Skip to main content
Log in

A behavioural analysis of the temporal organisation of walking movements in the 1st instar and adult stick insect (Carausius morosus)

  • Published:
Journal of comparative physiology Aims and scope Submit manuscript


The coordination of protraction leg movements in free walking first instar and adult stick insects has been examined by film and video-tape analysis. The first instar insect uses two alternative modes of walking which differ in the step frequency dependence of the time required to complete one metachronal cycle of protractions and the duration of the protraction cycle (Figs. 2, 4). The adult uses only one of these modes and its step patterns show a persistent right-left asymmetry in the timing of leg movements (Fig. 7). Two alternative methods of changing direction were found (Figs. 5, 9) and they appear to be related to the two modes of walking used by the animal. A model based on inhibitory interactions between oscillators representing leg activity has been constructed to explain the step pattern changes in both modes of walking and the temporal asymmetry of the adult pattern. This model appears to be applicable to the walking behaviour of other insects including the cockroach.

This is a preview of subscription content, log in via an institution to check access.

Access this article

Price excludes VAT (USA)
Tax calculation will be finalised during checkout.

Instant access to the full article PDF.

Similar content being viewed by others


  • Buddenbrock, W. V.: Der Rhythmus der Schreitbewegungen der Stabheuschrecke Dixippus. Biol. Zbl.41, 41–48 (1921).

    Google Scholar 

  • Davis, W. J.: Reflex organisation in the swimmeret system of the lobster. I Intrasegmental reflexes. J. exp. Biol51, 547–563 (1969).

    Google Scholar 

  • Delcomyn, F.: Reflexes and locomotion in the American cockroach. Ph. D. Thesis. University of Oregon (1969).

  • Delcomyn, F.: The locomotion of the cockroachPeriplaneta americana. J. exp. Biol.54, 453–469 (1971a).

    Google Scholar 

  • Delcomyn, F.: The effect of limb amputation on locomotion in the cockroachPeriplaneta americana. J. exp. Biol.54, 453–469 (1971b).

    Google Scholar 

  • Holst, E. von: Die Koordination der Bewegung bei den Arthropoden in Abhängigkeit von zentralen and peripheren Bedingungen. Biol. Rev.10, 234–261 (1935).

    Google Scholar 

  • Hoyle, G.: Exploration of neuronal mechanisms underlying behaviour in insects. Neural theory and modelling, p. 346–376. Reiss, R. Stanford University Press 1964.

    Google Scholar 

  • Hughes, G. M.: The coordination of insect movements. I. The walking movements of insects (Blatta orientalis). J. exp. Biol.29, 267–284 (1952).

    Google Scholar 

  • Pearson, K. G., Iies, J. F.: Discharge patterns of coxal levator and depressor motor neurons of the cockroach,Periplaneta americana. J. exp. Biol.52, 139–165 (1970).

    Google Scholar 

  • Roeder, K. D.: The control of tonus and locomotor activity in the praying mantis (Mantis religiosa). J. exp. Zool.76, 353–375 (1937).

    Google Scholar 

  • Usherwood, P. N. R., Runion, H. I., Campbell, J. I.: Structure and physiology of a chordotonal organ in the locust leg. J. exp. Biol.48, 305–323 (1968).

    Google Scholar 

  • Wendler, G.: Laufen und Stehen der StabheuschreckeCarausius morosus. Z. vergl. Physiol.48, 198–250 (1964a).

    Google Scholar 

  • Wendler, G.: Über die Fortbewegung der Larven vonCantharis fusca. Z. vergl. Physiol.48, 283–294 (1964b).

    Google Scholar 

  • Wendler, G.: Ein Analogmodell der Beinbewegungen eines laufenden Insekts. ‚Kybernetik 1968’, Beih. Elektronische Rechenanlagen18, 68–74. München: Oldenbourg 1968.

    Google Scholar 

  • Wilson, D. M.: Proprioceptive leg reflexes in cockroaches. J. exp. Biol.43, 379–409 (1965).

    Google Scholar 

  • Wilson, D. M.: Insect walking. Ann. Rev. Entomol.11, 103–122 (1966).

    Google Scholar 

  • Wilson, D. M.: Stepping patterns in Tarantula spiders. J. exp. Biol.47, 133–151 (1968a).

    Google Scholar 

  • Wilson, D. M.: Inherant asymmetry and reflex modulation of the locust flight motor pattern. J. exp. Biol.48, 631–641 (1968b).

    Google Scholar 

Download references

Author information

Authors and Affiliations


Additional information

This work was supported by a Public Health Service grant NB-06054 to Dr. R. K. Josephson and a fellowship F03-GM-42337 to the author.

I would like to thank Drs. P. N. R. Usherwood, F. Delcomyn, C. Walther, M. D. Burns and W. Winlow for their criticism of earlier drafts of this manuscript and I am particularly indebted to Dr. R. K. Josephson for his encouragement in this project.

Rights and permissions

Reprints and permissions

About this article

Cite this article

Graham, D. A behavioural analysis of the temporal organisation of walking movements in the 1st instar and adult stick insect (Carausius morosus). J. Comp. Physiol. 81, 23–52 (1972).

Download citation

  • Received:

  • Issue Date:

  • DOI: