Peer-reviewed Journal Arcticles


Smaczny, S., Klein, E., Jung, S., Moeller, K., & Karnath, H. O. (2024). The line bisection bias as a deficit of proportional reasoning− evidence from number line estimation in neglect. Neuropsychologia, 108848.


Klein, E. & Knops, A. (2023). The two-network framework of number processing - a step towards a better understanding of the neural origins of developmental dyscalculia. Journal of Neural Transmission.

Klein, E., Zamarian, L., & Kaufmann L. (2023). Challenges in Understanding Numerical Learning: Editorial for Brain Sciences Special Issue "Neurocognitive Signatures of Math (Learning) across the Lifespan and Their Interrelation with Other Aspects of Cognition and Emotion". Brain Sciences, 13(3):420.

Ninaus, M., Cortez, R. Hazin, I., Kiili, K., Wortha, S.M., Klein, E., Weiss, E.M. & Moeller, K. (2023). The added value of game elements: better training performance but comparable learning gains. Educational Technology Research and Development, 1-23.

Smaczny, S., Sperber, C., Jung, S, Moeller, K., Karnath, H.-O. & Klein, E. (2023). Disconnection in a left-hemispheric temporo-parietal network impairs multiplication fact retrieval. Neuroimage 268, 119840.

Wortha, S. M., Klein, E., Lambert, K., Dackermann, T., & Moeller, K. (2023). The relevance of basic numerical skills for fraction processing: Evidence from cross-sectional data. PLOS ONE, 18(1), e0281241.


Göbel, S. M., Terry, R., Klein, E., Hymers, M., & Kaufmann, L. (2022). Impaired Arithmetic Fact Retrieval in an Adult with Developmental Dyscalculia: Evidence from Behavioral and Functional Brain Imaging Data. Brain Sciences, 12(6), 735.

Jung, S., Janssen, R. J., & Klein, E. (2022). Laterality in simple multiplication: Assessing hemispheric specialization of arithmetic fact retrieval in a visual hemifield paradigm. Journal of Experimental Psychology: Human Perception and Performance, 48(4), 351.

Rosenzopf, H., Sperber, C., Wortha, F., Wiesen, D., Muth, A., Klein, E., Moeller, K. & Karnath, H.-O. (2022). Spatial Neglect in the digital Age - Influence of Presentation Format on Patients' Test Behavior. Journal of the International Neuropsychological Society, 1-10.


Greipl, S., Klein, E., Lindstedt, A., Kiili, K., Moeller, K., Karnath, H.- O., Bahnmueller, J., Bloechle, J., & Ninaus, M. (2021). When the brain comes into play: Neurofunctional correlates of emotions and reward in game-based learning. Computers in Human Behavior, 125, 106946.

Jung, S., Moeller, K., Klein, E., & Heller, J. (2021). Mode effect: An issue of perspective? Writing mode differences in a spelling assessment in German children with and without developmental dyslexia. Dyslexia, 1-38. DOI: 10.1002/dys.1675

Klein, E., Babai, R., Ischebeck, A. & Moeller, K. (2021). Editorial: Neuro-cognitive Architecture of Numerical Cognition and Its Development. Frontiers in Human Neuroscience, 15:670460. doi: 10.3389/fnhum.2021.670460


Colling, L. J., Szűcs, D., De Marco, D., Cipora, K., Ulrich, R., Nuerk, H. C., Soltanlou, M., Bryce, D., Chen, S.-C., Schroeder, P.A., Henare, D.T., Chrystall, C.K., Corballis, P.M., Ansari, D., Goffin, C., Sokolowski, H.M., Hancock P.J.B, Millen, A.E., Langton S.R.H., Holmes, K.J., Saviano, M.S., Tummino, T.A., Lindemann, O., Zwaan, R.A., Lukavský, J., Becková, A., Vranka, M.A., Cutini, S., Mammarella, I.C., Mulatti, C., Bell, R., Buchner, A., Mieth, L., Röer, J.P., Klein, E., Huber, S., Moeller, K., Ocampo, B., Lupiáñez, J., Ortiz-Tudela, J., de la Fuente, J., Santiago, S., Ouellet, M., Hubbard, E.M., Toomarian, E.Y., Job, R., Treccani, B., & McShane, B.B. (2020). Registered Replication Report on Fischer, Castel, Dodd, and Pratt (2003). Advances in Methods and Practices in Psychological Science, 1-12. DOI: 10.1177/2515245920903079.

Jung, S., Moeller, K., Karnath, H. O., & Klein, E. (2020). Hemispheric lateralization of arithmetic facts and magnitude processing for two-digit numbers. Frontiers in Human Neuroscience, 14, 88. [OA]

Jung, S., Roesch, S., Klein, E., Dackermann, T., Heller, J., & Moeller, K. (2020). The strategy matters: Bounded and unbounded number line estimation in secondary school children. Cognitive Development, 53, 100839.

Kaufmann, L., von Aster, M., Goebel., S.M., Marksteiner, J. & Klein, E. (2020). Developmental dyscalculia in adults: Current issues and open questions for future research. Learning and Learning Disorders, 9, 126-137. [OA]

Wortha, S.M, Bloechle, J., Ninaus, M., Kiili, K., Lindstedt, A., Bahnmueller, J., Moeller, K. & Klein, E. (2020). Neurofunctional plasticity in fraction learning: an fMRI training study. Trends in Neuroscience and Education, 21, 100141.


Dietrich, J. F., Nuerk, H. C., Klein, E., Moeller, K., & Huber, S. (2019). Set size influences the relationship between ANS acuity and math performance: a result of different strategies? Psychological Research, 83(3), 590-561.

Klein, E., Bieck, S.M., Bloechle, J., Huber, S., Bahnmueller, J., Willmes, K. & Moeller, K. (2019). Anticipation of difficult tasks - neural correlates of negative emotions and emotion regulation. Behavioral and Brain Functions, 15(1), 4.

Klein, E., Willmes, K., Bieck, S.M., Bloechle, J., & Moeller, K. (2019). White matter neuro-plasticity in mental arithmetic: changes in hippocampal connectivity following arithmetic drill training. Cortex, 114, 115-123.

Mock, J., Huber, S., Bloechle, J., Bahnmueller, J., Moeller, K., & Klein, E. (2019). Processing symbolic and non-symbolic proportions: Domain-specific numerical and domain-general processes in intraparietal cortex. Brain Research, 1714, 133-146.

Ninaus, M., Greipl, S., Kiili, K., Lindstedt, A., Huber, S., Klein, E., Karnath, H.-O., & Moeller, K. (2019). Increased emotional engagement in game-based learning - A machine learning approach on facial emotion detection data. Computers & Education, 142, 103641.


Bieck, S. M., Artemenko, C., Moeller, K., & Klein, E. (2018). Low to no effect: application of tRNS during two-digit addition. Frontiers in Neuroscience, 12, 176. [OA]

Bloechle, J., Huber, S., Klein, E., Bahnmueller, J., Moeller, K., & Rennig, J. (2018). Neuro-cognitive mechanisms of global Gestalt perception in visual quantification. NeuroImage, 181, 359-369.

Bloechle, J., Huber, J., Klein, E., Bahnmueller, J., Rennig, J., Moeller, K., & Huber, S. (2018). Spatial arrangement and set size influence the coding of non-symbolic quantities in the intraparietal sulcus. Frontiers in Human Neuroscience, 12, 54. [OA]

Klein, E., Moeller, K., Huber, S., Willmes, K., Kiechl-Kohlendorfer, U. & Kaufmann, L. (2018). Gestational age modulates neural correlates of intentional, but not automatic number magnitude processing in children born preterm. International Journal of Developmental Neuroscience, 65, 38-44.

