Selected Publications


Scott Frey, Binal Motawar, Kelli Buchanan, Christina Kaufman, Phil Stevens, Carmen Cirstea, Sean Morrow (2022). Greater and more natural use of the upper limbs during everyday life by former amputees versus prosthesis users. Neurorehabilitation and Neural Repair, 2022 Jan 7.


Benjamin A. Philip, , Mark P. McAvoy, Scott H. Frey (2020). Interhemispheric Parietal-Frontal Connectivity Predicts the Ability to Acquire a Nondominant Hand Skill. Brain Connectivity, 10 May 2021.

Randerath J, Finkel L, Shigaki C, Burris J, Nanda A, Hwang P, Frey SH. (2021). Is This Within Reach? Left but Not Right Brain Damage Affects Affordance Judgment Tendencies. Frontiers in Human Neuroscience, 27 January 2021.


Frey, Scott H. (2020). Harnessing Neuroplasticity to Enhance Functional Recovery in Allogeneic Hand Transplant and Heterotopic Hand Replant Recipients. Defense Technical Information Center,

Valyear KF, Philip BA, Cirstea CM, Chen PW, Baune NA, Marchal N, Frey S. H. (2020). Interhemispheric transfer of post-amputation cortical plasticity within the human somatosensory cortex. Neuroimage, 2020 Feb 1;206:116291.


Peng H, Cirstea CM, Kaufman CL, Frey S. H. (2019). Microstructural integrity of corticospinal and medial lemniscus tracts: insights from diffusion tensor tractography of right-hand amputees. J Neurophysiol, 2019 Jul 1;122(1):316-324.

M Ma, X Wang, Y Duan, SH Frey, X Gu (2019). Optimal mass transport based brain morphometry for patients with congenital hand deformities. The Visual Computer volume 35pages13111325(2019)


Randerath, J., Finkel, L., Shigaki, C., Burris, J., Nanda, A., Hwang, P., & Frey, S. H. (2018). Does it fit? - Impaired affordance perception after stroke. Neuropsychologia, 108, 92–102.

Lisa Finkel, Katharina Hogrefe, Scott H Frey, Georg Goldenberg, Jennifer Randerath (2018). It takes two to pantomime: Communication meets motor cognition. Neuroimage Clin., 2018 Jun 21;19:1008-1017.

Daniela Mattos, Scott H Frey (2018). How to Press a Button: Perspectives from the Science of Motor Control. Stevens' Handbook of Experimental Psychology and Cognitive Neuroscience 2, 1-14.

SH Frey (2018). Harnessing Neuroplasticity to Enhance Functional Recovery in Allogeneic Hand Transplant and Heterotopic Hand Replant Recipients.

Sainburg, R. L., Liew, S.-L., Frey, S. H., & Clark, F. (2017). Promoting Translational Research Among Movement Science, Occupational Science, and Occupational Therapy. Journal of Motor Behavior, 49(1), 1–7.

Randerath J, Valyear KF, Philip BA, Frey S. H. (2017) Contributions of the parietal cortex to increased efficiency of planning-based action selection. Neuropsychologia,2017 Oct;105:135-143.

Valyear, K. F., Mattos, D., Philip, B. A., Kaufman, C., & Frey, S. H. (2017). Grasping with a new hand: Improved performance and normalized grasp-selective brain responses despite persistent functional changes in primary motor cortex and low-level sensory and motor impairments. NeuroImage.

Cirstea, C. M., Choi, I.-Y., Lee, P., Peng, H., Kaufman, C. L., & Frey, S. H. (2017). Magnetic resonance spectroscopy of current hand amputees reveals evidence for neuronal-level changes in former sensorimotor cortex. Journal of Neurophysiology, 117(4), 1821–1830.

B Philip, K Valyear, C Cirstea, S Frey (2017). Reorganization of Primary Somatosensory Cortex After Upper Limb Amputation May Lack Functional Significance. Archives of Physical Medicine and Rehabilitation 98 (10), e103.


