Object Avoidance During Locomotion

  • David A. McVea
  • Keir G. Pearson
Part of the Advances in Experimental Medicine and Biology book series (AEMB, volume 629)


Many animals rely on vision for navigating through complex environments and for avoiding specific obstacles during locomotion. Navigation and obstacle avoidance are tasks that depend on gathering information about the environment by vision and using this information at later times to guide limb and body movements. Here we review studies demonstrating the use of short-term visual memory during walking in humans and cats. Our own investigations have demonstrated that cats have the ability to retain a memory of an obstacle they have stepped over with the forelegs for many minutes and to use this memory to guide stepping of the hindlegs to avoid the remembered obstacle. A brain region that may be critically involved in the retention of memories of the location of obstacles is the posterior parietal cortex. Recordings from neurons in area 5 in the posterior parietal cortex in freely walking cats have revealed the existence of neurons whose activity is strongly correlated with the location of an obstacle relative to the body. How these neurons might be used to regulate motor commands remains to be established. We believe that studies on obstacle avoidance in walking cats have the potential to significantly advance our understanding of visuo-motor transformations. Current knowledge about the brain regions and pathways underlying visuo-motor transformations during walking are reviewed.


Visual Information Motor Cortex Visual Feedback Optic Flow Visual Input 
These keywords were added by machine and not by the authors. This process is experimental and the keywords may be updated as the learning algorithm improves.