Mock, J., Huber, S., Bloechle, J., Dietrich, J.F., Bahnmueller, J., Rennig, J., Klein, E., & Moeller, K. (2018). Magnitude processing of symbolic and non-symbolic proportions: an fMRI study. Behavioral and Brain Functions, 14(1), 9. [OA]


Braga, L.W., Amemiya, E., Tauil, A., Suguieda, D., Lacerda, C., Klein, E., Dehaene-Lambertz, G., & Dehaene, S. (2017). Tracking adult literacy acquisition with functional MRI: a single-case study. Mind, Brain and Education, 11, 121-132. [OA]


Bloechle, J., Huber, S., Bahnmueller, J., Rennig, J., Willmes, K., Cavdaroglu, S., Moeller, K., & Klein, E. (2016). Fact Learning in Complex Arithmetic - The Role of the Angular Gyrus revisited. Human Brain Mapping, 37, 3061-3079.

Dietrich, J.F., Huber, S., Klein, E., Willmes, K., Pixner, S., & Moeller, K. (2016). A systematic investigation of accuracy and response time based measures used to index ANS acuity. PLoS One, 11(9):e0163076. [OA]

Huber, S., Klein, E., Moeller, K., & Willmes, K. (2016). Spatial-numerical and ordinal positional associations coexist in parallel. Frontiers in Psychology, 7:438. [OA]

Klein, E., Suchan, J., Moeller, K., Karnath, H.-O., Knops, A., Wood, G., Nuerk, H.-C., & Willmes K. (2016). Considering structural connectivity in the triple code model of numerical cognition - Differential connectivity for magnitude processing and arithmetic facts. Brain Structure and Function, 221(2), 979-95.

Klein, E., Willmes, K., Jung, S., Huber, S., Braga, L. W., & Moeller, K. (2016). Differing connectivity of Exner's area for numbers and letters. Frontiers in Human Neuroscience, 10:281. [OA]

Mock, J., Huber, S., Klein, E., & Moeller, K. (2016). Insights into numerical cognition - Considering eye-fixations in number processing and arithmetic. Psychological Research, 80, 334-359.

Morozova, M., Koschutnig, K., Klein, E., & Wood, G. (2016). Monotonic non-linear transformations as a tool to investigate age-related effects on brain white matter integrity: A Box-Cox investigation. Neuroimage, 125, 1119-30.

Scharinger, M., Domahs, U., Klein, E. & Domahs, F. (2016). Mental representations of vowel features asymmetrically modulate activity in superior temporal sulcus. Brain and Language, 163, 42-49.

Spüler, M., Walter, C., Rosenstiel, W., Gerjets, P., Moeller, K., & Klein, E. (2016). EEG-based prediction of cognitive workload induced by arithmetic - A step towards online adaptation in numerical learning. ZDM Mathematics and Education, 1-12. doi:10.1007/s11858-015-0754-8.


Artemenko, C., Moeller, K., Huber, S., & Klein, E. (2015). Differential influences of unilateral tDCS over the intraparietal cortex on numerical cognition. Frontiers in Human Neuroscience, 9:110. [OA]

Dietrich, J. F., Huber, S., Moeller, K., & Klein, E. (2015). The influence of math anxiety on symbolic and non-symbolic magnitude processing. Frontiers in Psychology, 6:1621. [OA]

Huber, S., Bahnmueller, J., Klein, E., & Moeller, K. (2015). Testing a model of componential processing of multi-symbol numbers - Evidence from measurement units. Psychonomic Bulletin & Review, 22, 1417-1423.

Huber, S., Klein, E., Graf, M., Nuerk, H.-C., Moeller, K., & Willmes, K. (2015). Embodied markedness of parity? Examining handedness effects on parity judgments. Psychological Research, 79, 963-977.

Huber, S., Klein, E., Moeller, K., & Willmes, K. (2015). Comparing a single case to a control group - Applying linear mixed effects models to repeated measures data. Cortex, 71, 148-159.