Kroliczak, G., Piper, B.K., & Frey, S.H. (2016). Specialization of the left supramarginal gyrus for hand-independent praxis representation is not related to hand dominance. Neuropsychologia Journal, DOI:10.1016/j.neuropsychologia.2016.03.023

Sachet, A. B., Frey, S. H., Jacobs, S., & Taylor, M. (2016). Development of the correspondence between real and imagined fine and gross motor actions. Journal of Cognition and Development. 17:1, 162-179, DOI: 0.1080/15248372.2014.963585

Povinelli, D. J., & Frey, S. H. (2016). Constraints on the exploitation of the functional properties of objects in expert tool-using chimpanzees (Pan troglodytes). Cortex; a Journal Devoted to the Study of the Nervous System and Behavior, 82, 11–23.

Philip, B. A., & Frey, S. H. (2016). Increased functional connectivity between cortical hand areas and praxis network associated with training-related improvements in non-dominant hand precision drawing. Neuropsychologia, 87, 157–168.

A Barrett, R Zhang, S Frey, M Oh-Park (2016). Spatial-Motor Flexibility (SMF) and Post-Stroke Cognitive Recovery (P6.014). Neurology 86 (16 Supplement), P6. 014,


Peng, H., Wang, X., Ye, D., Frey, S.H., & Gu, X. (2015).  Brain Morphometry on Congenital Hand Deformities based on Teichmuller Space Theory.  Computer-aided Design, 58, 84-91.

Frey, S.H.,  & Chen, Pin-Wei. (2015). The Mirror Neuron Analogy in Rehabilitation Neuroscience. Physics of Life Reviews. 12, 106-107.

Frey, S.H., Hansen, M., & Marchal, N. (2015).  Grasping with the press of a button: grasp selective responses in the human anterior intraparietal sulcus depend on non-arbitrary causal relationships between hand movements and end-effector actions. J. Cognitive Neuroscience, 27, 1146-1160.

Valyear, K. F. & Frey, S.H.  (2015).  fMRI repetition suppression reveals involvement of left rostral supramarginal gyrus and bilateral superior parietal cortex in hand selection.  Neuroimage. doi: 10.1016/j.neuroimage.2015.03.058.

Philip, B., Buckon, C. Sienko, S.; Aiona, M., Ross, S., & Frey, S.H.  (2015).  Maturation and experience in action representation: unilateral congenital amelia, bilateral deficits. Neuropsychologia, 75, 420-430.

Randerath, J. & Frey, S. H. (2015). Diagnostics and Training of Affordance Perception in Healthy Young Adults-Implications for Post-Stroke Neurorehabilitation. Frontiers in human neuroscience 9, 674, doi:10.3389/fnhum.2015.00674.

Randerath, J., Valyear, K., Hood, A., & Frey, S.H.  (2015). Two routes to the same action – an action repetition priming study.  J. of Motor Behavior, 47, 2, 142-152.

Valyear KF, Frey S. H. (2015). Human posterior parietal cortex mediates hand-specific planning. Neuroimage. 2015 Jul 1;114:226-38.

SH Frey (2015).  Why Brain Science is Essential to the Success of Hand Allotransplantation. The Science of Reconstructive Transplantation pp 361-375


Macuga, K.L., & Frey, S.H.”  (2014).  “Differential contributions of the superior and inferior parietal cortex to feedback versus feed forward control of tools.”  Neuroimage. 92,  36 – 52.

Philip, B.A., & Frey, S.H.  (2014).  Compensatory changes accompanying chronic forced use of the non-dominant hand by unilateral amputees.  J. of Neuroscience, 34(10):3622-3631.

Shigaki, C.L., Frey, S.H., & Barrett, A.M.   Rehabilitation of Post Stroke Cognition. Seminars in Neurology, 34, 5, 496-503.

Ricker JH, DeLuca J, Frey S. H. (2014). On the changing roles of neuroimaging in rehabilitation science. Brain Imaging Behav. 014 Sep;8(3):333-4.

Shigaki CL, Frey S. H., Barrett AM. (2014). Rehabilitation of poststroke cognition. Semin Neurol. 2014 Nov;34(5):496-503.

