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In nature, sounds from objects of interest arrive at the ears accompanied by sound waves from other actively emitting objects and by reflections off of nearby surfaces. Despite the fact that all of these waveforms sum at the eardrums, humans with normal hearing effortlessly segregate one sound source from another. Our laboratory is investigating the neural basis of this perceptual feat, often called the "cocktail party effect", using the barn owl as an animal model. The barn owl, renowned for its ability to localize sounds and its spatiotopic representation of auditory space, is an established model for spatial hearing. Here, we briefly review the neural basis of sound-localization of a single sound source in an anechoic environment and then generalize the ideas developed therein to cases in which there are multiple, concomitant sound sources and acoustical reflection.

All known human societies establish social order by punishing violators of social norms. However, little is known about how the brain processes the punishment threat associated with norm violations. We use fMRI to study the neural circuitry behind social norm compliance by comparing a treatment in which norm violations can be punished with a control treatment in which punishment is impossible. Individuals' increase in norm compliance when punishment is possible exhibits a strong positive correlation with activations in the lateral orbitofrontal cortex and right dorsolateral prefrontal cortex. Moreover, lateral orbitofrontal cortex activity is strongly correlated with Machiavellian personality characteristics. These findings indicate a neural network involved in social norm compliance that might constitute an important basis for human sociality. Different activations of this network reveal individual differences in the behavioral response to the punishment threat and might thus provide a deeper understanding of the neurobiological sources of pathologies such as antisocial personality disorder.

Reward probability has been shown to be coded by dopamine neurons in monkeys. Phasic neuronal activation not only increased linearly with reward probability upon expectation of reward, but also varied monotonically across the range of probabilities upon omission or receipt of rewards, therefore modeling discrepancies between expected and received rewards. Such a discrete coding of prediction error has been suggested to be one of the basic principles of learning. We used functional magnetic resonance imaging (fMRI) to show that the human dopamine system codes reward probability and prediction error in a similar way. We used a simple delayed incentive task with a discrete range of reward probabilities from 0%-100%. Activity in the nucleus accumbens of human subjects strongly resembled the phasic responses found in monkey neurons. First, during the expectation period of the task, the fMRI signal in the human nucleus accumbens (NAc) increased linearly with the probability of the reward. Second, during the outcome phase, activity in the NAc coded the prediction error as a linear function of reward probabilities. Third, we found that the Nac signal was correlated with individual differences in sensation seeking and novelty seeking, indicating a link between individual fMRI activation of the dopamine system in a probabilistic paradigm and personality traits previously suggested to be linked with reward processing. We therefore identify two different covariates that model activity in the Nac: specific properties of a psychological task and individual character traits.

Transcranial magnetic stimulation (TMS) is increasingly used as a research tool for functional brain mapping in cognitive neuroscience. Despite being mostly tolerable, side effects of TMS could influence task performance in behavioural TMS studies. In order to test this issue, healthy subjects assessed the discomfort caused by the stimulation during a verbal working memory task. We investigated the relation between subjective disturbance and task performance. Subjects were stimulated during the delay period of a delayed-match-to-sample task above cortical areas that had been identified before to be involved in working memory. Task performance and subjective disturbance due to side effects were monitored. The subjects' grade of discomfort correlated with the error rates: the higher the discomfort, the more errors were made. Conclusively, TMS side effects may bias task performance in cognitive neuroscience studies and may thereby lead to misinterpretation of results. We emphasize the importance of controlling side effects of the stimulation as a source of biasing effects in TMS studies.

In cognitive neuroscience, optically tracked frameless stereotaxic navigation has been successfully used to precisely guide transcranial magnetic stimulation (TMS) to desired cortical areas for brain-mapping purposes. Thereby, potential sources of imprecision are the fixation of a reference frame to the head of the subject and the referencing procedure according to certain landmarks (LM). The aim of our study was to evaluate the accuracy of frameless stereotaxic coil positioning in a standard experimental setting. A parameter for accuracy is the reproducibility of LM coordinates. In order to test the stability of the referencing for stereotaxic positioning within a single TMS session (within-session stability), the coordinates of six predefined facial LM in nine subjects were recorded first after the initial registration and second after a 20 minutes TMS session. The two sets of coordinates were then compared. The reliability of the positioning coordinates between different TMS sessions (inter-session repeatability) was addressed by registering the subjects LM coordinates in two independent TMS sessions. The variance of the recorded coordinates was analyzed. Altogether, LM were registered 1728 times (192 measures per subject). Within-session stability: The mean Euclidean distance (MED) between the LM position coordinates before and after a TMS session was 1.6 mm, when pooling over all LM. Inter-session repeatability: The MED between the LM positions recorded after the reference procedures of two different sessions showed an average deviation of 2.5 mm. In conclusion, optically tracked frameless stereotaxic coil positioning is from the technical viewpoint of high stability and repeatability. It is therefore a precise method for TMS brain mapping studies or for repeated TMS treatments, with the need of topographically exact stimulation.