Jung, S., Halm, K., Huber, W., Willmes, K., & Klein, E. (2015). What letters can "learn" from Arabic digits - fMRI-controlled single case therapy study of peripheral agraphia. Brain and Language, 149, 13-26.

Mihulowicz, K., Klein, E., Nuerk, H.-O., Willmes, K., & Karnath, H.-O. (2015). Spatial displacement of numbers on a vertical number line in spatial neglect. Frontiers in Human Neuroscience, 9:240. [OA]

Moeller, K., Willmes, K., & Klein, E. (2015). A review on functional and structural brain connectivity in numerical cognition. Frontiers in Human Neuroscience. 9:227. [OA]

Rath, D., Domahs, F., Dressel, K., Claros-Salinas, D., Klein, E., Willmes, K., & Krinzinger, H. (2015). Patterns of linguistic and numerical performance in aphasia. Behavioral and Brain Functions, 11:2. [OA]

Shikhare, S., Heim, S., Klein, E., Huber, S. & Willmes, K. (2015). Processing of numerical and proportional quantifiers. Cognitive Science, 39, 1504-1536.


Heisterueber, M., Klein, E., Willmes, K., Heim, S., & Domahs, F. (2014). Processing word prosody - behavioral and neuroimaging evidence for heterogeneous performance in a language with variable stress. Frontiers in Psychology, 5:365. [OA]

Huber, S., Klein, E., Willmes, K., Nuerk, H.-C., & Moeller, K. (2014). Decimal fraction representations are not distinct from natural number representations - Evidence from a combined eye-tracking and computational modelling approach. Frontiers in Human Neuroscience, 8:172. [OA]

Klein, E., Huber, S., Nuerk, H.-C., & Moeller, K.(2014). Operational momentum affects eye fixation behaviour. Quarterly Journal of Experimental Psychology, 67(8), 1614-1625.

Klein, E., Moeller., K., Kiechl-Kohlendorfer, U., Kremser, C., Starke, M., Cohen Kadosh, R., Pupp-Peglow, U., Schocke, M., & Kaufmann, L. (2014). Processing of intentional and automatic number magnitudes in children born prematurely - Evidence from fMRI. Developmental Neuropsychology, 39(5), 342-364.

Mihulowicz, U., Willmes, K., Karnath, H.-O., & Klein, E. (2014). Single-digit arithmetic processing - anatomical evidence from statistical voxel-based lesion analysis. Frontiers in Human Neuroscience, 8:286. [OA]

Moeller, K., Klein, E., Kucian, K., & Willmes, K. (2014). Numerical Development - From cognitive functions to neural underpinnings. Frontiers in Psychology, 5:1047. [OA]

Willmes, K., Moeller, K., & Klein, E. (2014). Where numbers meet words - A common ventral network for semantic classification. Scandinavian Journal of Psychology, 55, 202-211.


Domahs, U., Klein, E., Huber, W., & Domahs, F. (2013). Good, bad and ugly word stress - fMRI evidence for foot structure driven processing of prosodic violations. Brain and Language, 125(3), 272-282.

Heisterüber, M., Schäffner, S., Heim, S., Klein, E., Willmes, K., & Domahs, F. (2013). Funktionelle Bildgebung subklinischer Defizite in der Wortakzentverarbeitung [Neuroimaging Studies of Subclinical Deficits in Word Stress Processing]. Sprache, Stimme, Gehör, 37(1), 21-23.

Klein, E., Bahnmueller, J., Mann, A., Pixner, S., Kaufmann, L., Nuerk, H.-C. & Moeller, K. (2013). Language influences on numerical development - Inversion effects on multi-digit number processing. Frontiers in Psychology, 4:480. [OA]

Klein, E., Mann, A., Huber, S., Bloechle, J., Willmes, K., Karim, A.A., Nuerk, H.-C., & Moeller, K. (2013). Bilateral bi-cephalic tDCS with two active electrodes of the same polarity modulates bilateral cognitive processes differentially. PLoS ONE 8(8): e71607. [OA]