C BUCKON, S SIENKO, M AIONA, S FREY (2014). Motor imagery and planning deficits in children with congenital hemiplegia: G9. Developmental Medicine & Child Neurology, 2014

K Macuga, S Frey (2014). Tool use and representations of reachable space in the superior parietal lobe. Journal of Vision, 14:10 306-306
C Kaufman, K Valyear, B Philip, B Blair, J Kutz, R Ouseph, M Marvin, S Frey (2014). Abrogation of Phantom Limb Pain Following Hand Transplantation.: Abstract# D2830 Transplantation: July 15, 2014 - Volume 98 - Issue - p 411,
Scott H Frey (2014).  Cortical Organization in Allogeneic Hand Transplants or Heterotopic Hand Replants,
Scott H Frey, Scott T Grafton (2014). Finding the actor in reciprocal affordance. MIT Press,


Randerath, J., Martin, K., & Frey, S.H.  (2013).  Are tool properties always processed automatically? The role of tool use context and task complexity.  Cortex, 49,  1679-1693.

Philip, B.A., & Frey, S.H.  (2013).  Stimulus-response correspondence across peripersonal space is unaffected by chronic unilateral limb loss Experimental Brain Research, 224, 3, 373 – 382.

Valyear, K.F., & Frey, S.H.  (2013).  Hand selection for object grasping is influenced by recent motor history.  Psychonomic Bulletin & Rev., 21, 2, 566-573


Frey, S.H. & Povinelli, D. (2012). Comparative investigations of manual action representations: evidence that chimpanzees represent the costs of potential future actions involving tools. Phil. Trans. of the Royal Soc. B., 1585, 48-58.

Macuga, K., & Frey, S.H. (2012). “Neural representations involved in observed, imagined, and imitated actions are dissociable and hierarchically organized.” Neuroimage, 59, 3, 2798-2807.

Bogdanov S, Smith J, Frey SH. (2012) Former hand territory activity increases after amputation during intact hand movements, but Is unaffected by illusory visual feedback. Neurorehabilitation and Neural Repair, 26, 6, 604-615.

Macuga, K., & Frey, S.H. (2012). ”Motor imagery of tool use: Relationship to actual use and adherence to Fitts’ law across tasks.” Experimental Brain Research, 218, 169-179

Randerath J, Martin KR, Frey SH. (2012) Are tool properties always processed automatically? The role of tool use context and task complexity. Cortex, 2013 Jun;49(6):1679-93,

Philip BA, Frey S. H. (2012) Preserved grip selection planning in chronic unilateral upper extremity amputees. Experimental Brain Research, 2011 Oct;214(3):437-52,

Philip BA, Frey S. H. (2014) Erratum to: Preserved grip selection planning in chronic unilateral upper extremity amputees. Experimental Brain Research, 232pages345346(2014),


Povinelli, D.J., Reaux, J.E., & Frey, S.H.  (2011).  “Chimpanzees’ tool use within peripersonal space provides evidence for separable representations of hand and tool even during active use.” Neuropsychologia, 48, 243-247.  PMC19766665

Macuga, K. & Frey, S.H.  (2011).  “Selective responses in right inferior frontal and supramarginal gyri differentiate between observed movements of oneself vs. another”.  Neuropsychologia, 49, 5, 1202-1207.  PMC3078186

Kroliczak, G., & Frey, S.H.  (2011).  “Atypical lateralization of language predicts cerebral asymmetries in parietal gesture representations.” Neuropsychologia, 49, 7, 1698-1702. PMC3100506

Frey, S.H., Fogassi, L., Grafton, S.T., Picard, N., Rothwell, J., Schweighofer, N., Corbetta, M., & Fitzpatrick, S.M.  (2011).  “Neurological principles and rehabilitation of action disorders:  computation, anatomy and physiology (CAP) model.”  Neurorehabilitation and Neural Repair, 25, 5 suppl., 6S-20S.

Marangon, M., Jacobs, S., & Frey, S.H.  (2011).  “Context-sensitivity of grasp representations in human rostral inferior parietal lobule.”  J. of Neurophysiology, 105,  5, 2536-2546. PMC3094173

Martin, K., Jacobs, S., and Frey, S.H.  (2011).  “Handedness-related differences in contributions of anterior intraparietal and ventral premotor cortices to feed-forward grip selection involving the hands or a recently mastered tool.”  Neuroimage, 57, 2, 502-512. PMC3114104

Philip, B.A., & Frey, S.H.  (2011).  Preservation of movement planning in patients with chronic unilateral upper extremity amputations.  Experimental Brain Research, 214,  3, 437-452.

M Marangon, S Jacobs, SH Frey (2011). Evidence for context sensitivity of grasp representations in human parietal and premotor cortices.  Journal of neurophysiology, 2536-2546.