  1. Adkins RJ, Cegnar MR and Rafuse DD. Differential effects of lesions of the anterior and posterior sigmoid gyri in cats. Brain Res. 30(2): 411–414, 1971.PubMedCrossRefGoogle Scholar
  2. Andersen RA and Buneo CA. Sensorimotor integration in posterior parietal cortex. Adv. Neurol. 93: 159–177, 2003.PubMedGoogle Scholar
  3. Andersen RA and Buneo CA. Intentional maps in posterior parietal cortex. Annu. Rev. Neurosci. 25: 189–220, 2002.PubMedCrossRefGoogle Scholar
  4. Andujar JE and Drew T. Organization of posterior parietal projections to the forelimb and hindlimb representations in the motor cortex of the cat: A retrograde tracer study. Soc. Neurosci. Abstr. 288.13, 2005.Google Scholar
  5. Armstrong DM and Drew T. Forelimb electromyographic responses to motor cortex stimulation during locomotion in the cat. J. Physiol. 367: 327–351, 1985.PubMedGoogle Scholar
  6. Armstrong DM and Drew T. Discharges of pyramidal tract and other motor cortical neurones during locomotion in the cat. J. Physiol. 346: 471–495, 1984.PubMedGoogle Scholar
  7. Armstrong DM and Marple-Horvat DE. Role of the cerebellum and motor cortex in the regulation of visually controlled locomotion. Can. J. Physiol. Pharmacol. 74(4): 443–455, 1996.PubMedCrossRefGoogle Scholar
  8. Assaiante C, Marchand AR and Amblard B. Discrete visual samples may control locomotor equilibrium and foot positioning in man. J. Mot. Behav. 21(1): 72–91, 1989.PubMedGoogle Scholar
  9. Avendano C, Rausell E, Perez-Aguilar D and Isorna S. Organization of the association cortical afferent connections of area 5: A retrograde tracer study in the cat. J. Comp. Neurol. 278(1): 1–33, 1988.PubMedCrossRefGoogle Scholar
  10. Avendano C, Rausell E and Reinoso-Suarez F. Thalamic projections to areas 5a and 5b of the parietal cortex in the cat: A retrograde horseradish peroxidase study. J. Neurosci. 5(6): 1446–1470, 1985.PubMedGoogle Scholar
  11. Babb RS, Waters RS and Asanuma H. Corticocortical connections to the motor cortex from the posterior parietal lobe (areas 5a, 5b, 7) in the cat demonstrated by the retrograde axonal transport of horseradish peroxidase. Exp. Brain Res. 54(3): 476–484, 1984.PubMedCrossRefGoogle Scholar
  12. Beloozerova IN and Sirota MG. Role of motor cortex in control of locomotion. In: Stance and Motion: Facts and Concepts, edited by Gurfinkel VS, Ioffe ME, Massion J and Roll JP. New York: Prenum Press, 1986, pp. 163–176.Google Scholar
  13. Beloozerova IN and Sirota MG. Integration of motor and visual information in the parietal area 5 during locomotion. J. Neurophysiol. 90(2): 961–971, 2003.PubMedCrossRefGoogle Scholar
  14. Beloozerova IN and Sirota MG. The role of the motor cortex in the control of accuracy of locomotor movements in the cat. J. Physiol. 461: 1–25, 1993.PubMedGoogle Scholar
  15. Bretzner F and Drew T. Motor cortical modulation of cutaneous reflex responses in the hindlimb of the intact cat. J. Neurophysiol. 94(1): 673–687, 2005.PubMedCrossRefGoogle Scholar
  16. Brustein E and Rossignol S. Recovery of locomotion after ventral and ventrolateral spinal lesions in the cat. I. Deficits and adaptive mechanisms. J. Neurophysiol. 80(3): 1245–1267, 1998.PubMedGoogle Scholar
  17. Buneo CA and Andersen RA. The posterior parietal cortex: Sensorimotor interface for the planning and online control of visually guided movements. Neuropsychologia 44(13): 2594–2606, 2005.Google Scholar
  18. Capaday C. Neurophysiological methods for studies of the motor system in freely moving human subjects. J. Neurosci. Methods 74(2): 201–218, 1997.PubMedCrossRefGoogle Scholar
  19. Colby CL and Goldberg ME. Space and attention in parietal cortex. Annu. Rev. Neurosci. 22: 319–349, 1999.PubMedCrossRefGoogle Scholar
  20. Dietz V and Duysens J. Significance of load receptor input during locomotion: A review. Gait Posture 11(2): 102–110, 2000.PubMedCrossRefGoogle Scholar
  21. Donelan JM and Pearson KG. Contribution of sensory feedback to ongoing ankle extensor activity during the stance phase of walking. Can. J. Physiol. Pharmacol. 82(8–9): 589–598, 2004.PubMedCrossRefGoogle Scholar
  22. Drew T. Motor cortical cell discharge during voluntary gait modification. Brain Res. 457:181–187, 1988.PubMedCrossRefGoogle Scholar
  23. Drew T. Motor cortical activity during voluntary gait modifications in the cat. I. Cells related to the forelimbs. J. Neurophysiol. 70(1): 179–199, 1993.PubMedGoogle Scholar
