Journal of Comparative Physiology A

, Volume 156, Issue 2, pp 153–164 | Cite as

Spinal reflex pattern in the tortoise (Testudo graeca, T. hermanni)

  • H. Steffens
  • E. D. Schomburg
  • W. J. Koehler


  1. 1.

    In high spinal anaemically decapitated tortoises (Testudo graeca andT. hermanni) lumbar reflex pathways were investigated. The responses to peripheral stimulation of identified ipsi- and contralateral hindlimb nerves were recorded intracellularly in lumbar motoneurones. Early, presumably monosynaptic, and later polysynaptic responses were observed.

  2. 2.

    Presumed monosynaptic projections from low threshold afferents to motoneurones were found not only in homonymous motoneurones but also in heteronymous motoneurones.

  3. 3.

    The polysynaptic responses in a given motoneurone to stimulation of ipsilateral cutaneous, mixed and muscle nerves generally were uniform, i.e. the responses were almost always in the same direction independent of the stimulated nerve. I.p.s.p.s were clearly dominating in number and often also in size. A response pattern corresponding to the flexion reflex, i.e. excitation of flexor motoneurones and inhibition of extensor motoneurones could occur, but other patterns were frequently observed.

  4. 4.

    Responses to stimulation of contralateral nerves were in most cases in the same direction as those to stimulation of ipsilateral nerves but were generally smaller, i.e., the reflex pathways from both sides evoked qualitatively the same but quantitatively different effects. Reciprocal effects corresponding to the pattern of the contralateral crossed extension reflex occurred, but were the exception.

  5. 5.

    The spinal reflex pattern found in tortoise in many aspects resembles that one of higher vertebrates, but there are specializations found according to the phylogenetic position and to the behaviour of the tortoise.



High Spinal Response Pattern Phylogenetic Position Reciprocal Effect Muscle Nerve 
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.