J Smith, SH Frey (2011). Use of independent component analysis to define regions of interest for fMRI studies.  International Society for Magnet Resonance in Medicine (ISMRM) Montreal,


Jacobs, S., Danielmeier, C. & Frey, S.H. (2010). Human anterior intraparietal and ventral premotor cortices support representations of grasping with the hand or a novel tool. Journal of Cognitive Neuroscience, 2010.

SH Frey (2010). Cortical Mechanisms of Multi-step Action Planning.


Povinelli, D.J., Reaux, J.E., & Frey, S.H. (2009). Chimpanzees context-dependent tool use provides evidence for separable representations of hand and tool even during active use within peripersonal space. Neuropsychologia, 2009.

Arbib, M.A., Bonaiuto, J.B., Jacobs, S. & Frey, S.H. (2009). Tool use and the distalization of the end-effector Psychological Research, 73, 441-462 Kroliczak, G. & Frey, S.H. (2009). A Common Network in the Left Cerebral Hemisphere Represents Planning of Tool Use Pantomimes and Familiar Intransitive Gestures at the Hand-Independent Level Cerebral Cortex, 19(10): 2396-410.

J Smith, S Bogdanov, S Watrous, S Frey (2009). Rapid digit mapping in the human brain at 3T. Proc. Intl. Soc. Mag. Reson. Med. 17,


Frey, S.H., Bogdanov, S.B., Smith, J.C., Watrous, S. & Breidenbach, W.C. (2008). Chronically Deafferented Sensory Cortex Recovers a Grossly Typical Organization after Allogenic Hand Transplantation Current Biology 18, 1530–1534.

Frey, S.H. (2008). Tool use, communicative gesture and cerebral asymmetries in the modern human brain. Philos Trans R Soc Lond B Biol Sci 363.


Farrer, C., Frey, S.H., Tunik, E., Turk, D., Inati, S., & Grafton. (2007). The angular gyrus computes action awareness representations. Cerebral Cortex.

Frey, S.H. (2007). What Puts the How in Where: Semantic and Sensori-Motor Bases of Everyday Tool Use.. Cortex, 43, 368-375.

Frey, S.H. (2007). Neurological specialization for manual gesture and tool use in humans. To appear in Jon H. Kaas (Ed.), Evolution of the nervous systems.New York: Elsevier.


Frey, S.H. & Gerry, V.E. (2006). Modulation of neural activity during observational learning of actions and their sequential orders. J. Neuroscience, 26, 13194-13201.

Newman-Norlund, R., Frey, S.H., Petitto, L-A., & Grafton, S.T. (2006). Anatomic distinctions between visual and auditory second language acquisition. Journal of Cognitive Neuroscience, 18, 1984-1997.

Roger D Newman-Norlund, Scott H Frey, Laura-Ann Petitto, Scott T Grafton (2006). Anatomical Substrates of Visual and Auditory Miniature Second-language Learning. Journal of Cognitive Neuroscience, Volume 18, Issue 12, December 2006, p.1984-1997,


Frey, S.H., Newman-Norlund, R.N., & Grafton, S.T. (2005). A distributed network in the left cerebral hemisphere for planning everyday tool use skills. Cerebral Cortex.

Frey, S.H., Vinton, D., Newman-Norlund, R., & Grafton. S.T. (2005). Cortical topography of human anterior intraparietal cortex active during visually-guided grasping. Cognitive Brain Research.

Tunik E, Frey, S.H., & Grafton ST. (2005). Virtual lesions of the anterior intraparietal area disrupt goal-dependent on-line adjustments of grasp. Nature Neuroscience,8, 505-511.

Frey, S.H., Funnell, M.G., Gerry, V.E. & Gazzaniga, M.S. (2005). A dissociation between tool use skills and hand dominance: Insights from left and right-handed callosotomy patients. Journal of Cognitive Neuroscience, 17, 262-272.

Müsseler, J., Wühr, P., Danielmeier, C. & Zysset, S. (2005). Action-induced blindness with lateralized stimuli and responses. Experimental Brain Research, 160, 214-222.

Buxbaum, L.J., Frey, S.H., & Bartlett-Williams, M. (2005). Deficient internal models for planning hand-object interactions in apraxia. Neuropsychologia, 43, 917-929.


