Experimental Brain Research

, Volume 233, Issue 5, pp 1625–1637 | Cite as

Neural processes mediating the preparation and release of focal motor output are suppressed or absent during imagined movement

  • Jeremy S. Eagles
  • Anthony N. Carlsen
  • Colum D. MacKinnonEmail author
Research Article


Movements that are executed or imagined activate a similar subset of cortical regions, but the extent to which this activity represents functionally equivalent neural processes is unclear. During preparation for an executed movement, presentation of a startling acoustic stimulus (SAS) evokes a premature release of the planned movement with the spatial and temporal features of the tasks essentially intact. If imagined movement incorporates the same preparatory processes as executed movement, then a SAS should release the planned movement during preparation. This hypothesis was tested using an instructed-delay cueing paradigm during which subjects were required to rapidly release a handheld weight while maintaining the posture of the arm or to perform first-person imagery of the same task while holding the weight. In a subset of trials, a SAS was presented at 1500, 500, or 200 ms prior to the release cue. Task-appropriate preparation during executed and imagined movements was confirmed by electroencephalographic recording of a contingent negative variation waveform. During preparation for executed movement, a SAS often resulted in premature release of the weight with the probability of release progressively increasing from 24 % at −1500 ms to 80 % at −200 ms. In contrast, the SAS rarely (<2 % of trials) triggered a release of the weight during imagined movement. However, the SAS frequently evoked the planned postural response (suppression of bicep brachii muscle activity) irrespective of the task or timing of stimulation (even during periods of postural hold without preparation). These findings provide evidence that neural processes mediating the preparation and release of the focal motor task (release of the weight) are markedly attenuated or absent during imagined movement and that postural and focal components of the task are prepared independently.


Imagined movement Movement preparation Startle Contingent negative variation 



We thank the volunteers for these experiments, and the technical assistance from Mr. Di Zhang. J.E. was supported by a Grant from NIH T32 HD057845 and ANC by a Grant from the Natural Sciences and Engineering Research Council of Canada (PDF-357177).