We describe a principal component analysis (PCA) method for functional magnetic resonance imaging (fMRI) data based on functional data analysis, an advanced nonparametric approach. The data delivered by the fMRI scans are viewed as continuous functions of time sampled at the interscan interval and subject to observational noise, and are used accordingly to estimate an image in which smooth functions replace the voxels. The techniques of functional data analysis are used to carry out PCA directly on these functions. We show that functional PCA is more effective than is its ordinary counterpart in recovering the signal of interest, even if limited or no prior knowledge of the form of hemodynamic function or the structure of the experimental design is specified. We discuss the rationale and advantages of the proposed approach relative to other exploratory methods, such as clustering or independent component analysis, as well as the differences from methods based on expanded design matrices.

Neuroimaging studies in humans have shown that different working memory (WM) tasks recruit a common bilateral fronto-parietal cortical network. Animal studies as well as neuroimaging studies in humans have suggested that this network, in particular the prefrontal cortex, is preferentially recruited when material from different domains (e.g. spatial information or verbal/object information) has to be memorized. Early imaging studies have suggested qualitative dissociations in the prefrontal cortex for spatial and object/verbal WM, either in a left-right or a ventral-dorsal dimension. However, results from different studies are inconsistent. Moreover, recent fMRI studies have failed to find evidence for domain dependent dissociations of WM-related activity in prefrontal cortex. Here we present evidence from two independent fMRI studies using physically identical stimuli in a verbal and spatial WM task showing that domain dominance for WM does indeed exist, although only in the form of quantitative differences in activation and not in the form of a dissociation with different prefrontal regions showing mutually exclusive activation in different domains. Our results support a mixed dimension model of domain dominance for WM within the prefrontal cortex, with left ventral prefrontal cortex (PFC) supporting preferentially verbal WM and right dorsal PFC supporting preferentially spatial WM. The concept of domain dominance is discussed in the light of recent theories of prefrontal cortex function.

Objectives: Transcranial magnetic stimulation (TMS) is progressively gaining relevance as a tool in cognitive neuroscience and clinical research. However, most studies in this field do not consider individual anatomy. Neuronavigational devices allow to guide the coil to a specific cortical area, predetermined by functional magnetic resonance imaging (fMRI). Therefore, it is crucial to know whether the area of a certain function as identified by fMRI corresponds to the area where the TMS should be placed in order to influence this function., Methods: We investigated the spatial relation between the cortical area activated by a motor task in fMRI and the area of magnetically evoked motor potentials (MEP) in 8 subjects, using a spacing of 5x5 mm. A neuronavigational system was adapted for coil positioning and for the registration of the stimulation coordinates., Results: A spatial divergence of the centers of gravity from fMRI and MEP was found with a mean distance of about 10 mm, with the MEP centers being, by a mean derivation of 7.5 mm, consistently anterior to the center of fMRI activation. However, regarding MEP areas and fMRI activities, a large overlap was found for stimulation intensities of both 110 and 120% motor threshold., Conclusions: The combination of fMRI and neuronavigated TMS is useful for non-invasive investigation of individual cortical functions predetermined by fMRI. Whereas both are spatially by and large congruent, discrepencies in the exact spatial relation between MEP and fMRI areas should be considered and further studied.

Spontaneous and tone-evoked changes in light reflectance were recorded from primary auditory cortex (A1) of anesthetized cats (barbiturate induction, ketamine maintenance). Spontaneous 0.1-Hz oscillations of reflectance of 540- and 690-nm light were recorded in quiet. Stimulation with tone pips evoked localized reflectance decreases at 540 nm in 3/10 cats. The distribution of patches "activated" by tones of different frequencies reflected the known tonotopic organization of auditory cortex. Stimulus-evoked reflectance changes at 690 nm were observed in 9/10 cats but lacked stimulus-dependent topography. In two experiments, stimulus-evoked optical signals at 540 nm were compared with multiunit responses to the same stimuli recorded at multiple sites. A significant correlation (P < 0.05) between magnitude of reflectance decrease and multiunit response strength was evident in only one of five stimulus conditions in each experiment. There was no significant correlation when data were pooled across all stimulus conditions in either experiment. In one experiment, the spatial distribution of activated patches, evident in records of spontaneous activity at 540 nm, was similar to that of patches activated by tonal stimuli. These results suggest that local cerebral blood volume changes reflect the gross tonotopic organization of A1 but are not restricted to the sites of spiking neurons.

