Memory and Consciousness


Prof. Dr. Alan Richardson-Klavehn

Student members

Angelika Blümel
Judith Kehl


Dr. Jasmin Kizilirmak
Dr. Catherine Sweeney-Reed 
Hannes Thürich

About our group

The Memory and Consciousness Research Group, led by Professor Alan Richardson-Klavehn, conducts research on the neural basis of human learning and memory, using a wide range of cutting edge techniques to measure and visualize the activity and structure of the living brain. These include 3T and 7T magnetic resonance imaging (MRI), magnetoencepalography (MEG), electroencephalography (EEG), intracranial recordings in patients with electrode implants in the thalamus and the nucleus accumbens for epilepsy therapy, and recordings in patients with electrode grids placed on the surface of the cortex for presurgical epilepsy monitoring. The group also carries on a parallel line of purely behavioural (psychological) research. The current major methodological emphasis of the group is on examining connectivity between different brain areas during memory formation and retrieval, using functional MRI, and in real time using measures of oscillatory synchrony in electrophysiological signals.

The name of the group reflects the fact that human learning and memory is not unitary, but consists of different subtypes, which are accompanied by different kinds of consciousness during memory formation and memory retrieval, and may have different neural substrates. Thus understanding of memory, consciousness, and brain are inextricably interlinked.


Current and past research themes of the group include:

  • The relationship between priming (facilitated perceptual or conceptual processing as a result of past experience with a stimulus) and conscious recollection of past events
  • Voluntary suppression and interference control in episodic memory
  • The role of dopaminergic genes in learning and memory
  • Learning from insights during problem solving
  • The relationship between novelty and schema-consistency effects on memory formation
  • The roles of the anterior and dorsomedial thalamic nuclei in episodic memory formation and retrieval, and their relation to neocortical and medial temporal lobe function
  • The roles of hippocampal and thalamic connectivity in episodic memory formation and retrieval
  • Memory deficits in individuals with mild cognitive impairment (a possible precursor to Alzheimer's disease) and Korsakoff's syndrome (memory loss owing to B-vitamin deficiency in chronic alcoholics)

Selected publications in Cognitive Neuroscience

Additional psychological publications here

Kizilirmak, J. M., Wiegmann, B., & Richardson-Klavehn, A. (2016). Problem solving as an encoding task: A special case of the generation effect. Journal of Problem Solving, 9, 59-76. doi: 10.7771/1932-6246.1182

Soch, J., Deserno, L., Assman, A., Barman, A., Walter, H., Richardson-Klavehn, A., & Schott, B. H. (2016). Inhibition of information flow to the Default Mode Network during self-reference versus reference to others. Cerebral Cortex, epub ahead of print. doi: 10.1093/cercor/bhw206

Sweeney-Reed, C. M., Zaehle T., Voges, J., Schmitt, F. C., Buentjen, L., Kopitzki, K., Richardson-Klavehn, A., Hinrichs, H., Heinze, H. J., Knight, R. T., & Rugg, M. D. (2016). Pre-stimulus thalamic theta power predicts human memory formation. Neuroimage, epub ahead of print. doi: 10.1016/j.neuroimage.2016.05.042

Sweeney-Reed, C. M., Zaehle T., Voges, J., Schmitt, F. C., Buentjen, L., Kopitzki, K., Richardson-Klavehn, A., Hinrichs, H., Heinze, H. J., Knight, R. T., & Rugg, M. D. (2016). Clinical, neuropsychological, and pre-stimulus dorsomedial thalamic nucleus electrophysiological data in deep brain stimulation patients. Data in Brief, epub ahead of print. doi: 10.1016/j.dib.2016.06.008

Sweeney-Reed, C. M., Zaehle T., Voges, J., Schmitt, F. C., Buentjen, L., Kopitzki, K., Hinrichs, H., Heinze, H. J., Rugg, M. D., Knight, R. T., & Richardson-Klavehn, A. (2015). Thalamic theta phase alignment predicts human memory formation and anterior thalamic cross-frequency coupling. eLife, 4, e07578, doi: 10.7554/eLife.07578.

