. In contrast posterior regions of the superior temporal cortex are recruited during both articulation conditions. In dorsal regions of the posterior superior temporal gyrus, responses to mouthing and reading aloud were equivalent, and in more ventral posterior superior temporal sulcus, responses were greater for reading aloud compared with mouthing while listening. These data demonstrate an anterior-posterior division of superior temporal regions where anterior fields are suppressed during motor output, potentially for the purpose of enhanced detection of the speech of others. We suggest posterior fields are engaged in auditory processing for the guidance of articulation by auditory information.

Activity monitoring and motion classification of the lizard Chamaeleo jacksonii using multiple Doppler radars.

We describe a simple, non-contact and efficient tool for monitoring the natural activity of a small lizard (Chamaeleo jacksonii) to yield valuable information about their metabolic activity and energy expenditure. It allows monitoring in a non-confined laboratory environment and uses multiple Doppler radars operating at 10.525 GHz. We developed a classification algorithm that can differentiate between fidgeting and locomotion by processing the quadrature baseband signals from the radars. The results have been verified by visual inspection and indicate that the tool could also be used for automated monitoring of the activities of reptiles and other small animals.

Human brain mechanisms for the early analysis of voices.

In this functional magnetic resonance imaging study, we investigated human brain mechanisms that are involved in the analysis of voices as sound sources and in the pre-semantic analysis of voice information. The source of the voice was altered by changing the speaker, and the salience of the voice was altered by changing the amount of spectrotemporal detail. We identified a mechanism for detecting a change in the source of the voice in the posterior superior temporal lobe and anatomically distinct mechanisms for the detailed analysis of voice information in a bilateral network extending from the posterior to the anterior superior temporal lobe surrounding the superior temporal sulcus. The findings are consistent with a processing hierarchy in which general source attributes are analyzed in the posterior superior temporal lobe, abstraction of voice identity features occurs in posterior superior temporal sulcus, and further analysis of voice information occurs in anterior superior temporal sulcus and higher order cortices in the middle and anterior temporal lobe.

Complex behavior in the formaldehyde-sulfite reaction.

The formaldehyde-sulfite reaction is an example of an "acid-to-alkali" clock. It displays an induction period, during which the pH varies only slowly in time, followed by a reaction event, during which the pH increases rapidly by several units. When the reaction is performed in a closed (batch) reactor, the clock time is found to increase with a decrease in initial concentrations of formaldehyde and sulfite and an increase in the total initial concentration of S(IV). At long times, following the clock event, there is a slow decrease in pH. In an open (flow) reactor, bistability between a low-pH steady state (pH approximately 6-8) and a high-pH steady state (pH approximately 11) is observed. Additionally, we report the existence of sustained, small-amplitude oscillations in pH in this system. An extended kinetic mechanism reproduces the batch behavior but fails to account for the complex behavior observed in the flow reactor. Possible additional reaction steps are discussed.

When the central executive lets us down: Schemas, attention, and load in a generative working memory task.

Participants were asked to generate a single sequence of numbers in between two bounds. By varying the requested sequence length and way in which the question is posed, this paradigm enables assessment of the contributions to central executive functioning of schema, focus of attention, and load. With sequences of three or four numbers, a quarter of the sample failed fully to comply with the instructions. They generated an incorrect number of numbers or went outside the specified bounds. With sequences of six numbers, more than half of the sample infringed one or more of the constraints. Participants consistently generated sequences with particular generic properties. The overall frequency and patterns of infringements suggest that a substantial proportion of participants focused their conceptual attention on sequence content and often neglected the problem of how the length and boundary constraints were going to be evaluated either before or during response delivery.

When the central executive lets us down: schemas, attention, and load in a generative working memory task.

Participants were asked to generate a single sequence of numbers in between two bounds. By varying the requested sequence length and way in which the question is posed, this paradigm enables assessment of the contributions to central executive functioning of schema, focus of attention, and load. With sequences of three or four numbers, a quarter of the sample failed fully to comply with the instructions. They generated an incorrect number of numbers or went outside the specified bounds. With sequences of six numbers, more than half of the sample infringed one or more of the constraints. Participants consistently generated sequences with particular generic properties. The overall frequency and patterns of infringements suggest that a substantial proportion of participants focused their conceptual attention on sequence content and often neglected the problem of how thelength and boundary constraints were going to be evaluated either before or during response delivery.

The planning and guiding of reading saccades: a repetitive transcranial magnetic stimulation study.