  1. Alibiglou L, MacKinnon CD (2012) The early release of planned movement by acoustic startle can be delayed by transcranial magnetic stimulation over the motor cortex. J Physiol 590:919–936CrossRefPubMedCentralPubMedGoogle Scholar
  2. Aruin AS, Shiratori T, Latash ML (2001) The role of action in postural preparation for loading and unloading in standing subjects. Exp Brain Res 138:458–466CrossRefPubMedGoogle Scholar
  3. Bakker M, de Lange FP, Stevens JA, Toni I, Bloem BR (2007) Motor imagery of gait: a quantitative approach. Exp Brain Res 179:497–504CrossRefPubMedGoogle Scholar
  4. Bakker M, De Lange FP, Helmich RC, Scheeringa R, Bloem BR, Toni I (2008) Cerebral correlates of motor imagery of normal and precision gait. Neuroimage 41:998–1010CrossRefPubMedGoogle Scholar
  5. Barclay-Goddard RE, Stevenson TJ, Poluha W, Thalman L (2011) Mental practice for treating upper extremity deficits in individuals with hemiparesis after stroke. Cochrane Database Syst Rev 5:1–47Google Scholar
  6. Bateni H, Zecevic A, McIlroy WE, Maki BE (2004) Resolving conflicts in task demands during balance recovery: does holding an object inhibit compensatory grasping? Exp Brain Res 157:49–58CrossRefPubMedGoogle Scholar
  7. Braun S, Kleynen M, Schols J, Schack T, Beurskens A, Wade D (2008) Using mental practice in stroke rehabilitation: a framework. Clin Rehabil 22:579–591CrossRefPubMedGoogle Scholar
  8. Braun S, Kleynen M, van Heel T, Kruithof N, Wade D, Beurskens A (2013) The effects of mental practice in neurological rehabilitation; a systematic review and meta-analysis. Front Hum Neurosci 7:390CrossRefPubMedCentralPubMedGoogle Scholar
  9. Brown P, Rothwell JC, Thompson PD, Britton TC, Day BL, Marsden CD (1991) New observations on the normal auditory startle reflex in man. Brain 114(Pt 4):1891–1902CrossRefPubMedGoogle Scholar
  10. Butler AJ, Page SJ (2006) Mental practice with motor imagery: evidence for motor recovery and cortical reorganization after stroke. Arch Phys Med Rehabil 87:S2–S11CrossRefPubMedCentralPubMedGoogle Scholar
  11. Caldara R, Deiber MP, Andrey C, Michel CM, Thut G, Hauert CA (2004) Actual and mental motor preparation and execution: a spatiotemporal ERP study. Exp Brain Res 159:389–399CrossRefPubMedGoogle Scholar
  12. Carlsen AN, MacKinnon CD (2010) Motor preparation is modulated by the resolution of the response timing information. Brain Res 1322:38–49CrossRefPubMedGoogle Scholar
  13. Carlsen AN, Chua R, Inglis JT, Sanderson DJ, Franks IM (2004a) Can prepared responses be stored subcortically? Exp Brain Res 159:301–309CrossRefPubMedGoogle Scholar
  14. Carlsen AN, Chua R, Inglis JT, Sanderson DJ, Franks IM (2004b) Prepared movements are elicited early by startle. J Mot Behav 36:253–264CrossRefPubMedGoogle Scholar
  15. Carlsen AN, Dakin CJ, Chua R, Franks IM (2007) Startle produces early response latencies that are distinct from stimulus intensity effects. Exp Brain Res 176:199–205CrossRefPubMedGoogle Scholar
  16. Carlsen AN, Almeida QJ, Franks IM (2012a) Startle decreases reaction time to active inhibition. Exp Brain Res 217:7–14CrossRefPubMedGoogle Scholar
  17. Carlsen AN, Maslovat D, Franks IM (2012b) Preparation for voluntary movement in healthy and clinical populations: evidence from startle. Clin Neurophysiol 123:21–33CrossRefPubMedGoogle Scholar
  18. Carrillo-de-la-Pena MT, Galdo-Alvarez S, Lastra-Barreira C (2008) Equivalent is not equal: primary motor cortex (MI) activation during motor imagery and execution of sequential movements. Brain Res 1226:134–143CrossRefPubMedGoogle Scholar
  19. Cisek P, Kalaska JF (2004) Neural correlates of mental rehearsal in dorsal premotor cortex. Nature 431:993–996CrossRefPubMedGoogle Scholar
  20. Confais J, Kilavik BE, Ponce-Alvarez A, Riehle A (2012) On the anticipatory precue activity in motor cortex. J Neurosci 32:15359–15368CrossRefPubMedGoogle Scholar
  21. Cordo PJ, Nashner LM (1982) Properties of postural adjustments associated with rapid arm movements. J Neurophysiol 47:287–302PubMedGoogle Scholar
  22. Cressman EK, Carlsen AN, Chua R, Franks IM (2006) Temporal uncertainty does not affect response latencies of movements produced during startle reactions. Exp Brain Res 171:278–282CrossRefPubMedGoogle Scholar
  23. Cunnington R, Iansek R, Bradshaw JL, Phillips JG (1996) Movement-related potentials associated with movement preparation and motor imagery. Exp Brain Res 111:429–436CrossRefPubMedGoogle Scholar
  24. Decety J (1993) Analysis of actual and mental movement times in graphic tasks. Acta Psychol (Amst) 82:367–372CrossRefGoogle Scholar
  25. Decety J, Jeannerod M (1995) Mentally simulated movements in virtual reality: does Fitts’s law hold in motor imagery? Behav Brain Res 72:127–134CrossRefPubMedGoogle Scholar
