Papers by Roman Freunberger
NeuroImage, 2012
Efficient encoding of relevant information and suppression of irrelevant information influence wo... more Efficient encoding of relevant information and suppression of irrelevant information influence working memory (WM) performance, which is limited and declines in adulthood. A cued Sternberg WM task and electroencephalographic recordings (EEG) were used to investigate encoding and control operations in response to to-be-remembered (REM) and not-to-be-remembered (NREM) stimuli in younger and older adults. Younger and older adults selectively remembered REM items in a final recognition memory test. During early stages of stimulus processing, inter-trial phase stability was higher for REM than for NREM items in younger and older adults, presumably reflecting preferential encoding of REM items. At later stages, the oscillatory power of oscillations in the alpha/beta frequency range was higher for NREM than for REM, presumably reflecting the inhibitory top-down suppression of task-irrelevant information. Early phase stability was selectively related to working memory performance in younger adults and high-functioning older adults. The results of this study reveal the differential contributions of low-level feature binding and strategic control components to adult age differences in WM, and show that older adults with more youth-like processing dynamics tend to achieve higher levels of performance.
Klinische Neurophysiologie, 2007
Neuroscience Letters, 2005
This study investigated the question whether good and bad performance in a visual discrimination ... more This study investigated the question whether good and bad performance in a visual discrimination task is related to resting alpha power in a different way as it is known from memory tasks. The results show that good perceptual but not memory performance is related to low alpha amplitudes. In addition, we found that large phase resetting in the alpha band, and enhanced early components in the ERP are related to good performance in the discrimination task. The conclusion of this study is that in contrast to memory performance which is related to large resting alpha activity low alpha amplitudes are an indicator for good perceptual performance.
Neuroscience Letters, 2007
In this study we investigated the hypothesis whether P2-related differences tested in a visual pr... more In this study we investigated the hypothesis whether P2-related differences tested in a visual priming paradigm are associated with theta phaselocking. We recorded the EEG from 31 electrodes and calculated phase-locking index and total power differences for frequencies between 2 and 20 Hz. ERPs (event-related potentials) were analyzed for P1, N1 and P2 components. P2 showed strongest task-related amplitude differences between congruent and incongruent targets. A source analyses was performed for the P2 component using sLoreta that revealed local generators of the P2 in parieto-occipital regions. Phase-locking analyses showed specific effects in the theta range (4-6 Hz) appearing in time windows at around the P2 component. We draw the conclusion that phase-locked theta reflect top-down regulation processes mediating information between memory systems and is in part involved in the modulation of the P2 component.
Neuroscience & Biobehavioral Reviews, 2007
The traditional view holds that event-related potentials (ERPs) reflect fixed latency, fixed pola... more The traditional view holds that event-related potentials (ERPs) reflect fixed latency, fixed polarity evoked responses that appear superimposed on the 'background EEG'. The validity of the evoked model has been questioned by studies arguing that ERPs are generated at least in part by a reset of ongoing oscillations. But a proof of phase reset that is distinct from the 'artificial' influence of evoked components on EEG phase-has been proven difficult for a variety of methodological reasons. We argue that a theoretical analysis of the assumptions and empirical evaluation of predictions of the evoked and oscillatory ERP model offer a promising way to shed new light on mechanisms generating ERPs that goes well beyond attempts to prove phase reset. Research on EEG oscillations documents that oscillations are task relevant and show a common operating principle, which is the control of the timing of neural activity. Both findings suggest that phase reorganization of task relevant oscillations is a theoretical necessity. We further argue and show evidence that (i) task relevant oscillations exhibit a typical interactive and task relevant relationship between pre- and poststimulus power in the theta and alpha frequency range in a way that small prestimulus power is related to large poststimulus power and vice versa, (ii) ERP (interpeak) latencies and (iii) ERP amplitudes reflect frequency characteristics of alpha and theta oscillations. We emphasize that central assumptions of the evoked model cannot be substantiated and conclude that the ERPR model offers a new way for an integrative interpretation of ongoing and event-related EEG phenomena.