Functional magnetic resonance imaging (fMRI) was used to examine the pattern of cortical activity during a picture naming task. Subjects (n=12) had to covertly name either animals or furniture items. Functional scanning was performed using a conventional 1.5-Tesla whole-body MRI system. Images obtained during naming the two categories were compared using a non-parametric test. The study revealed evidence for domain-specific lexical regions in left middle, right middle and inferior frontal areas, as well as in superior and middle temporal areas. The results corroborate neuropsychological data and demonstrate directly and non-invasively in human volunteers that semantic representations in frontal and temporal areas are, to some degree, localized and possibly implemented as multiple maps. A completely distributed storage of semantic information is rendered unlikely., (Copyright 1998 Elsevier Science B.V.)

The cortical organization of executive control was investigated using event-related potentials (ERPs). ERPs were collected while subjects performed a go/no go task that required response inhibition. First, around 260 ms after stimulus onset, an effect of response inhibition on ERPs was observed over inferior prefrontal areas. Generators in these regions were confirmed by source analysis. Later, between 300-600 ms after stimulus onset, a left lateralized fronto-central ERP effect was found which differed in topography from a non-specific effect of task difficulty. Source analysis indicated that generators in anterior cingulate and left premotor areas also contributed to this effect. Orchestrated activation of prefrontal areas and the anterior cingulate subserves executive function whereas relatively late activity of the left premotor cortex is involved in motor control.

The term "phantom limb" denotes the sensation that an extremity is present although it has been lost. A number of clinical features and recent findings of cortical map plasticity after destruction of afferent pathways (deafferentation) suggest that phantom limbs are caused by large-scale cortical reorganization processes. However, in paraplegics, who also suffer from cortical deafferentation, phantom sensations rarely develop, and if they do, they are weak, lacking in detail, and delayed, occurring after months. This has been taken to suggest a non-cortical genesis of phantom limbs. This article proposes a biologically plausible minimal neural network model to solve this apparent puzzle. Deafferentation was simulated in trained self-organizing feature maps. Reorganization was found to be directed by input noise. According to the model, the production of input noise by the deafferented primary sensory neuron promotes cortical reorganization in amputees. No such noise is generated or conducted to the cortex in paraplegics.

Neuropsychological, computational, and psycholinguistic data suggest the existence of semantic maps, i.e. localized representations of semantic information in the brain. Using functional magnetic resonance imaging, this hypothesis was directly tested with a picture naming task involving items from four different semantic categories. Small left lateralized fronto-temporal cortical sites of category-specific activation were found when brain activation signals were averaged. Data suggest the existence of multiple maps coding high-level representations of objects, such that meaningful distinctions, at least in part, govern the physical distribution of cortical semantic storage.

Functional magnetic resonance imaging (fMRI) offers a powerful experimental tool for mapping activated cortical regions in man. Thereby, the paramagnetic deoxyhemoglobin in the red blood cells acts as an endogenous susceptibility contrast agent, which allows the noninvasive detection of stimulus-induced transient changes in regional cerebral blood flow and volume. Fifteen normal subjects were examined on a conventional 1.5-T MR system to visualize cortical activation during the performance of high-level cognitive tasks. A computer-controlled videoprojector was employed to present psychometrically optimized activation paradigms. Reaction times and error rates of the volunteers were acquired online during stimulus presentation. The time course of cortical activation was measured in a series of strongly T2*-weighted gradient-echo images from three or four adjacent slices. For anatomical correlation, picture elements showing a stimulus-related significant signal increase were color-coded and superimposed on T1-weighted spin-echo images. Analysis of the fMRI data revealed a subtle (range 2-5%), but statistically significant (P < 0.05) increase in signal intensity during the periods of induced cortical activation. Judgment of semantic relatedness of word pairs, for example, activated selectively cortical areas in left frontal and left temporal brain regions. The strength of cortex activation in the semantic task decreased significantly in the course of stimulus presentation and was paralleled by a decrease in the corresponding reaction times. With its move into the area of cognitive neuroscience, fMRI calls both for the careful design of activation schemes and for the acquisition of behavioral data. For example, brain regions involved in language processing could only be identified clearly when psychometrically matched activation paradigms were employed. The reaction time data correlated well with selective learning and thus helped to facilitate interpretation of the fMRI data sets.