Notger G. Müller

Object Familiarity and Semantic Congruency Modulate Responses in Cortical Audiovisual Integration Areas

The cortical integration of auditory and visual features is crucial for efficient object recognition. Previous studies have shown that audiovisual (AV) integration is affected by where and when auditory and visual features occur. However, because relatively little is known about the impact of what is integrated, we here investigated the impact of semantic congruency and object familiarity on the neural correlates of AV integration. We used functional magnetic resonance imaging to identify regions involved in the integration of both (congruent and incongruent) familiar animal sounds and images and of arbitrary combinations of unfamiliar artificial sounds and object images. Unfamiliar object images and sounds were integrated in the inferior frontal cortex (IFC), possibly reflecting learning of novel AV associations. Integration of familiar, but semantically incongruent combinations also correlated with IFC activation and additionally involved the posterior superior temporal sulcus (pSTS). For highly familiar semantically congruent AV pairings, we again found AV integration effects in pSTS and additionally in superior temporal gyrus. These findings demonstrate that the neural correlates of object-related AV integration reflect both semantic congruency and familiarity of the integrated sounds and images.

Expectations Change the Signatures and Timing of Electrophysiological Correlates of Perceptual Awareness

Previous experience allows the brain to predict what comes next. How these expectations affect conscious experience is poorly understood. In particular, it is unknown whether and when expectations interact with sensory evidence in granting access to conscious perception, and how this is reflected electrophysiologically. Here, we parametrically manipulate sensory evidence and expectations while measuring event-related potentials in human subjects to assess the time course of evoked responses that correlate with subjective visibility, the properties of the stimuli, and/or perceptual expectations. We found that expectations lower the threshold of conscious perception and reduce the latency of neuronal signatures differentiating seen and unseen stimuli. Without expectations, this differentiation occurs ∼300 ms and with expectations ∼200 ms after stimulus in occipitoparietal sensors. The amplitude of this differentiating response component (P2) decreases as visibility increases, regardless of whether this increase is attributable to enhanced sensory evidence and/or the gradual buildup of perceptual expectations. Importantly, at matched performance levels, responses to seen and unseen stimuli differed regardless of the physical stimulus properties. These findings indicate that the latency of the neuronal correlates of access to consciousness depend on whether access is driven by stimulus saliency or by a combination of expectations and sensory evidence.

The attentional 'spotlight's' penumbra: center-surround modulation in striate cortex.

By enhancing neural activity in respective retinotopic cortical representations attention increases the efficiency with which visual information at a selected location is processed. Behavioral data also suggest that information from the vicinity of the attended region is actively suppressed. In search for a physiological correlate of this ‘spotlights penumbra’ we assessed neural responses in retinotopic representations of an attended location and of locations at different distances to it. Relative to passive viewing we found suppressed striate activity for the nearby but not for the far locations. This attention-driven center-surround distribution of neural activity may enhance the contrast between attended and non-attended objects. We relate the different behavior of extrastriate areas to their lower spatial resolution, i.e. larger receptive fields.

Keeping Memory Clear and Stable—The Contribution of Human Basal Ganglia and Prefrontal Cortex to Working Memory

Successful remembering involves both hindering irrelevant information from entering working memory (WM) and actively maintaining relevant information online. Using a voxelwise lesion–behavior brain mapping approach in stroke patients, we observed that lesions of the left basal ganglia render WM susceptible to irrelevant information. Lesions of the right prefrontal cortex on the other hand make it difficult to keep more than a few items in WM. These findings support basal ganglia–prefrontal cortex models of WM whereby the basal ganglia play a gatekeeper role and allow only relevant information to enter prefrontal cortex where this information then is actively maintained in WM.

Vascular hippocampal plasticity after aerobic exercise in older adults

Aerobic exercise in young adults can induce vascular plasticity in the hippocampus, a critical region for recall and recognition memory. In a mechanistic proof-of-concept intervention over 3 months, we investigated whether healthy older adults (60–77 years) also show such plasticity. Regional cerebral blood flow (rCBF) and volume (rCBV) were measured with gadolinium-based perfusion imaging (3 Tesla magnetic resonance image (MRI)). Hippocampal volumes were assessed by high-resolution 7 Tesla MRI. Fitness improvement correlated with changes in hippocampal perfusion and hippocampal head volume. Perfusion tended to increase in younger, but to decrease in older individuals. The changes in fitness, hippocampal perfusion and volume were positively related to changes in recognition memory and early recall for complex spatial objects. Path analyses indicated that fitness-related changes in complex object recognition were modulated by hippocampal perfusion. These findings indicate a preserved capacity of the aging human hippocampus for functionally relevant vascular plasticity, which decreases with progressing age.

Cross-modal processing in early visual and auditory cortices depends on expected statistical relationship of multisensory information.

Previous studies have shown that processing information in one sensory modality can either be enhanced or attenuated by concurrent stimulation of another modality. Here, we reconcile these apparently contradictory results by showing that the sign of cross-modal interactions depends on whether the content of two modalities is associated or not. When concurrently presented auditory and visual stimuli are paired by chance, cue-induced preparatory neural activity is strongly enhanced in the task-relevant sensory system and suppressed in the irrelevant system. Conversely, when information in the two modalities is reliably associated, activity is enhanced in both systems regardless of which modality is task relevant. Our findings illustrate an ecologically optimal flexibility of the neural mechanisms that govern multisensory processing: facilitation occurs when integration is expected, and suppression occurs when distraction is expected. Because thalamic structures were more active when the senses needed to operate separately, we propose them to serve gatekeeper functions in early cross-modal interactions.