  26. Decety J, Jeannerod M, Prablanc C (1989) The timing of mentally represented actions. Behav Brain Res 34:35–42CrossRefPubMedGoogle Scholar
  27. Dickstein R, Dunsky A, Marcovitz E (2004) Motor imagery for gait rehabilitation in post-stroke hemiparesis. Phys Ther 84:1167–1177PubMedGoogle Scholar
  28. Dunsky A, Dickstein R, Ariav C, Deutsch J, Marcovitz E (2006) Motor imagery practice in gait rehabilitation of chronic post-stroke hemiparesis: four case studies. Int J Rehabil Res 29:351–356CrossRefPubMedGoogle Scholar
  29. Duque J, Ivry RB (2009) Role of corticospinal suppression during motor preparation. Cereb Cortex 19:2013–2024CrossRefPubMedCentralPubMedGoogle Scholar
  30. Duque J, Lew D, Mazzocchio R, Olivier E, Ivry RB (2010) Evidence for two concurrent inhibitory mechanisms during response preparation. J Neurosci 30:3793–3802CrossRefPubMedCentralPubMedGoogle Scholar
  31. Fontani G, Migliorini S, Benocci R, Facchini A, Casini M, Corradeschi F (2007) Effect of mental imagery on the development of skilled motor actions. Percept Mot Skills 105:803–826PubMedGoogle Scholar
  32. Frank C, Land WM, Popp C, Schack T (2014) Mental representation and mental practice: experimental investigation on the functional links between motor memory and motor imagery. PLoS ONE 9:e95175CrossRefPubMedCentralPubMedGoogle Scholar
  33. Gahery Y, Ioffe ME, Massion J, Polit A (1981) Postural support for local movements in cats and dogs. Zh Vyssh Nerv Deiat Im I P Pavlova 31:232–241PubMedGoogle Scholar
  34. Gentili R, Papaxanthis C, Pozzo T (2006) Improvement and generalization of arm motor performance through motor imagery practice. Neuroscience 137:761–772CrossRefPubMedGoogle Scholar
  35. Gerardin E, Sirigu A, Lehericy S et al (2000) Partially overlapping neural networks for real and imagined hand movements. Cereb Cortex 10:1093–1104CrossRefPubMedGoogle Scholar
  36. Green JB, Bialy Y, Sora E, Thatcher RW (1997) An electroencephalographic study of imagined movement. Arch Phys Med Rehabil 78:578–581CrossRefPubMedGoogle Scholar
  37. Hanakawa T, Dimyan MA, Hallett M (2008) Motor planning, imagery, and execution in the distributed motor network: a time-course study with functional MRI. Cereb Cortex 18:2775–2788CrossRefPubMedCentralPubMedGoogle Scholar
  38. Hodges PW, Bui BH (1996) A comparison of computer-based methods for the determination of onset of muscle contraction using electromyography. Electroencephalogr Clin Neurophysiol 101:511–519PubMedGoogle Scholar
  39. Holmes P, Calmels C (2008) A neuroscientific review of imagery and observation use in sport. J Mot Behav 40:433–445CrossRefPubMedGoogle Scholar
  40. Hugon M, Massion J, Wiesendanger M (1982) Anticipatory postural changes induced by active unloading and comparison with passive unloading in man. Pflugers Arch 393:292–296CrossRefPubMedGoogle Scholar
  41. Ietswaart M, Johnston M, Dijkerman HC, Joice S, Scott CL, MacWalter RS, Hamilton SJ (2011) Mental practice with motor imagery in stroke recovery: randomized controlled trial of efficacy. Brain 134:1373–1386CrossRefPubMedCentralPubMedGoogle Scholar
  42. Jackson PL, Doyon J, Richards CL, Malouin F (2004) The efficacy of combined physical and mental practice in the learning of a foot-sequence task after stroke: a case report. Neurorehabil Neural Repair 18:106–111CrossRefPubMedGoogle Scholar
  43. Jankelowitz SK, Colebatch JG (2002) Movement-related potentials associated with self-paced, cued and imagined arm movements. Exp Brain Res 147:98–107CrossRefPubMedGoogle Scholar
  44. Jeannerod M (2001) Neural simulation of action: a unifying mechanism for motor cognition. Neuroimage 14:S103–S109CrossRefPubMedGoogle Scholar
  45. Kearns DW, Crossman J (1992) Effects of a cognitive intervention package on the free-throw performance of varsity basketball players during practice and competition. Percept Mot Skills 75:1243–1253CrossRefPubMedGoogle Scholar
  46. Kho AY, Liu KP, Chung RC (2014) Meta-analysis on the effect of mental imagery on motor recovery of the hemiplegic upper extremity function. Aust Occup Ther J 61:38–48CrossRefPubMedGoogle Scholar
  47. Kranczioch C, Mathews S, Dean PJ, Sterr A (2009) On the equivalence of executed and imagined movements: evidence from lateralized motor and nonmotor potentials. Hum Brain Mapp 30:3275–3286CrossRefPubMedGoogle Scholar
  48. Kranczioch C, Mathews S, Dean P, Sterr A (2010) Task complexity differentially affects executed and imagined movement preparation: evidence from movement-related potentials. PLoS ONE 5:e9284CrossRefPubMedCentralPubMedGoogle Scholar
  49. Kumru H, Valls-Solé J (2006) Excitability of the pathways mediating the startle reaction before execution of a voluntary movement. Exp Brain Res 169:427–432CrossRefPubMedGoogle Scholar