Guggenmos, M., Thoma, V., Haynes, J. D., Richardson-Klavehn, A., Cichy, R. M., & Sterzer, P. (2015). Spatial attention enhances object coding in local and distributed representations of the lateral occipital complex. NeuroImage, 116, 149-157. doi: 10.1016/j.neuroimage.2015.04.004.

Kizilirmak, J. M., Galvao Gomes da Silva, J., Imamoglu, F., & Richardson-Klavehn, A. (2015). Generation and the subjective feeling of „aha!“ are indipendently related to learning from insight. Psychological Research. doi: 10.1007/s00426-015-0697-2

Guggenmos, M., Thoma, V., Cichy, R. M., Haynes, J.-D., Sterzer, P.*, & Richardson-Klavehn, A.* (2015). Non-holistic coding of objects in lateral occipital complex with and without attention. NeuroImage, 107, 356-363. *Equal contribution. doi: 10.1016/j.neuroimage.2014.12.013

Sweeney-Reed, C. M.*, Zaehle, T., Voges, J., Schmitt, F. C., Buentjen, L., Kopitzki, K., Esslinger, C., Hinrichs, H., Heinze, H.-J., Knight, R. T., & Richardson-Klavehn, A.* (2014). Corticothalamic phase synchrony and cross-frequency coupling predict human memory formation. eLife, 3, e05352. *Corresponding authors. doi: 10.7554/eLife.05352

Kizilirmak, J. M., Rösler, F., & Khader, P. H. (2014). Trial-to-trial dynamics of selective long-term-memory retrieval with continuously changing retrieval targets. Brain and Cognition, 90, 8-18. doi: 10.1016/j.bandc.2014.04.013

Daly, I., Faller, J., Scherer, R., Sweeney-Reed, C. M., Nasuto, S. J., Billinger, M., & Müller-Putz, G. (2014). Exploration of the neural correlates of cerebral palsy for sensorimotor BCI control. Frontiers in Neuroengineering, 7, e20. doi: 10.3389/fneng.2014.00020

Bergström, Z. M., Anderson, M. C., Buda, M., Simons, J. S., & Richardson-Klavehn, A. (2013). Intentional retrieval suppression can conceal guilty knowledge in ERP memory detection tests. Biological Psychology, 94, 1-11. doi: 10.1016/j.biopsycho.2013.04.012

Schott, B. H.*, Wüstenberg, T., Wimber, M., Fenker, D. B., Zierhut, K. C., Seidenbecher, C. I., Heinze, H.-J., Walter, H., Düzel, E., & Richardson-Klavehn, A.* (2013). The relationship between level of processing and hippocampal-cortical functional connectivity during episodic memory formation in humans. Human Brain Mapping, 34, 407-424. *Corresponding authors. doi: 10.1002/hbm.21435

Richardson-Klavehn, A., & Badenschier, F. (2013). Schon wieder dieses Wort - wie kommt das? (This word already again: How does that happen?). Answer to reader enquiry concerning psychological and brain processes of perceptual priming and conscious recollection in the German web portal (

Staudigl, T., Zaehle, T., Voges, J., Hanslmayr, S., Esslinger, C., Hinrichs, H., Schmitt, F. C., Heinze, H.-J., & Richardson-Klavehn, A. (2012). Memory signals from the thalamus: Early thalamocortical phase synchronization entrains gamma oscillations during long-term memory retrieval. Neuropsychologia, 50, 3519-3527. doi: 10.1016/j.neuropsychologia.2012.08.023

Wimber, M., Maaß, A., Staudigl, T., Richardson-Klavehn, A., & Hanslmayr, S. (2012). Rapid memory reactivation revealed by oscillatory entrainment. Current Biology, 22, 1482-1486. doi: 10.1016/j.cub.2012.05.054

Kizilirmak, J. M., Rösler, F., & Khader, P. H. (2012). Control processes during selective long-term memory retrieval. NeuroImage, 59, 1830-1841. doi: 10.1016/j.neuroimage.2011.08.041

Sweeney-Reed, C. M., Riddell, P. M., Ellis, J. A., Freeman, J. E., & Nasuto, S. J. (2012). Neural correlates of true and false memory in mild cognitive impairment. PLoS One, 7, e48357. doi: 10.1371/journal.pone.0048357