Danielmeier, C., Zysset, S., Müsseler, J. & von Cramon, D.Y. (2004). Where action impairs visual encoding: an event-related fMRI study. Cognitive Brain Research, 21(1), 39-48.

Johnson-Frey, S.H. (2004). The neural bases of complex human tool use. Trends in Cognitive Sciences, 8, 71-78.

Johnson-Frey, S.H., McCarty, M., & Keen, R. (2004). Reaching beyond spatial perception: effects of intended future actions on visually-guided prehension.  Visual Cognition, 11, 371-399.

Leslie, K., Johnson-Frey, S.H., & Grafton, S.T. (2004). Functional imaging of face and hand imitation: Toward a motor theory of empathy. Neuroimage, 21, 601607 .

Johnson-Frey, S.H. (2004). Simulation through stimulation? Motor imagery and functional reorganization in hemiplegic stroke patients. Brain and Cognition, 55, 328-331.

Johnson-Frey, S.H .(2004). The organization of action representations in posterior parietal cortex. Behavioral and Brain Sciences, 27, 40-41.

Taking Action: Cognitive Neuroscience Perspectives on intentional acts. MIT Press. Edited by Scott H. Johnson-Frey


Johnson-Frey, S.H. (2003). What’s so special about human tool use? Neuron, 39, 201-204.

Johnson-Frey, S.H., Maloof, F.R, Newman-Norlund, R., Farrer, C., Inati, S. & Grafton, S.T. (2003). Actions or Hand-Object Interactions? Human Inferior Frontal Cortex and Action Observation. Neuron, 39, 1053-1058.

Johnson-Frey, S.H. (2003). Mirror neurons, Broca’s area and language: reflecting on the evidence. Behavioral and Brain Sciences 26, 226-227.

Johnson-Frey, S.H. (2003). Cortical Representations of Tool Use in Humans. In S.H. Johnson-Frey (Ed.) . Cognitive neuroscience perspectives on the problem of intentional action. MIT Press.

Johnson-Frey, S.H. & Grafton S.T. (2003). From “Acting On” to “Acting With”: the functional anatomy of action representation. In C. Prablanc, D. Pelisson, & Y. Rossetti (Eds.), Space Coding and Action Production. New York: Elsevier.


Johnson, S.H., Saykin, A.J., Sprehn, G. (2002). Intact motor imagery in chronic upper limb hemiplegics: evidence for activity-independent action representations. Journal of Cognitive Neuroscience, 14:6, pp. 841-852

Nakatani, C., Pollatsek, A. & Johnson, S.H. (2002). Viewpoint-dependent recognition of scenes. Quarterly Journal of Experimental Psychology: Human Experimental Psychology, 55A, 115-139.

Johnson, S.H., Rotte, M., Grafton, S.T., Hinrichs, H., Gazzaniga, M.S., & Heinze, H-J. (2002) Selective Activation of a Parietofrontal Circuit during Implicitly Imagined Prehension. NeuroImage, 17, 1693-1704.


Johnson, S.H. (2001). Seeing both sides at once: Effects of object structure and orientation of recognizing novel views of three-dimensional objects. Journal of Experimental Psychology: Human Perception and Performance, 27, 1468-1484 .

Johnson, S.H., & Corballis, P.M, &Gazzaniga, M.S. (2001). Within grasp but out of reach: Evidence for a double dissociation imagined hand and arm movements in the left cerebral hemisphere. Neuropsychologia,39, 36-50.


Johnson, S.H. (2000). Thinking ahead: The Case For Motor Imagery in Prospective Judgments of Prehension. Cognition, 74, 33-70.

Johnson, S.H . (2000). Imagining the impossible: Intact motor representations in hemiplegic adults. Neuroreport, 11, 729-732.


Johnson, S.H. (1998). Cerebral organization of motor imagery: Contralateral control of grip selection in mentally-represented prehension. Psychological Science, 9, 219-222.


James E Cutting, Ken Springer, Paul A Braren, Scott H Johnson (1992). Wayfinding on foot from information in retinal, not optical, flow. Journal of Experimental Psychology, : General, 121(1), 41–72.

*Please note that Dr. Frey has also published previously as “Scott H. Johnson” and “Scott H. Johnson-Frey.” For complete list of publications, please inquire at