Klein, E., Moeller, K., Glauche, V., Weiller, C., & Willmes, K. (2013). Processing pathways in mental arithmetic - Evidence from probabilistic fiber tracking. PLoS ONE, 8(1): e55455. [OA]

Klein, E., Moeller, K., Zinsberger, D., Zauner, H., Wood, G., Willmes, K., Haider, C., Gassner, A., & Nuerk, H.-C. (2013). Object-based neglect in number processing. Behavioral and Brain Functions, 9:5. [OA]

Klein, E.,Moeller, K., & Willmes, K.(2013).A neural disconnection hypothesis on impaired numerical processing. Frontiers in Human Neuroscience, 7:663. [OA]

Moeller, K., Klein, E., & Nuerk, H.-C. (2013). Influences of cognitive control in numerical cognition - Adaptation by binding for implicit learning. Topics in Cognitive Science, 5(2), 335-353.

Moeller, K., Klein, E., Willmes, K., & Nuerk, H.-C. (2013). Magnitude representation in sequential comparison of two-digit numbers is not holistic either. Cognitive Processing, 14, 51-62.

Ochtrup, M.-T., Rath, D., Klein, E., Krinzinger, H., Willmes, K. & Domahs, F. (2013). Are Number Words Fundamentally Different? A Qualitative Analysis of Aphasic Errors in Word and Number Word Production. International Journal of Speech & Language Pathology and Audiology, 1, 12-28.


Domahs, F., Klein, E., Moeller, K., Nuerk, H.-C., Yoon, B.-C., & Willmes, K. (2012). Multimodal semantic quantity representations Further evidence from Korean Sign Language. Frontiers in Psychology, 2, 389. [OA]

Moeller, K., Klein, E., Nuerk, H.-C., & Cohen Kadosh, R. (2012). A unitary or multiple representations of numerical magnitude? - The case of symbolic and non-symbolic two-digit quantities. Frontiers in Psychology, 3, 191. [OA]


Domahs, F., Rath, D., Klein, E., Krinzinger, H. & Willmes, K. (2011). Numerische und sprachliche Kognition bei Patienten mit Aphasie. Zeitschrift für Neuropsychologie, 3, 199-200.

Klein, E., Domahs, U, Grande, M., & Domahs, F. (2011). Neuro-cognitive foundations of word stress processing - evidence from fMRI. Behavioral and Brain Functions, 7:15. [OA]

Klein, E., Moeller, K., Willmes, K., Nuerk, H.-C., & Domahs, F. (2011). The influence of implicit hand-based representations on mental arithmetic. Frontiers in Psychology, 2, 197. [OA]

Moeller K., Klein, E., Fischer, M. H., Nuerk, H.-C., & Willmes, K. (2011). Representation of Multiplication Facts - Evidence for partial verbal coding. Behavioral and Brain Functions, 7:25. [OA]

Moeller, K., Klein, E., & Nuerk, H.-C. (2011). Three processes underlying the carry effect in addition - Evidence from eye-tracking. British Journal of Psychology, 102, 623-645.

Moeller, K., Klein, E., & Nuerk, H.-C. (2011). (No) small adults - Children's processing of carry addition problems. Developmental Neuropsychology, 36(6), 702-720.

Nuerk, H.-C., Moeller, K., Klein, E., Willmes, K., & Fischer, M. H. (2011). Extending the Mental Number Line - A Review of Multi-Digit Number Processing. Journal of Psychology, 219, 3-22.


Klein, E., Moeller, K., Dressel, K., Domahs, F., Wood, G., Willmes, K., & Nuerk, H.-C. (2010). To carry or not to carry - is this the question? Disentangling the carry effect in multi-digit addition. Acta Psychologica, 135, 67-76.