Neuroscience & Biobehavioral Reviews, 2010
A central topic in cognitive neuroscience is the question, which processes underlie large scale c... more A central topic in cognitive neuroscience is the question, which processes underlie large scale communication within and between different neural networks. The basic assumption is that oscillatory phase synchronization plays an important role for process binding--the transient linking of different cognitive processes--which may be considered a special type of large scale communication. We investigate this question for memory processes on the basis of different types of oscillatory synchronization mechanisms. The reviewed findings suggest that theta and alpha phase coupling (and phase reorganization) reflect control processes in two large memory systems, a working memory and a complex knowledge system that comprises semantic long-term memory. It is suggested that alpha phase synchronization may be interpreted in terms of processes that coordinate top-down control (a process guided by expectancy to focus on relevant search areas) and access to memory traces (a process leading to the activation of a memory trace). An analogous interpretation is suggested for theta oscillations and the controlled access to episodic memories.
Neuroscience & Biobehavioral Reviews, 2010
Neural correlates of control mechanisms in human working memory are discussed at two levels in th... more Neural correlates of control mechanisms in human working memory are discussed at two levels in this review: (i) at 'item level', where in multi-item working memory information needs to be organized into sequential memory representations, and (ii) at a 'process level', indicating the integration and control of a variety of cognitive functions involved in working memory, independent of item representations per se. It will be discussed that at both levels electroencephalographic theta activity is responsible for control of working memory functions. On item level, exact phase coding, e.g., approached by coupling between theta and gamma oscillations or phase resetting of theta frequency, is suggested to integrate information into working memory representations. At process level interregional theta synchronization is discussed to integrate brain structures necessary for working memory. When discussing the specificity of theta activity for control of working memory processes it will be suggested that theta oscillations might play an important general integrative role in organization of brain activity. And as working memory often involves a variety of cognitive processes which need to be coordinated there is particular need for an integrative brain mechanism like theta activity as suggested in this review.
Neuroscience, 2007
The event-related potential (ERP) is one of the most popular measures in human cognitive neurosci... more The event-related potential (ERP) is one of the most popular measures in human cognitive neuroscience. During the last few years there has been a debate about the neural fundamentals of ERPs. Two models have been proposed: The evoked model states that additive evoked responses which are completely independent of ongoing background electroencephalogram generate the ERP. On the other hand the phase reset model suggests a resetting of ongoing brain oscillations to be the neural generator of ERPs. Here, evidence for either of the two models is presented and validated, and their possible impact on cognitive neuroscience is discussed. In addition, future prospects on this field of research are presented.
NeuroImage, 2012
In the present study we have tested the hypothesis that evoked traveling alpha waves are behavior... more In the present study we have tested the hypothesis that evoked traveling alpha waves are behaviorally significant. The results of a visual-semantic categorization task show that three early ERP components including the P1-N1 complex had a dominant frequency characteristic in the alpha range and behaved like traveling waves do. They exhibited a traveling direction from midline occipital to right lateral parietal sites. Phase analyses revealed that this traveling behavior of ERP components could be explained by phase-delays in the alpha but not theta and beta frequency range. Most importantly, we found that the speed of the traveling alpha wave was significantly and negatively correlated with reaction time indicating that slow traveling speed was associated with fast picture-categorization. We conclude that evoked alpha oscillations are functionally associated with early access to visual-semantic information and generateor at least modulatethe early waveforms of the visual ERP.
NeuroImage, 2008
In the present study, we investigate the role of upper alpha oscillations for semantic access and... more In the present study, we investigate the role of upper alpha oscillations for semantic access and retrieval processes. In each of a series of trials, subjects were presented trains of distorted pictures (with decreasing levels of degradation), and were asked to respond as quickly as possible when they recognize the meaning of the picture. The results show that during the time-window of picture recognition, upper alpha power decreased but inter-areal phase synchronization increased as compared to meaningless control pictures. We assume that synchronous alpha oscillations--particularly during a decrease in alpha power--reflect topographically specific neural network activity that is related to the access of semantic information in LTM.