  24. ∗Drew T. Visuomotor coordination in locomotion. Curr. Opin. Neurobiol. 1(4): 652–657, 1991.PubMedCrossRefGoogle Scholar
  25. Drew T, Dubuc R and Rossignol S. Discharge patterns of reticulospinal and other reticular neurons in chronic, unrestrained cats walking on a treadmill. J. Neurophysiol. 55(2): 375–401, 1986.PubMedGoogle Scholar
  26. Drew T, Jiang W, Kably B and Lavoie S. Role of the motor cortex in the control of visually triggered gait modifications. Can. J. Physiol. Pharmacol. 74(4): 426–442, 1996.PubMedCrossRefGoogle Scholar
  27. ∗Drew T, Jiang W and Widajewicz W. Contributions of the motor cortex to the control of the hindlimbs during locomotion in the cat. Brain Res. Brain Res. Rev. 40(1–3): 178–191, 2002.PubMedCrossRefGoogle Scholar
  28. Drew T, Prentice S and Schepens B. Cortical and brainstem control of locomotion. Prog. Brain Res. 143: 251–261, 2004.PubMedCrossRefGoogle Scholar
  29. Drew T and Rossignol S. Phase-dependent responses evoked in limb muscles by stimulation of medullary reticular formation during locomotion in thalamic cats. J. Neurophysiol. 52(4): 653–675, 1984.PubMedGoogle Scholar
  30. Earhart GM, Horak FB, Jones GM, Block EW, Weber KD, Suchowersky O and Fletcher WA. Is the cerebellum important for podokinetic adaptation? Ann. NY. Acad. Sci. 978: 511–512, 2002.PubMedCrossRefGoogle Scholar
  31. Elliott D. Continuous visual information may be important after all: A failure to replicate Thomson (1983). J. Exp. Psychol. Hum. Percept. Perform. 12(3): 388–391, 1986.PubMedCrossRefGoogle Scholar
  32. Fabre M and Buser P. Effects of lesioning the anterior suprasylvian cortex on visuo-motor guidance performance in the cat. Exp. Brain Res. 41(2): 81–88, 1981.PubMedCrossRefGoogle Scholar
  33. Fiset S and Dore FY. Duration of cats' (Felis catus) working memory for disappearing objects. Anim. Cogn. 31: 1–9, 2005.Google Scholar
  34. Fowler GA and Sherk H. Gaze during visually-guided locomotion in cats. Behav. Brain Res. 139(1–2): 83–96, 2003.PubMedCrossRefGoogle Scholar
  35. Georgopoulos AP and Ashe J. One motor cortex, two different views. Nat. Neurosci. 3(10): 963; author reply 964–965, 2000.CrossRefGoogle Scholar
  36. Ghosh S. Comparison of the cortical connections of areas 4 gamma and 4 delta in the cat cerebral cortex. J. Comp. Neurol. 388(3): 371–396, 1997.PubMedCrossRefGoogle Scholar
  37. Gibson JJ. Visually controlled locomotion and visual orientation in animals. Br. J. Psychol. 49(3): 182–194, 1958.PubMedGoogle Scholar
  38. Graziano MSA and Botvinick MM. How the brain represents the body: Insights from neurophysiology and psychology. Attention Perform 19: 136–157, 2002.Google Scholar
  39. Graziano MS, Cooke DF and Taylor CS. Coding the location of the arm by sight. Science 290(5497): 1782–1786, 2000.PubMedCrossRefGoogle Scholar
  40. Grillner S. Possible analogies in the control of innate motor acts and the production of sound in speech. In: Speech Motor Control, edited by Grillner S. Oxford: Pergamon Press, 1982, pp. 217–229.Google Scholar
  41. Hollands MA, Marple-Horvat DE, Henkes S and Rowan AK. Human Eye Movements During Visually Guided Stepping. J. Mot. Behav. 27(2): 155–163, 1995.PubMedCrossRefGoogle Scholar
  42. Jiang W and Drew T. Effects of bilateral lesions of the dorsolateral funiculi and dorsal columns at the level of the low thoracic spinal cord on the control of locomotion in the adult cat. I. Treadmill walking. J. Neurophysiol. 76(2): 849–866, 1996.PubMedGoogle Scholar
  43. Kably B and Drew T. Corticoreticular pathways in the cat. II. Discharge activity of neurons in area 4 during voluntary gait modifications. J. Neurophysiol. 80(1): 406–424, 1998.PubMedGoogle Scholar
  44. Kakei S, Futami T and Shinoda Y. Projection pattern of single corticocortical fibers from the parietal cortex to the motor cortex. Neuroreport 7(14): 2369–2372, 1996.PubMedCrossRefGoogle Scholar
  45. Kakei S and Shinoda Y. Parietal projection of thalamocortical fibers from the ventroanterior-ventrolateral complex of the cat thalamus. Neurosci. Lett. 117(3): 280–284, 1990.PubMedCrossRefGoogle Scholar
  46. Kakei S, Yagi J, Wannier T, Na J and Shinoda Y. Cerebellar and cerebral inputs to corticocortical and corticofugal neurons in areas 5 and 7 in the cat. J. Neurophysiol. 74(1): 400–412, 1995.PubMedGoogle Scholar