A previous positron emission tomography study that investigated the cortical areas involved in directing eye movements during text reading showed two areas of extra-occipital asymmetry: left > right posterior parietal cortex (PPC), and right > left frontal eye-field (FEF). We used the temporal resolution of repetitive TMS (rTMS) to isolate the contributions of the left and right PPC and FEF to the planning and execution of rightward reading saccades. We present eye-movement data collected during text reading, which involves the initiation and maintenance of a series of saccades (scanpath). rTMS over the left but not right PPC slowed reading speeds for the whole array of words, indicating that this area is involved throughout the scanpath. rTMS over the right but not the left FEF slowed the time to make the first saccade, but only when triggered before the stimuli appeared, demonstrating that the role of this region is in the preparation of the scanpath. Our results are compatible with the hypotheses that the left PPC maintains reading saccades along a line of text while the right FEF is involved in the preparation of the motor plan for the scanpath at the start of each new line of text.

Specifying executive representations and processes in number generation tasks.

The Interacting Cognitive Subsystems framework, ICS (Barnard, 1985) proposes that central executive phenomena can be accounted for by two autonomous subsystems, which process different forms of meaning: propositional and schematic (implicational) meanings. The apparent supervisory role of the executive arises from limitations on the exchange of information between these and other cognitive subsystems. This general proposal is elaborated in four experiments in which a total of 1,293 participants are asked to spontaneously generate a large verbal number to varying task constraints, with the intention of specifying the representations of number and task that underlie responses. Responses change systematically according to participants' use of explicit propositional information provided by the instructions, and inferred implicational information about what the experimenter is requesting. There was a high error rate (between 6% and 24%), participants producing responses that did not fall within the large range indicated by the instructions. The studies support the distinction between propositional and implicational processing in executive function, and provide a framework for understanding normal executive representations and processes.

The functional anatomy of single-word reading in patients with hemianopic and pure alexia.

We investigated single-word reading in normal subjects and patients with alexia following a left occipital infarct, using PET. The most posterior brain region to show a lateralized response was at the left occipitotemporal junction, in the inferior temporal gyrus. This region was activated when normal subjects, patients with hemianopic alexia and patients with an incomplete right homonymous hemianopia, but no reading deficit, viewed single words presented at increasing rates. This same area was damaged in a patient with pure alexia ("alexia without agraphia") and no hemianopia, who read words slowly using a letter-by-letter strategy. Although the exact level of the functional deficit is controversial, pure alexia is the result of an inability to map a percept of all the letters in a familiar letter string on to the mental representation of the whole word form. However, the commonest deficit associated with "pure" alexia is a right homonymous field defect; an impairment that may, by itself, interfere with single-word reading because of inability to see the letters towards the end of a word. The relative contributions of pure and hemianopic alexia in individual patients needs to be assessed, as the latter has been shown to respond well to specific rehabilitation programmes.

Factors influencing the use of breast reconstruction postmastectomy: a National Cancer Database study.

BACKGROUND: Advances in surgical techniques and changes in our understanding of the biology of breast cancer have made immediate or early breast reconstruction a viable option for the majority of women with breast cancer. Little is known about national patterns of use of reconstruction. This study was undertaken to determine national patterns of care and factors that influence the use of breast reconstruction. STUDY DESIGN: A large convenience sample reported to the National Cancer Data Base was studied. Patients coded as undergoing mastectomy between 1985 and 1990 (n = 155,463) and between 1994 and 1995 (n = 68,348) were evaluated. The use of reconstruction in the two time periods was compared, and patient and tumor factors influencing the use of the procedure were compared. RESULTS: Between 1985 and 1990, 3.4% of mastectomy patients had early or immediate reconstruction, increasing to 8.3% in 1994-5. Patient age, income, geographic location, type of hospital where treatment occurred, and tumor stage all influenced the use of reconstruction in univariate analysis. In multivariate analysis, patients age 50 or under had a 4.3-fold greater likelihood of having reconstruction than their older counterparts. Patients with ductal carcinoma in situ were twice as likely as those with invasive cancer to have reconstruction. Family income of $40,000 or more (Odds Ratio 2.0), ethnicity other than African-American (Odds Ratio 1.6), surgery in a National Cancer Institute-designated cancer center (Odds Ratio 1.4), and surgery in a geographic region other than the Midwest or South (Odds Ratio 1.3) remained significant predictors of the use of reconstruction in multivariate analysis. CONCLUSIONS: Breast reconstruction is an underused option in breast cancer management. Predictors of the use of reconstruction do not reflect contraindications to the procedure, and indicate the need for both physician and patient education.

Separate neural subsystems within 'Wernicke's area'.