Human Brain Mapping, 2009
The functions of human alpha oscillations ($10 Hz) were related to cognitive processes such as me... more The functions of human alpha oscillations ($10 Hz) were related to cognitive processes such as memory and top-down control. Recent models suggest that alpha phase serves as a mechanism especially relevant for the timing of neural activity, whereas alpha amplitude is important for the inhibition of task-irrelevant brain areas. This study investigates directly the influence of top-down modulation on phase-locked and nonphase-locked alpha rhythms. We conducted an EEG experiment where subjects performed a working memory task. In the encoding phase of the task subjects had to learn presented pictures of nonliving objects that could later be asked to be retrieved. We varied the top-down modulation by including cues indicating either to remember or to forget (not to remember) the next following item. Spectral analyses showed that nonremember cues elicited pronounced alpha amplitude increase compared to remember cues. Furthermore, phase-locking in low frequencies, especially in the alpha range (7-12 Hz), was stronger for remember as opposed to not-to-remember items. In conclusion, we propose that alpha amplitude reflects top-down modulated inhibition and that alpha phase is important for the exact timing of neural activity and can be related to binding processes. Hum Brain Mapp 30:3417-3425,
Human Brain Mapping, 2005
Electroencephalogram (EEG) ␣ (ϳ10 Hz) is the dominant rhythm in the human brain during conditions... more Electroencephalogram (EEG) ␣ (ϳ10 Hz) is the dominant rhythm in the human brain during conditions of mental inactivity. High amplitudes as observed during rest usually diminish during cognitive effort. During retention of information in working memory, however, power increase of ␣ oscillations can be observed. This ␣ synchronization has been interpreted as cortical idling or active inhibition. The present study provides evidence that during top-down processing in a working memory task, ␣ power increases at prefrontal but decreases at occipital electrode sites, thereby reaching a state in which ␣ power and frequency become very similar over large distances. Two experimental conditions were compared. In the first, visuospatial information only had to be retained in memory whereas the second condition additionally demanded manipulation of the information. During the second condition, stronger ␣ synchronization at prefrontal sites and larger occipital ␣ suppression was observed as compared to that for pure retention. This effect was accompanied by assimilation of prefrontal and occipital ␣ frequency, stronger functional coupling between prefrontal and occipital brain areas, and ␣ latency shifts from prefrontal cortex to primary visual areas, possibly indicating the control of posterior cortical activation by anterior brain areas. An increase of prefrontal EEG ␣ amplitudes, which is accompanied by a decrease at posterior sites, thus may not be interpreted in terms of idling or "global" inhibition but may enable a tight functional coupling between prefrontal cortical areas, and thereby allows the control of the execution of processes in primary visual brain regions. Hum Brain Mapp 26:148 -155, 2005.
Frontiers in Psychology, 2011
For a long time alpha oscillations have been functionally linked to the processing of visual info... more For a long time alpha oscillations have been functionally linked to the processing of visual information. Here we propose an new theory about the functional meaning of alpha. The central idea is that synchronized alpha reflects a basic processing mode that controls access to information stored in a complex long-term memory system, which we term knowledge system in order to emphasize that it comprises not only declarative memories but any kind of knowledge comprising also procedural information. Based on this theoretical background, we assume that during early stages of perception, alpha "directs the flow of information" to those neural structures which represent information that is relevant for encoding. The physiological function of alpha is interpreted in terms of inhibition. We assume that alpha enables access to stored information by inhibiting task-irrelevant neuronal structures and by timing cortical activity in task relevant neuronal structures. We discuss a variety findings showing that evoked alpha and phase locking reflect successful encoding of global stimulus features in an early post-stimulus interval of about 0-150 ms.
Frontiers in Psychology, 2011
Visual attention can be shifted in space without moving the eyes. Amplitude decrease of rhythmica... more Visual attention can be shifted in space without moving the eyes. Amplitude decrease of rhythmical brain activity around 10 Hz (so called alpha activity) at contralateral posterior sites has been reported during covered shifts of visuospatial attention to one visual hemi-field. Alpha amplitude increase, on the other hand, can be found at ipsilateral visual cortex. There is some evidence suggesting an involvement of prefrontal brain areas during the control of attention-related anticipatory alpha amplitude asymmetry. This open question has been studied in detail using a multimodal approach combining transcranial magnetic stimulation (TMS) and multichannel electroencephalography (EEG) in healthy humans. Slow (1 Hz) repetitive TMS leading to reduced excitability of the stimulation site was delivered either to right frontal eye field (FEF) or a control site (vertex). Subsequently, participants had to perform a spatial cuing task in which covert shifts of attention were required to either the left or the right visual hemi-field. After stimulation at the vertex (control condition) a pattern of anticipatory, attention-related ipsilateral alpha increase/contralateral alpha decrease over posterior recording sites could be obtained. Additionally, there was pronounced coupling between (in particular right) FEF and posterior brain sites at EEG alpha frequency. When, however, right prefrontal cortex had been virtually lesioned preceding the task, these EEG correlates of visuospatial attention were attenuated. Notably, the effect of TMS at the right FEF on interregional fronto-parietal alpha coupling predicted the effect of TMS on response times. This suggests that visual attention processes associated with posterior EEG alpha activity are at least partly top-down controlled by the prefrontal cortex.
