Integrating when and what information in the left parietal lobe allows language rule generalization.

A crucial aspect when learning a language is discovering the rules that govern how words are combined in order to convey meanings. Because rules are characterized by sequential co-occurrences between elements (e.g., "These cupcakes are unbelievable"), tracking the statistical relationships between these elements is fundamental. However, purely bottom-up statistical learning alone cannot fully account for the ability to create abstract rule representations that can be generalized, a paramount requirement of linguistic rules. Here, we provide evidence that, after the statistical relations between words have been extracted, the engagement of goal-directed attention is key to enable rule generalization. Incidental learning performance during a rule-learning task on an artificial language revealed a progressive shift from statistical learning to goal-directed attention. In addition, and consistent with the recruitment of attention, functional MRI (fMRI) analyses of late learning stages showed left parietal activity within a broad bilateral dorsal frontoparietal network. Critically, repetitive transcranial magnetic stimulation (rTMS) on participants' peak of activation within the left parietal cortex impaired their ability to generalize learned rules to a structurally analogous new language. No stimulation or rTMS on a nonrelevant brain region did not have the same interfering effect on generalization. Performance on an additional attentional task showed that this rTMS on the parietal site hindered participants' ability to integrate "what" (stimulus identity) and "when" (stimulus timing) information about an expected target. The present findings suggest that learning rules from speech is a two-stage process: following statistical learning, goal-directed attention-involving left parietal regions-integrates "what" and "when" stimulus information to facilitate rapid rule generalization.

Common behavioral clusters and subcortical anatomy in stroke.

A long-held view is that stroke causes many distinct neurological syndromes due to damage of specialized cortical and subcortical centers. However, it is unknown if a syndrome-based description is helpful in characterizing behavioral deficits across a large number of patients. We studied a large prospective sample of first-time stroke patients with heterogeneous lesions at 1-2 weeks post-stroke. We measured behavior over multiple domains and lesion anatomy with structural MRI and a probabilistic atlas of white matter pathways. Multivariate methods estimated the percentage of behavioral variance explained by structural damage. A few clusters of behavioral deficits spanning multiple functions explained neurological impairment. Stroke topography was predominantly subcortical, and disconnection of white matter tracts critically contributed to behavioral deficits and their correlation. The locus of damage explained more variance for motor and language than memory or attention deficits. Our findings highlight the need for better models of white matter damage on cognition.

Large-scale changes in network interactions as a physiological signature of spatial neglect.

The relationship between spontaneous brain activity and behaviour following focal injury is not well understood. Here, we report a large-scale study of resting state functional connectivity MRI and spatial neglect following stroke in a large (n=84) heterogeneous sample of first-ever stroke patients (within 1-2 weeks). Spatial neglect, which is typically more severe after right than left hemisphere injury, includes deficits of spatial attention and motor actions contralateral to the lesion, and low general attention due to impaired vigilance/arousal. Patients underwent structural and resting state functional MRI scans, and spatial neglect was measured using the Posner spatial cueing task, and Mesulam and Behavioural Inattention Test cancellation tests. A principal component analysis of the behavioural tests revealed a main factor accounting for 34% of variance that captured three correlated behavioural deficits: visual neglect of the contralesional visual field, visuomotor neglect of the contralesional field, and low overall performance. In an independent sample (21 healthy subjects), we defined 10 resting state networks consisting of 169 brain regions: visual-fovea and visual-periphery, sensory-motor, auditory, dorsal attention, ventral attention, language, fronto-parietal control, cingulo-opercular control, and default mode. We correlated the neglect factor score with the strength of resting state functional connectivity within and across the 10 resting state networks. All damaged brain voxels were removed from the functional connectivity:behaviour correlational analysis. We found that the correlated behavioural deficits summarized by the factor score were associated with correlated multi-network patterns of abnormal functional connectivity involving large swaths of cortex. Specifically, dorsal attention and sensory-motor networks showed: (i) reduced interhemispheric functional connectivity; (ii) reduced anti-correlation with fronto-parietal and default mode networks in the right hemisphere; and (iii) increased intrahemispheric connectivity with the basal ganglia. These patterns of functional connectivity:behaviour correlations were stronger in patients with right- as compared to left-hemisphere damage and were independent of lesion volume. Our findings identify large-scale changes in resting state network interactions that are a physiological signature of spatial neglect and may relate to its right hemisphere lateralization.

Dorsal and ventral attention systems underlie social and symbolic cueing.