Daly, I., Sweeney-Reed, C. M., & Nasuto, S. J. (2012). Testing for significance of phase synchronisation dynamics in the EEG. Journal of Computational Neuroscience, 34, 411-432. doi: 10.1007/s10827-012-0428-2

Wimber, M., Schott, B. H., Wendler, F., Seidenbecher, C. I., Behnisch, G., Macharadze, T., Bäuml, K.-H., & Richardson-Klavehn, A. (2011). Prefrontal dopamine and the dynamic control of human long-term memory. Translational Psychiatry, 1, e15. doi: 10.1038/tp.2011.15

Schott, B. H., Seidenbecher, C. I., Richter, S., Wüstenberg, T., Debska-Vielhaber, G., Schubert, H., Heinze, H.-J., Richardson-Klavehn, A., & Düzel, E. (2011). Genetic variation of the serotonin 2a receptor affects hippocampal novelty processing in humans. PLoS One, 6, e15984. doi: 10.1371/journal.pone.0015984

Zweynert, S., Pade, J. P., Wüstenberg, T., Sterzer, P., Walter, H., Seidenbecher, C. I., Richardson-Klavehn, A., Düzel, E., & Schott, B. H. (2011). Motivational salience modulates hippocampal repetition suppression and functional connectivity in humans. Frontiers in Human Neuroscience, 5, e144. doi: 10.3389/fnhum.2011.00144

Richardson-Klavehn, A., & Schott, B. H. (2011). Homo sapiens: An animal made of memory. In K. Hermann (Ed.), Here and now: Amnesia (pp. 52-57). Berlin, Germany: Savvy Contemporary Gallery.

Wimber, M., Heinze, H.-J., & Richardson-Klavehn, A. (2010). Distinct fronto-parietal networks set the stage for later perceptual identification priming and episodic recognition memory. Journal of Neuroscience, 30, 13272-13280. doi: 10.1523/JNEUROSCI.0588-10.2010 

Richardson-Klavehn, A. (2010). Priming, automatic recollection, and control of retrieval: Toward an integrative retrieval architecture. In J. H. Mace (Ed.), The act of remembering: Toward an understanding of how we recall the past (pp. 111-179). Oxford, UK: Wiley-Blackwell. doi: 10.1002/9781444328202

Richardson-Klavehn, A. (2010). On the fruitful relationship between functional neuroimaging and cognitive theories of human learning and memory. In A. S. Benjamin (Ed.), Successful remembering and successful forgetting: A Festschrift in honor of Robert A. Bjork (pp. 477- 503). New York: Psychology Press. doi: 10.4324/9780203842539

Richardson-Klavehn, A. (2010). Prestimulus neural oscillations and their haemodynamic correlates elucidate the cognitive and neural processes of memory formation. Clinical Neurophysiology, 41, 137-146. doi: 10.1055/s-0030-1252031

Kopp, B., Kizilirmak, J., Liebscher, C., Runge, J., & Wessel, K. (2010). Event-related brain potentials and the efficiency of visual search for vertically and horizontally oriented stimuli. Cognitive, Affective, & Behavioral Neuroscience, 10, 523-540. doi: 10.3758/CABN.10.4.523

Bergström, Z. M., de Fockert, J. W., & Richardson-Klavehn, A. (2009). ERP and behavioural evidence for direct suppression of unwanted memories. NeuroImage, 48, 726-737. doi: 10.1016/j.neuroimage.2009.06.051

Bergström, Z. M., de Fockert, J. W., & Richardson-Klavehn, A. (2009). Event related potential evidence that automatic recollection can be voluntarily avoided. Journal of Cognitive Neuroscience, 21, 1280-1301. doi: 10.1162/jocn.2009.21075

Guderian, S., Schott, B. H., Richardson-Klavehn, A., & Düzel, E. (2009). Medial temporal theta state before an event predicts episodic encoding success in humans. Proceedings of the National Academy of Sciences USA, 106, 5365-5370. doi: 10.1073/pnas.0900289106

Richardson-Klavehn, A., Bergström, Z. M., Magno, E., Markopoulos, G., Sweeney-Reed, C. M., & Wimber, M. (2009). On the intimate relationship between neurobiology and function in the theoretical analysis of human learning and memory. In F. Rösler, C. Ranganath, B. Röder, & R. H. Kluwe (Eds.), Neuroimaging in human memory: Linking cognitive processes to neural systems (pp. 127-165). New York: Oxford University Press. doi: 10.1093/acprof:oso/9780199217298.001.0001