Experimental Brain Research, 2006
In a task switching design, we investigated the question whether long-range theta coupling primar... more In a task switching design, we investigated the question whether long-range theta coupling primarily reflects top-down control processes. Switch and stay trials did not differ with respect to memory load or global working memory (WM) demands. The results revealed significantly stronger theta coupling (in a range of 4-7 Hz) between prefrontal and posterior regions during switch as compared to stay trials. Power differences, reflecting more local effects, were largest in the upper alpha band (10-13 Hz) and over posterior brain areas, possibly reflecting long-term memory activation. The conclusion of the present study is that long-range coherent oscillatory activity in the theta band reflects top-down activation rather than global WM functions.
European Journal of Neuroscience, 2008
The present study attempts to demonstrate functional similarities between the P1 component of eve... more The present study attempts to demonstrate functional similarities between the P1 component of event-related potentials and alpha oscillations that are predicted by the 'alpha inhibition-timing' hypothesis. On the basis of findings showing that the frequency characteristic of the P1 component lies in the alpha range and that alpha oscillation is functionally associated with inhibition, we predict that the P1 component also reflects inhibitory processes. This hypothesis is tested in two experiments, a spatial-cuing task and a visual-semantic categorization task. The results of the cuing task demonstrate that in a similar way as alpha power, the P1 component is larger over task-irrelevant ipsilateral sites. For the categorization task, we found that the P1 component, in a similar way to alpha oscillations, is larger for task-irrelevant, distorted pictures. We conclude that the P1 component may be generated at least in part by evoked alpha oscillations and reflects inhibition in the sense of suppressing task-irrelevant processes.
Cerebral Cortex, 2006
An unresolved question in electroencephalogram (EEG) research is whether event-related potentials... more An unresolved question in electroencephalogram (EEG) research is whether event-related potentials (ERPs) are generated by phase-reset or evoked response. We analyzed data of a visual feature detection task and will show 1) phase concentration in the alpha frequency range, 2) ongoing alpha activity prior to stimulus onset, 3) evoked alpha oscillation in the ERP, 4) lack of power increase during phase concentration, 5) decrease in amplitude variance during early evoked components preceding a decrease in power, and 6) the same cortical sources for induced prestimulus power and evoked poststimulus power. Because none of these data provide unequivocal evidence for phase reset, we additionally tested the basic assumption of the evoked model, which is the additivity of the evoked response on the basis of a simulation approach. Our findings suggest that nonadditive processes-typical for a phase reset-are involved in the generation of the ERP. Thus, together with the other findings this study provides unequivocal evidence for phase resetting in the human EEG.
Brain Research Bulletin, 2007
We tested the hypothesis whether images of real objects elicit stronger gamma (&a... more We tested the hypothesis whether images of real objects elicit stronger gamma (>25 Hz) synchronization, when compared with scrambled objects. The background of this study is a recent debate about the functional meaning of evoked and induced gamma oscillations. Brain electrical source analysis (BESA) and low resolution electromagnetic tomography analysis (LORETA) was performed on the basis of the event-related potential (ERP) data. A component at around 230 ms (termed C230) showed strongest differences between objects and scrambled objects. Time-frequency analyses were run across electrodes and within the dipole sources. We found increased gamma event-related synchronization (ERS) between 200 and 300 ms for real objects. This effect was strongest in a fronto-medial source. Induced gamma, as also shown in previous studies, reflects the more task-relevant mechanism where object representations become activated.
Brain Research Bulletin, 2011
► Pre-stimulus alpha phase is not randomly distributed in time across trials. ► Ongoing oscillato... more ► Pre-stimulus alpha phase is not randomly distributed in time across trials. ► Ongoing oscillatory activity is not random noise. ► Pre-stimulus oscillatory activity contributes to the appearance of event-related potentials.
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Papers by Roman Freunberger