Eye gaze is a powerful cue for orienting attention in space. Studies examining whether gaze and symbolic cues recruit the same neural mechanisms have found mixed results. We tested whether there is a specialized attentional mechanism for social cues. We separately measured BOLD activity during orienting and reorienting attention following predictive gaze and symbolic cues. Results showed that gaze and symbolic cues exerted their influence through the same neural networks but also produced some differential modulations. Dorsal frontoparietal regions in left intraparietal sulcus (IPS) and bilateral MT(+)/lateral occipital cortex only showed orienting effects for symbolic cues, whereas right posterior IPS showed larger validity effects following gaze cues. Both exceptions may reflect the greater automaticity of gaze cues: Symbolic orienting may require more effort, while disengaging attention during reorienting may be more difficult following gaze cues. Face-selective regions, identified with a face localizer, showed selective activations for gaze cues reflecting sensory processing but no attentional modulations. Therefore, no evidence was found linking face-selective regions to a hypothetical, specialized mechanism for orienting attention to gaze cues. However, a functional connectivity analysis showed greater connectivity between face-selective regions and right posterior IPS, posterior STS, and inferior frontal gyrus during gaze cueing, consistent with proposals that face-selective regions may send gaze signals to parts of the dorsal and ventral frontoparietal attention networks. Finally, although the default-mode network is thought to be involved in social cognition, this role does not extend to gaze orienting as these regions were more deactivated following gaze cues and showed less functional connectivity with face-selective regions during gaze cues.

False Belief vs. False Photographs: A Test of Theory of Mind or Working Memory?

Theory of mind (ToM), the ability to reason about other people's thoughts and beliefs, has been traditionally studied in behavioral and neuroimaging experiments by comparing performance in "false belief" and "false photograph" (control) stories. However, some evidence suggests that these stories are not matched in difficulty, complicating the interpretation of results. Here, we more fully evaluated the relative difficulty of comprehending these stories and drawing inferences from them. Subjects read false belief and false photograph stories followed by comprehension questions that probed true ("reality" questions) or false beliefs ("representation" questions) appropriate to the stories. Stories and comprehension questions were read and answered, respectively, more slowly in the false photograph than false belief conditions, indicating their greater difficulty. Interestingly, accuracy on representation questions for false photograph stories was significantly lower than for all other conditions and correlated positively with participants' working memory span scores. These results suggest that drawing representational inferences from false photo stories is particularly difficult and places heavy demands on working memory. Extensive naturalistic practice with ToM reasoning may enable a more flexible and efficient mental representation of false belief stories, resulting in lower memory load requirements. An important implication of these results is that the differential modulation of right temporal-parietal junction (RTPJ) during ToM and "false photo" control conditions may reflect the documented negative correlation of RTPJ activity with working memory load rather than a specialized involvement in ToM processes.

Attention and anxiety: different attentional functioning under state and trait anxiety.

Anxiety modulates the functioning of attention. Although the existence of this relationship is clear, its nature is still poorly defined. Added are the facts that different types of anxiety--state or trait--may influence attention differently and that attention is not a unitary system. We studied the influence of such types of anxiety by means of a task that, using emotionally neutral information, assesses the efficiency of three attentional networks: orienting, alerting, and executive control. Results showed a double dissociation. Trait anxiety was related to deficiencies in the executive control network, but state anxiety was associated with an overfunctioning of the alerting and orienting networks.

Green love is ugly: emotions elicited by synesthetic grapheme-color perceptions.

Synesthetes who experience grapheme-color synesthesia often report feeling uneasy when dealing with incongruently colored graphemes although no empirical data is available to confirm this phenomenon. We studied this affective reaction related to synesthetic perceptions by means of an evaluation task. We found that the perception of an incorrectly colored word affects the judgments of emotional valence. Furthermore, this effect competed with the word's emotional valence in a categorization task thus supporting the automatic nature of this synesthetically elicited affective reaction. When manipulating word valence and word color-photism congruence, we found that responses were slower (and less accurate) for inconsistent conditions than for consistent conditions. Inconsistent conditions were defined as those where semantics and color-photism congruence did not produce a similar assessment and therefore gave rise to a negative affective reaction (i.e., positive-valence words presented in a color different from the synesthete's photism or negative-valence words presented in the photism's color). We therefore observed a modulation of the congruency effect (i.e., faster reaction times to congruently colored words than incongruently colored words). Although this congruence effect has been taken as an index of the true experience of synesthesia, we observed that it can be reversed when the experimental manipulations turn an incongruently colored word into a consistent stimulus. To our knowledge, this is the first report of an affective reaction elicited by the congruency between the synesthetically induced color of a word and the color in which the word is actually presented. The underlying neural mechanisms that might be involved in this phenomenon are discussed.

Ovals of time: time-space associations in synaesthesia.