Klein, E., Moeller, K., Nuerk, H.-C., & Willmes K. (2010). On the neuro-cognitive foundations of basic auditory number processing. Behavioral and Brain Functions, 6:42. [OA]

Klein, E., Willmes, K., Dressel, K., Domahs, F., Wood, G., Nuerk, H.-C., & Moeller, K. (2010). Categorical and continuous - Disentangling the neural correlates of the carry effect in multi-digit addition. Behavioral and Brain Functions, 6:70. [OA]


Klein, E., Nuerk, H.-C., Wood, G., Knops, A., & Willmes, K. (2009). The exact vs. approximate distinction in numerical cognition may not be exact, but only approximate - How different processes work together in multi-digit addition. Brain and Cognition, 2, 369-381.

Moeller, K., Pixner, S., Klein, E., Cress, U., & Nuerk, H.-C. (2009). Zahlenverarbeitung ist nicht gleich Rechnen - Eine Beschreibung basisnumerischer Repräsentationen und spezifischer Interventionsansätze. Prävention und Rehabilitation, 21(3), 121-136.

Zaunmüller, L., Domahs, F., Dressel, K., Lonnemann, J., Klein, E., Ischebeck, A., & Willmes, K. (2009). Rehabilitation of arithmetic fact retrieval via extensive practice A combined fMRI and behavioural case-study. Neuropsychological Rehabilitation, 19, 422-443.

Book chapters

Klein, E., Jung, S., & Kaufmann, L. (2021). A Neuropsychological Perspective on Mathematics and Language Processing. In: Diversity Dimensions in Mathematics and Language (pp. 40-64). Eds.: Fritz-Stratmann, A., Herzog, M., & Gürsoy, E. Berlin. De Gruyter. DOI: 10.1515/9783110661941-003

Fischer, U., Klein, E., Dackermann, T., & Moeller, K. (2017). The Physiology of Numerical Learning - From Neural Correlates to Embodied Trainings. In: The Psychology of Digital Learning (pp.21-40).Heidelberg. Springer-Verlag.

Moeller, K., Klein, E., & Kaufmann, L. (2017). Bedeutung der neurokognitiven und bildgebenden Befunde für ein besseres Verständnis von Rechenschwierigkeiten. In A. Fritz, S. Schmidt, & G. Ricken (Eds.), Handbuch Rechenschwäche (S. 80-95, 3. völlig überarbeitete Auflage). Weinheim: Beltz. [Handbook Dyscalculia. 3rd. revised edition].

Roesch, S., Klein, E., Kaufmann, L., & Moeller, K. (2017). Zahlen, Rechnen und das Gehirn: Können neurokognitive Befunde zu einem besseren Verständnis des Rechnens und der Rechenstörung beitragen? Lernwege bei Dyskalkulie und Legasthenie - Landesverband NRW e.V.

Willmes, K. & Klein, E. (2014). Akalkulie [Acalculia]. In H.-O. Karnath, W. Ziegler, & G. Goldenberg (Eds.), Klinische Neuropsychologie - Kognitive Neurologie (pp. 133-146). Stuttgart: Thieme-Verlag.

Moeller, K., Pixner, S., Klein, E., Cress, U., & Nuerk, H.-C. (2013). Zahlenverarbeitung ist nicht gleich Rechnen - Eine Beschreibung basisnumerischer Repräsentationen und spezifischer Interventionsansätze. In S. Pixner & K. Moeller (Hrsg.): Lernstörungen. München. Hachinger Verlag, 1. Auflage, 70-94.

Nuerk, H.-C., Klein, E. & Willmes, K. (2012). Zahlenverarbeitung und Rechnen [Number processing and calculation]. In F. Schneider & G. Fink (Hrsg.): Funktionelle MRT in Psychiatrie und Neurologie. Heidelberg. Springer-Verlag, 2. Auflage, 443-455.

Willmes K., Klein, E. & Nuerk, H.-C. (2012): Akalkulie [Acalculia]. In F. Schneider & G. Fink (Hrsg.) Funktionelle MRT in Psychiatrie und Neurologie. Heidelberg. Springer-Verlag, 2. Auflage, 577-586.

Elise Klein - MD PhD
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