  50. la Fougere C, Zwergal A, Rominger A et al (2010) Real versus imagined locomotion: a [18F]-FDG PET-fMRI comparison. Neuroimage 50:1589–1598CrossRefPubMedGoogle Scholar
  51. Lotze M, Cohen LG (2006) Volition and imagery in neurorehabilitation. Cogn Behav Neurol 19:135–140CrossRefPubMedGoogle Scholar
  52. Lotze M, Montoya P, Erb M et al (1999) Activation of cortical and cerebellar motor areas during executed and imagined hand movements: an fMRI study. J Cogn Neurosci 11:491–501CrossRefPubMedGoogle Scholar
  53. MacKinnon CD, Bissig D, Chiusano J et al (2007) Preparation of anticipatory postural adjustments prior to stepping. J Neurophysiol 97:4368–4379CrossRefPubMedGoogle Scholar
  54. MacKinnon CD, Allen DP, Shiratori T, Rogers MW (2013) Early and unintentional release of planned motor actions during motor cortical preparation. PLoS ONE 8:e63417CrossRefPubMedCentralPubMedGoogle Scholar
  55. Malouin F, Richards CL, Jackson PL, Dumas F, Doyon J (2003) Brain activations during motor imagery of locomotor-related tasks: a PET study. Hum Brain Mapp 19:47–62CrossRefPubMedGoogle Scholar
  56. Maslovat D, Chua R, Hodges NJ (2013) When unintended movements “leak” out: a startling acoustic stimulus can elicit a prepared response during motor imagery and action observation. Neuropsychologia 51:838–844CrossRefPubMedGoogle Scholar
  57. Mason SG, Bashashati A, Fatourechi M, Navarro KF, Birch GE (2007) A comprehensive survey of brain interface technology designs. Ann Biomed Eng 35:137–169CrossRefPubMedGoogle Scholar
  58. Olsson CJ, Jonsson B, Nyberg L (2008) Internal imagery training in active high jumpers. Scand J Psychol 49:133–140CrossRefPubMedGoogle Scholar
  59. Rogers MW, Kennedy R, Palmer S et al (2011) Postural preparation prior to stepping in patients with Parkinson’s disease. J Neurophysiol 106:915–924CrossRefPubMedGoogle Scholar
  60. Roland PE, Larsen B, Lassen NA, Skinhoj E (1980) Supplementary motor area and other cortical areas in organization of voluntary movements in man. J Neurophysiol 43:118–136PubMedGoogle Scholar
  61. Schack T, Essig K, Frank C, Koester D (2014) Mental representation and motor imagery training. Front Hum Neurosci 8:328CrossRefPubMedCentralPubMedGoogle Scholar
  62. Sharma N, Pomeroy VM, Baron JC (2006) Motor imagery: a backdoor to the motor system after stroke? Stroke 37:1941–1952CrossRefPubMedGoogle Scholar
  63. Sirigu A, Duhamel JR (2001) Motor and visual imagery as two complementary but neurally dissociable mental processes. J Cogn Neurosci 13:910–919CrossRefPubMedGoogle Scholar
  64. Smith JL, Jamadar S, Provost AL, Michie PT (2013) Motor and non-motor inhibition in the Go/NoGo task: an ERP and fMRI study. Int J Psychophysiol 87:244–253CrossRefPubMedGoogle Scholar
  65. Snijders AH, Leunissen I, Bakker M, Overeem S, Helmich RC, Bloem BR, Toni I (2011) Gait-related cerebral alterations in patients with Parkinson’s disease with freezing of gait. Brain 134:59–72CrossRefPubMedGoogle Scholar
  66. Stephan KM, Fink GR, Passingham RE, Silbersweig D, Ceballos-Baumann AO, Frith CD, Frackowiak RS (1995) Functional anatomy of the mental representation of upper extremity movements in healthy subjects. J Neurophysiol 73:373–386PubMedGoogle Scholar
  67. Valls-Solé J, Rothwell JC, Goulart F, Cossu G, Munoz E (1999) Patterned ballistic movements triggered by a startle in healthy humans. J Physiol 516(Pt 3):931–938CrossRefPubMedCentralPubMedGoogle Scholar
  68. Valls-Solé J, Kumru H, Kofler M (2008) Interaction between startle and voluntary reactions in humans. Exp Brain Res 187:497–507CrossRefPubMedGoogle Scholar
  69. Walter WG, Cooper R, Aldridge VJ, McCallum WC, Winter AL (1964) Contingent negative variation: an electric sign of sensorimotor association and expectancy in the human brain. Nature 203:380–384CrossRefPubMedGoogle Scholar
  70. Wriessnegger SC, Steyrl D, Koschutnig K, Muller-Putz GR (2014) Short time sports exercise boosts motor imagery patterns: implications of mental practice in rehabilitation programs. Front Hum Neurosci 8:469CrossRefPubMedCentralPubMedGoogle Scholar

Copyright information

© Springer-Verlag Berlin Heidelberg 2015

Authors and Affiliations

  • Jeremy S. Eagles
    • 1
  • Anthony N. Carlsen
    • 1
    • 2
  • Colum D. MacKinnon
    • 1
    • 3
    Email author
  1. 1.Department of Physical Therapy and Human Movement Sciences, Feinberg School of MedicineNorthwestern UniversityChicagoUSA
  2. 2.School of Human Kinetics, Faculty of Health SciencesUniversity of OttawaOttawaCanada
  3. 3.Department of NeurologyUniversity of MinnesotaMinneapolisUSA

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