  47. ∗Kalaska JF. Parietal cortex area 5 and visuomotor behavior. Can. J. Physiol. Pharmacol. 74(4): 483–498, 1996.PubMedCrossRefGoogle Scholar
  48. Kalaska JF and Crammond DJ. Deciding not to GO: Neuronal correlates of response selection in a GO/NOGO task in primate premotor and parietal cortex. Cereb. Cortex 5(5): 410–428, 1995.PubMedCrossRefGoogle Scholar
  49. Kalaska JF and Drew T. Motor cortex and visuomotor behavior. Exerc. Sport Sci. Rev. 21: 397–436, 1993.PubMedCrossRefGoogle Scholar
  50. Kalaska JF, Scott SH, Cisek P and Sergio LE. Cortical control of reaching movements. Curr. Opin. Neurobiol. 7(6): 849–859, 1997.PubMedCrossRefGoogle Scholar
  51. Kawamura K. Corticortical fiber connections of the cat cerebrum. II. The parietal region. Brain Res. 51: 23–40, 1973.PubMedCrossRefGoogle Scholar
  52. Lacquaniti F, Guigon E, Bianchi L, Ferraina S and Caminiti R. Representing spatial information for limb movement: Role of area 5 in the monkey. Cereb. Cortex 5(5): 391–409, 1995.PubMedCrossRefGoogle Scholar
  53. Lajoie K and Drew T. The contribution of the posterior parietal cortex to the control of visually guided locomotion in the cat: A lesion study. Soc. Neurosci. Abstr. 287.23, 2005.Google Scholar
  54. Lajoie K and Drew T. Lesions in area 5 of the posterior parietal cortex in the cat produce errors in the accuracy of paw placement during visually-guided locomotion. J. Neurophysiol. 97: 2339–2354, 2007.PubMedCrossRefGoogle Scholar
  55. Lajoie K, Andujar J, Pearson KG and Drew T. Persistent neuronal activity in posterior parietal cortex area 5 related to long-lasting memories of obstacles in walking cats. Soc. Neurosci. Abst. 37: 397.8, 2007.Google Scholar
  56. Lavoie S, McFadyen B and Drew T. A kinematic and kinetic analysis of locomotion during voluntary gait modification in the cat. Exp. Brain Res. 106(1): 39–56, 1995.PubMedCrossRefGoogle Scholar
  57. Liddell EGT and Phillips CG. Pyramidal section in the cat. Brain 67: 1–9, 1944.CrossRefGoogle Scholar
  58. Marple-Horvat DE and Criado JM. Rhythmic neuronal activity in the lateral cerebellum of the cat during visually guided stepping. J. Physiol. 518(Pt 2): 595–603, 1999.PubMedCrossRefGoogle Scholar
  59. Marple-Horvat DE, Criado JM and Armstrong DM. Neuronal activity in the lateral cerebellum of the cat related to visual stimuli at rest, visually guided step modification, and saccadic eye movements. J. Physiol. 506 ( Pt 2): Pt 2: 489–514, 1998.Google Scholar
  60. ∗McVea DA and Pearson KG. Long-lasting memories of obstacles guide leg movements in the walking cat. J. Neurosci. 26(4): 1175–1178, 2006.PubMedCrossRefGoogle Scholar
  61. McVea DA and Pearson KG. Stepping of the forelegs over obstacles establishes long-lasting memories in cats. Curr. Biol. 17: R621–623, 2007.Google Scholar
  62. Mori S, Matsuyama K, Kohyama J, Kobayashi Y and Takakusaki K. Neuronal constituents of postural and locomotor control systems and their interactions in cats. Brain Dev. 14 Suppl: S109–20, 1992.PubMedGoogle Scholar
  63. Morton SM, Dordevic GS and Bastian AJ. Cerebellar damage produces context-dependent deficits in control of leg dynamics during obstacle avoidance. Exp. Brain Res. 156(2): 149–163, 2004.PubMedCrossRefGoogle Scholar
  64. Patla AE. How is human gait controlled by vision? Ecol Psychol 10(3–4): 287–302, 1998.CrossRefGoogle Scholar
  65. ∗Patla AE. Understanding the roles of vision in the control of human locomotion. Gait Posture 5: 54–69, 1997.CrossRefGoogle Scholar
  66. Patla AE, Adkin A, Martin C, Holden R and Prentice S. Characteristics of voluntary visual sampling of the environment for safe locomotion over different terrains. Exp. Brain Res. 112(3): 513–522, 1996.PubMedCrossRefGoogle Scholar
  67. Patla AE and Greig M. Any way you look at it, successful obstacle negotiation needs visually guided on-line foot placement regulation during the approach phase. Neurosci. Lett. 397(1–2): 110–114, 2006.PubMedCrossRefGoogle Scholar
  68. Patla AE, Prentice SD, Robinson C and Neufeld J. Visual control of locomotion: Strategies for changing direction and for going over obstacles. J. Exp. Psychol. Hum. Percept. Perform. 17(3): 603–634, 1991.PubMedCrossRefGoogle Scholar
  69. Patla AE and Vickers JN. How far ahead do we look when required to step on specific locations in the travel path during locomotion? Exp. Brain Res. 148(1): 133–138, 2003.PubMedCrossRefGoogle Scholar