Cortical Plasticity of Audio–Visual Object Representations

Several regions in human temporal and frontal cortex are known to integrate visual and auditory object features. The processing of audio–visual (AV) associations in these regions has been found to be modulated by object familiarity. The aim of the present study was to explore training-induced plasticity in human cortical AV integration. We used functional magnetic resonance imaging to analyze the neural correlates of AV integration for unfamiliar artificial object sounds and images in naïve subjects (PRE training) and after a behavioral training session in which subjects acquired associations between some of these sounds and images (POST-training). In the PRE-training session, unfamiliar artificial object sounds and images were mainly integrated in right inferior frontal cortex (IFC). The POST-training results showed extended integration-related IFC activations bilaterally, and a recruitment of additional regions in bilateral superior temporal gyrus/sulcus and intraparietal sulcus. Furthermore, training-induced differential response patterns to mismatching compared with matching (i.e., associated) artificial AV stimuli were most pronounced in left IFC. These effects were accompanied by complementary training-induced congruency effects in right posterior middle temporal gyrus and fusiform gyrus. Together, these findings demonstrate that short-term cross-modal association learning was sufficient to induce plastic changes of both AV integration of object stimuli and mechanisms of AV congruency processing.

Relationships of peripheral IGF-1, VEGF and BDNF levels to exercise-related changes in memory, hippocampal perfusion and volumes in older adults

Animal models point towards a key role of brain-derived neurotrophic factor (BDNF), insulin-like growth factor-I (IGF-I) and vascular endothelial growth factor (VEGF) in mediating exercise-induced structural and functional changes in the hippocampus. Recently, also platelet derived growth factor-C (PDGF-C) has been shown to promote blood vessel growth and neuronal survival. Moreover, reductions of these neurotrophic and angiogenic factors in old age have been related to hippocampal atrophy, decreased vascularization and cognitive decline. In a 3-month aerobic exercise study, forty healthy older humans (60 to 77years) were pseudo-randomly assigned to either an aerobic exercise group (indoor treadmill, n=21) or to a control group (indoor progressive-muscle relaxation/stretching, n=19). As reported recently, we found evidence for fitness-related perfusion changes of the aged human hippocampus that were closely linked to changes in episodic memory function. Here, we test whether peripheral levels of BDNF, IGF-I, VEGF or PDGF-C are related to changes in hippocampal blood flow, volume and memory performance. Growth factor levels were not significantly affected by exercise, and their changes were not related to changes in fitness or perfusion. However, changes in IGF-I levels were positively correlated with hippocampal volume changes (derived by manual volumetry and voxel-based morphometry) and late verbal recall performance, a relationship that seemed to be independent of fitness, perfusion or their changes over time. These preliminary findings link IGF-I levels to hippocampal volume changes and putatively hippocampus-dependent memory changes that seem to occur over time independently of exercise. We discuss methodological shortcomings of our study and potential differences in the temporal dynamics of how IGF-1, VEGF and BDNF may be affected by exercise and to what extent these differences may have led to the negative findings reported here.

Interaction between Bottom-up Saliency and Top-down Control: How Saliency Maps Are Created in the Human Brain

Whether an object captures our attention depends on its bottom-up salience, that is, how different it is compared with its neighbors, and top-down control, that is, our current inner goals. At which neuronal stage they interact to guide behavior is still unknown. In a functional magnetic resonance imaging study, we found evidence for a hierarchy of saliency maps in human early visual cortex (V1 to hV4) and identified where bottom-up saliency interacts with top-down control: V1 represented pure bottom-up signals, V2 was only responsive to top-down modulations, and in hV4 bottom-up saliency and top-down control converged. Two distinct cerebral networks exerted top-down control: distractor suppression engaged the left intraparietal sulcus, while target enhancement involved the frontal eye field and lateral occipital cortex. Hence, attentional selection is implemented in integrated maps in visual cortex, which provide precise topographic information about target-distractor locations thus allowing for successful visual search.

Repetition Suppression versus Enhancement—It's Quantity That Matters

Upon repetition, certain stimuli induce reduced neural responses (i.e., repetition suppression), whereas others evoke stronger signals (i.e., repetition enhancement). It has been hypothesized that stimulus properties (e.g., visibility) determine the direction of the repetition effect. Here, we show that the very same stimuli can induce both repetition suppression and enhancement, whereby the only determining factor is the number of repetitions. Repeating the same, initially novel low-visible pictures of scenes for up to 5 times enhanced the blood oxygen level-dependent (BOLD) response in scene-selective areas, that is, the parahippocampal place area (PPA) and the transverse occipital sulcus (TOS), presumably reflecting the strengthening of the internal representation. Additional repetitions (6-9) resulted in progressively attenuated neural responses indicating a more efficient representation of the now familiar stimulus. Behaviorally, repetition led to increasingly faster responses and higher visibility ratings. Novel scenes induced the largest BOLD response in the PPA and also higher activity in yet another scene-selective region, the retrospenial cortex (RSC). We propose that 2 separable processes modulate activity in the PPA: one process optimizes the internal stimulus representation and involves TOS and the other differentiates between familiar and novel scenes and involves RSC.