We examine a condition in which units of time, such as months of the year, are associated with specific locations in space. For individuals with this time-space synaesthesia, contiguous time units such as months are spatially linked forming idiosyncratically shaped patterns such as ovals, oblongs or circles. For some individuals, each time unit appears in a highly specific colour. For instance, one of the synaesthetes we studied experienced December as a red area located at arms length to the left of their body. For the same individual May was a blue area located roughly at arms length to the right of their body. We studied four synaesthetes who report spatial associations for the months of the year. We found that the time-space associations experienced by these individuals were consistent across test-retest. In addition, month names directed visual attention to particular locations in space. For some synaesthetes, this directing of spatial attention was quite rapid-in accord with their reports that month names involuntarily bring to mind spatial locations.

Automatic perception and synaesthesia: evidence from colour and photism naming in a stroop-negative priming task.

It is widely assumed that synaesthetic perception is highly automatic, as shown by Stroop test. Furthermore, it has been shown that, although automatic, it can be suppressed leading to Negative Priming (NP). However, these assumptions have not been consistently investigated, as not many papers have measured Stroop in synaesthesia, and only one used a NP procedure. Two experiments were carried out in a female synaesthete (MA), and 13 control participants, in which numbers and letters were displayed in colours either congruent or incongruent with MA's photisms. In contrast to control participants, MA showed significant Stroop effect both when naming the colours and when naming the photisms (slower RT when naming a colour or photism that was incongruently coloured versus congruently coloured). For comparison, we also report a control experiment in which the first letters of colour names were displayed in either congruent (e.g., B in blue) or incongruent (e.g., B in red) colours. Significant Stroop and NP effects were found when a control group named the displayed colour of these letters. The synaesthesic Stroop effect shown by MA was greater than that observed in the Control Experiment when MA was to name the displayed colour, but smaller when she was to name the photism of the stimuli. Regarding NP, MA showed an effect similar to that observed in the Control Experiment, but only when she was to name the photisms of the stimuli. Altogether, these results show that synaesthetic perception is highly automatic and difficult to inhibit.

Modulations among the alerting, orienting and executive control networks.

This paper reports a series of experiments that were carried out in order to study the attentional system. Three networks make up this system, and each of them specializes in particular processes. The executive control network specializes in control processes, such as conflict resolution or detection of errors; the orienting network directs the processing system to the source of input and enhances its processing; the alerting network prepares the system for a fast response by maintaining an adequate level of activation in the cognitive system. Recently, Fan and collaborators [J Cogn Neurosci 14(3):340-347, 2002] designed a task to measure the efficiency of each network. We modified Fan's task to test the influences among the networks. We found that the executive control network is inhibited by the alerting network, whereas the orienting network raises the efficiency of the executive control network (Experiment 1). We also found that the alerting network influences the orienting network by speeding up its time course function (Experiment 2). Results were replicated in a third experiment, proving the effects to be stable over time, participants and experimental context, and to be potentially important as a tool for neuropsychological assessment.

The three attentional networks: on their independence and interactions.

The present investigation was aimed to the study of the three attentional networks (Alerting, Orienting, and Executive Function) and their interactions. A modification of the task developed by Fan, McCandliss, Sommer, Raz, and Posner (2002) was used, in which a cost and benefit paradigm was combined with a flanker task and an alerting signal. We obtained significant interactions as predicted. The alerting network seemed to inhibit the executive function network (a larger flanker-congruency effect was found on trials where an alerting signal had been previously presented). The orienting network influenced the executive function network in a positive way (the flanker effect was smaller for cued than for uncued trials). Finally, alertness increased orienting (the cueing effect was bigger after the alerting signal). This last result, taken together with previous findings, points to an influence in the sense of a faster orienting under alertness, rather than a larger one. These results offer new insight into the functioning of the attentional system.

Normas Asociativas Intracategoriales para 612 Palabras de Seis Categorías Semánticas en Español / Intracategorical associative norms for 612 words from six semantic categories in Spanish

Se obtuvieron datos de fuerza asociativa intracategorial y de familiaridad subjetiva para 612 palabras pertenecientes a seis categorías diferentes. Estas palabras fueron evaluadas por un grupo diferente de personas para cada una de las medidas. Se realizó un análisis de los siguientes índices por categorías: número de asociados, fuerza asociativa del primer y segundo asociados, índice resultante de la sustracción de los dos primeros asociados, porcentaje de respuestas idiosincrásicas, porcentaje de respuestas en blanco o no válidas e índice de la familiaridad subjetiva. Los resultados apuntan a una mayor consistencia interna (menor número de asociados, mayor fuerza asociativa de éstos y menor porcentaje de respuestas idiosincrásicas) de dos de las seis categorías ("Partes del Cuerpo Humano" y "Animales") y a una menor consistencia interna de la categoría "Ropa, Calzado y Complementos". Estas normas asociativas son de interés para investigadores dedicados al estudio de la memoria semántica así como de las diferencias hemisféricas en la ejecución de tareas relacionadas con estímulos verbales (AU)