  70. Patla AE and Vickers JN. Where and when do we look as we approach and step over an obstacle in the travel path? Neuroreport 8(17): 3661–3665, 1997.PubMedCrossRefGoogle Scholar
  71. Prentice SD and Drew T. Contributions of the reticulospinal system to the postural adjustments occurring during voluntary gait modifications. J. Neurophysiol. 85(2): 679–698, 2001.PubMedGoogle Scholar
  72. Rauschecker JP, von Grunau MW and Poulin C. Centrifugal organization of direction preferences in the cat's lateral suprasylvian visual cortex and its relation to flow field processing. J. Neurosci. 7(4): 943–958, 1987.PubMedGoogle Scholar
  73. Robinson FR, Cohen JL, May J, Sestokas AK and Glickstein M. Cerebellar targets of visual pontine cells in the cat. J. Comp. Neurol. 223(4): 471–482, 1984.PubMedCrossRefGoogle Scholar
  74. Rossignol S. Locomotion and its recovery after spinal injury in animal models. Neurorehabil. Neural Repair 16(2): 201–206, 2002.PubMedCrossRefGoogle Scholar
  75. Rossignol S and Dubuc R. Spinal pattern generation. Curr. Opin. Neurobiol. 4(6): 894–902, 1994.PubMedCrossRefGoogle Scholar
  76. ∗Rossignol S, Dubuc R and Gossard JP. Dynamic sensorimotor interactions in locomotion. Physiol. Rev. 86(1): 89–154, 2006.PubMedCrossRefGoogle Scholar
  77. Sasaki K, Matsuda Y, Kawaguchi S and Mizuno N. On the cerebello-thalamo-cerebral pathway for the parietal cortex. Exp. Brain Res. 16(1): 89–103, 1972.PubMedGoogle Scholar
  78. Scott SH. The role of primary motor cortex in goal-directed movements: Insights from neurophysiological studies on non-human primates. Curr. Opin. Neurobiol. 13(6): 671–677, 2003.PubMedCrossRefGoogle Scholar
  79. Scott SH. Role of motor cortex in coordinating multi-joint movements: Is it time for a new paradigm? Can. J. Physiol. Pharmacol. 78(11): 923–933, 2000.PubMedCrossRefGoogle Scholar
  80. Scott SH, Sergio LE and Kalaska JF. Reaching movements with similar hand paths but different arm orientations. II. Activity of individual cells in dorsal premotor cortex and parietal area 5. J. Neurophysiol. 78(5): 2413–2426, 1997.PubMedGoogle Scholar
  81. Sherk H and Fowler GA. Lesions of extrastriate cortex and consequences for visual guidance during locomotion. Exp. Brain Res. 144(2): 159–171, 2002.PubMedCrossRefGoogle Scholar
  82. Sherk H and Fowler GA. Neural analysis of visual information during locomotion. Prog. Brain Res. 134: 247–264, 2001a.CrossRefGoogle Scholar
  83. Sherk H and Fowler GA. Visual analysis and image motion in locomoting cats. Eur. J. Neurosci. 13(6): 1239–1248, 2001b.CrossRefGoogle Scholar
  84. Steenhuis RE and Goodale MA. The effects of time and distance on accuracy of target-directed locomotion: Does an accurate short-term memory for spatial location exist? J. Mot. Behav. 20(4): 399–415, 1988.PubMedGoogle Scholar
  85. Symonds LL, Rosenquist AC, Edwards SB and Palmer LA. Projections of the pulvinar-lateral posterior complex to visual cortical areas in the cat. Neuroscience 6(10): 1995–2020, 1981.PubMedCrossRefGoogle Scholar
  86. Thomson JA. Is continuous visual monitoring necessary in visually guided locomotion? J. Exp. Psychol. Hum. Percept. Perform. 9(3): 427–443, 1983.PubMedCrossRefGoogle Scholar
  87. Ung RV, Imbeault MA, Ethier C, Brizzi L and Capaday C. On the potential role of the corticospinal tract in the control and progressive adaptation of the soleus h-reflex during backward walking. J. Neurophysiol. 94(2): 1133–1142, 2005.PubMedCrossRefGoogle Scholar
  88. Whelan PJ. Control of locomotion in the decerebrate cat. Prog. Neurobiol. 49(5): 481–515, 1996.PubMedCrossRefGoogle Scholar
  89. Widajewicz W, Kably B and Drew T. Motor cortical activity during voluntary gait modifications in the cat. II. Cells related to the hindlimbs. J. Neurophysiol. 72(5): 2070–2089, 1994.PubMedGoogle Scholar
  90. Wilkinson EJ and Sherk HA. The use of visual information for planning accurate steps in a cluttered environment. Behav. Brain Res. 164(2): 270–274, 2005.PubMedCrossRefGoogle Scholar
  91. The references marked with an asterisk (∗) are specifically recommended for further introduction or background to the topic.Google Scholar

Copyright information

© Springer Science+Business Media, LLC 2009

Authors and Affiliations

  1. 1.Department of Physiology and Centre for NeuroscienceUniversity of AlbertaEdmonton ABCanada

Personalised recommendations