Over time, both the functional and anatomical boundaries of 'Wernicke's area' have become so broad as to be meaningless. We have re-analysed four functional neuroimaging (PET) studies, three previously published and one unpublished, to identify anatomically separable, functional subsystems in the left superior temporal cortex posterior to primary auditory cortex. From the results we identified a posterior stream of auditory processing. One part, directed along the supratemporal cortical plane, responded to both non-speech and speech sounds, including the sound of the speaker's own voice. Activity in its most posterior and medial part, at the junction with the inferior parietal lobe, was linked to speech production rather than perception. The second, more lateral and ventral part lay in the posterior left superior temporal sulcus, a region that responded to an external source of speech. In addition, this region was activated by the recall of lists of words during verbal fluency tasks. The results are compatible with an hypothesis that the posterior superior temporal cortex is specialized for processes involved in the mimicry of sounds, including repetition, the specific role of the posterior left superior temporal sulcus being to transiently represent phonetic sequences, whether heard or internally generated and rehearsed. These processes are central to the acquisition of long- term lexical memories of novel words.

Identification of a pathway for intelligible speech in the left temporal lobe.

It has been proposed that the identification of sounds, including species-specific vocalizations, by primates depends on anterior projections from the primary auditory cortex, an auditory pathway analogous to the ventral route proposed for the visual identification of objects. We have identified a similar route in the human for understanding intelligible speech. Using PET imaging to identify separable neural subsystems within the human auditory cortex, we used a variety of speech and speech-like stimuli with equivalent acoustic complexity but varying intelligibility. We have demonstrated that the left superior temporal sulcus responds to the presence of phonetic information, but its anterior part only responds if the stimulus is also intelligible. This novel observation demonstrates a left anterior temporal pathway for speech comprehension.

A thalamo-prefrontal system for representation in executive response choice.

This study demonstrates the neural system potentially involved in the representation of, and choice between, stimulus classifications in an ambiguous, novel, decision-making task. This difficult choice behaviour is taken as an example of a basic executive processing task. Subjects heard sounds that were consonant-vowel combinations that had been distorted and were required to categorize each stimulus as speech-like or not-speech-like. Cerebral activity was measured with positron emission tomography. A neural system (thalamic and medial prefrontal cortical regions) was demonstrated; there was greater activity involved in assigning the sound to the larger class of not-speech-like sounds than to the more restricted category of speech-like sounds. We interpret this activity as reflecting process and representation in a simple central executive task.

Noun imageability and the temporal lobes.

We used positron emission tomography to investigate brain activity in response to hearing or reading nouns of varying imageability. Three experiments were performed. Activity increased with noun imageability in the left mid-fusiform gyrus, the lateral parahippocampal area in humans, and in the rostral medial temporal lobes close to or within perirhinal cortex. The left mid-fusiform activation has been observed in previous imaging studies of single word processing. Its functional significance was variously attributed to semantic processing, visual imagery, encoding episodic memories, or the integration of lexical inputs from different sensory modalities. These hypotheses are not mutually exclusive. The more rostral medial lobe response to noun imageability has not been observed previously. However, lesions in perirhinal cortex impair knowledge about objects in non-human primates, and bilateral rostral ventromedial temporal lobe potentials in response to object nouns were observed with human intracranial recordings. Imageable (object) nouns are learnt with reference to sensory experiences of living and non-living objects, whereas acquisition of the meaning of low imageable (abstract) nouns is more dependent on their context within sentences. Parahippocampal and perirhinal cortices are reciprocally connected with, respectively, second and third order sensory association cortices. We conclude that access to the representations of word meaning is dependent on heteromodal temporal lobe cortex, and that during the acquisition of object nouns one route is established through ventromedial temporal cortical regions that have reciprocal connections with all sensory association cortices.

Impaired reading in patients with right hemianopia.

A left occipital stroke may result in alexia for two reasons, which may coexist depending on the distribution of the lesion. A lesion of the left lateroventral prestriate cortex or its afferents impairs word recognition ("pure" alexia). If the left primary visual cortex or its afferents are destroyed, resulting in a complete right homonymous hemianopia, rightward saccades during text reading are disrupted ("hemianopic" alexia). By using functional imaging, we showed two separate but interdependent systems involved in reading. The first, subserving word recognition, involved the representation of foveal vision in the left and right primary visual cortex and the ventral prestriate cortex. The second system, responsible for the planning and execution of reading saccades, consisted of the representation of right parafoveal vision in the left visual cortex, the bilateral posterior parietal cortex (left > right), and the frontal eye fields (right > left). Disruption of this distributed neural system was demonstrated in patients with severe right homonymous hemianopia, commensurate with their inability to perform normal reading eye movements. Text reading, before processes involved in comprehension, requires the integration of perceptual and motor processes. We have demonstrated these distributed neural systems in normal readers and have shown how a right homonymous hemianopia disrupts the motor preparation of reading saccades during text reading.