Sweeney-Reed, C. M., & Nasuto, S. J. (2009). Detection of neural correlates of self-paced motor activity using empirical mode decomposition phase locking analysis. Journal of Neuroscience Methods, 184, 54-70. doi: 10.1016/j.jneumeth.2009.07.023

Wimber, M., Bäuml, K.-H., Bergström, Z. M., Markopoulos, G., Heinze, H.-J., & Richardson-Klavehn, A. (2008). Neural markers of inhibition in human memory retrieval. Journal of Neuroscience, 28, 13419-13427. doi: 10.1523/JNEUROSCI.1916-08.2008

Bergström, Z. M., Velmans, M., de Fockert, J. W., & Richardson-Klavehn, A. (2007). ERP evidence for successful voluntary control of conscious recollection. Brain Research, 1151, 119-133. doi: 10.1016/j.brainres.2007.03.014

Ramponi, C., Richardson-Klavehn, A., & Gardiner, J. M. (2007). Component processes of conceptual priming and associative cued recall: The roles of preexisting representation and depth of processing. Journal of Experimental Psychology: Learning, Memory, and Cognition, 33, 843-862. doi: 10.1037/0278-7393.33.5.843

Sweeney-Reed, C. M., & Nasuto, S. J. (2007). A novel approach to the detection of synchronisation in EEG based on empirical mode decomposition. Journal of Computational Neuroscience, 23, 79-111. doi: 10.1007/s10827-007-0020-3

Schott, B. H.*, Richardson-Klavehn, A.*, Henson, R. N. A., Becker, C., Heinze, H.-J., & Düzel, E. (2006). Neuroanatomical dissociation of encoding processes related to priming and explicit memory. Journal of Neuroscience, 26, 792-800. *Equal contribution. doi: 10.1523/JNEUROSCI.2402-05.2006

Andrade, A. O., Nasuto, S. J., Kyberd, P., Sweeney-Reed, C. M., & van Kanijn, F. R. (2006). EMG signal filtering based on empirical mode decomposition. Biomedical Signal Processing and Control, 1, 44-55. doi: 10.1016/j.bspc.2006.03.003

Schott, B. H.*, Henson, R. N. A.*, Richardson-Klavehn, A.*, Becker, C., Thoma, V., Heinze, H.-J., & Düzel, E. (2005). Redefining implicit and explicit memory: The functional neuroanatomy of priming, remembering, and control of retrieval. Proceedings of the National Academy of Sciences USA, 102, 1257-1262. *Equal contribution. doi: 10.1073/pnas.0409070102

Düzel, E., Richardson-Klavehn, A., Neufang, M., Schott, B. H., Scholz, M., & Heinze, H.-J. (2005). Early, partly anticipatory, neural oscillations during identification set the stage for priming. NeuroImage, 25, 690-700. doi: 10.1016/j.neuroimage.2004.11.038

Fenker, D. B., Schott, B. H., Richardson-Klavehn, A., Heinze, H.-J., & Düzel, E. (2005). Recapitulating emotional context: Activity of amygdala, hippocampus and fusiform cortex during recollection and familiarity. European Journal of Neuroscience, 21, 1993-1999. doi: 10.1111/j.1460-9568.2005.04033.x

Andrade, A. O., Nasuto, S. J., Kyberd, P., & Sweeney-Reed, C. M. (2005). Generative topographic mapping applied to clustering and visualization of motor unit action potentials. Biosystems, 82, 273-284. doi: 10.1016/j.biosystems.2005.09.004

Ramponi, C., Richardson-Klavehn, A., & Gardiner, J. M. (2004). Level of processing and age affect involuntary conceptual priming of weak but not strong associates. Experimental Psychology, 51, 159-164. doi: 10.1027/1618-3169.51.3.159

Schott, B. H., Richardson-Klavehn, A., Heinze, H.-J., & Düzel, E. (2002). Perceptual priming versus explicit memory: Dissociable neural correlates at encoding. Journal of Cognitive Neuroscience, 14, 578-592. doi: 10.1162/08989290260045828

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