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  1. Änggård L, Bergström R, Bernhard CG (1961) Analysis of prenatal spinal reflex activity in sheep. Acta Physiol Scand 53:128–136Google Scholar
  2. Baldissera F, Hultborn H, Illert M (1981) Integration in spinal neuronal systems. In: Brooks VB (ed) Motor control. Am Physiol Soc, Bethesda (Handbook of physiology, sect 1, vol II, pp 509–595)Google Scholar
  3. Bergström RM, Hellström P-E, Stenberg D (1962) Studies in reflex irradiations in the foetal guinea-pig. Ann Chir Gynaecol 51:171–178Google Scholar
  4. Bojanus LH (1819–1821) Anatome Testudinis Europaeae. Vilnae Typographi UniversitatisGoogle Scholar
  5. Crowe A, Valk-Fai T (1979) The mechanism of spinal control of reflex hind limb movements in the terrapin Pseudemys scripta elegans. J Physiol (Lond) 292:19PGoogle Scholar
  6. Eccles RM, Lundberg A (1959a) Synaptic actions in motoneurones by afferents which may evoke the flexion reflex. Arch Ital Biol 97:199–221Google Scholar
  7. Eccles RM, Lundberg A (1959b) Supraspinal control of interneurones mediating spinal reflexes. J Physiol (Lond) 147:565–584Google Scholar
  8. Eccles JC, Eccles RM, Lundberg A (1957) The convergence of monosynaptic excitatory afferents onto many different species of alpha motoneurones. J Physiol (Lond) 137:22–50Google Scholar
  9. Feenstra BWA, Hofman F, Van Leeuwen JJ (1984) Syntheses of spinal cord field potentials in the terrapin. Biol Cybern 50:409–418Google Scholar
  10. Fritz N (1981) Ia-Synergismus an der vorderen Extremität der Katze. Dissertation, Fakultät für Biologie, Ludwig-Maximilians-Universität MünchenGoogle Scholar
  11. Holmquist B, Lundberg A (1961) Differential supraspinal control of synaptic actions evoked by volleys in the flexion reflex afferents in alpha motoneurones. Acta Physiol Scand 54 (Suppl 186):1–51Google Scholar
  12. Kenins P (1977) Reflex response to stretch of limb muscles in the Australian blue tongue lizard. Comp Biochem Physiol 57 A: 383–390Google Scholar
  13. Kirkwood PA, Sears TA (1974) Monosynaptic excitation of motoneurones from secondary endings of muscle spindles. Nature 252:243–244Google Scholar
  14. Kniffki K-D, Schomburg ED, Steffens H (1981) Synaptic effects from chemically activated fine muscle afferents uponα-motoneurones in decerebrate and spinal cats. Brain Res 206:361–370Google Scholar
  15. Kusuma A (1979) The organization of the spinal cord in reptiles with different locomotor patterns. Proefschrift ter Verkrijging van de Graad van Doctor in de Geneeskunde aan de Katholieke Universiteit te Nijmegen. Stichting Studentenpers NijmegenGoogle Scholar
  16. Kusuma A, Donkelaar HJ ten (1979) Staining of the dorsal root primary afferent fibers by anterograde movement of horseradish peroxidase and retrograde labelling of motoneurons and preganglionic autonomic cells in the turtle spinal cord. Neurosci Lett 14:141–146Google Scholar
  17. Kusuma A, Donkelaar HJ ten (1980) Dorsal root projections in various types of reptiles. Brain Behav Evol 17:291–309Google Scholar
  18. Kusuma A, Donkelaar HJ ten, Nieuwenhuys R (1979) Intrinsic organization of the spinal cord. In: Gans C, Northcutt RG, Ulinski P (eds) Biology of the reptilia, vol 10, Neurology B. Academic Press London pp 59–109Google Scholar
  19. Lloyd DPC (1943) Neuron patterns controlling transmission of ipsilateral hind limb reflexes in cat. J Neurophysiol 6:293–315Google Scholar
  20. Lundberg A, Voorhoeve P (1962) Effects from the pyramidal tract on spinal reflex arcs. Acta Physiol Scand 56:201–219Google Scholar
  21. Lundberg A, Winsbury G (1960) Selective adequate activation of large afferents from muscle spindles and Golgi tendon organs. Acta Physiol Scand 49:155–164Google Scholar
  22. Lundberg A, Malmgren K, Schomburg ED (1977) Comments on reflex actions evoked by electrical stimulation of group II muscle afferents. Brain Res 122:551–555Google Scholar
  23. Nieuwenhuys R (1964) Comparative anatomy of the spinal cord. Prog Brain Res 11:1–57Google Scholar
  24. Perl ER (1957) Crossed reflexes of cutaneous origin. Am J Physiol 188:609–615Google Scholar
  25. Rosenberg ME (1972) Excitation and inhibition of motoneurones in the tortoise. J Physiol (Lond) 221:715–730Google Scholar
  26. Rosenberg ME (1974) The distribution of the sensory input in the dorsal spinal cord of the tortoise. J Comp Neurol 156:29–38Google Scholar
  27. Rosenberg ME (1977) Conduction in the dorsal white columns of tortoise. Comp Biochem Physiol A 56:487–494Google Scholar
  28. Rosenberg ME (1978) Thermal relations of nervous conduction in the tortoise. Comp Biochem Physiol A 60:57–63Google Scholar
  29. Rosenberg ME (1980) Central responses to mechanical and electrical stimulation of the carapace in the tortoise. Comp Biochem Physiol A 66:227–231Google Scholar
  30. Ruigrok TJH, Crowe A, Donkelaar HJ ten (1982) The distribution of motoneurones innervating hindlimb muscles in the terrapinPseudemys scripta elegans. Neurosci Lett 28:157–162Google Scholar
  31. Schmidt RF, Willis WD (1963) Intracellular recording from motoneurons of the cervical spinal cord of the cat. J Neurophysiol 26:28–43Google Scholar
  32. Schomburg ED, Behrends HB (1978) Phasic control of the transmission in the excitatory and inhibitory reflex pathways from cutaneous afferents toα-motoneurones during fictive locomotion in cats. Neurosci Lett 8:277–282Google Scholar
  33. Schomburg ED, Roesler J, Meinck H-M (1977) Phase-dependent transmission in the excitatory propriospinal reflex pathway from forelimb afferents to lumbar motoneurones during fictive locomotion. Neurosci Lett 5:249–252Google Scholar
  34. Schomburg ED, Behrends HB, Steffens H (1981) Changes in segmental and propriospinal reflex pathways during spinal locomotion. In: Taylor A, Prochazka A (eds) Muscle receptors and movement. Macmillan, London, pp 413–425Google Scholar
  35. Sherrington CS (1910) Flexion-reflex of the limb, crossed extension reflex, and reflex stepping and standing. J Physiol (Lond) 40:28–121Google Scholar
  36. Steffens H, Schomburg ED, Behrends HB (1978) Segmental reflex pathways from cutaneous afferents toα-motoneurones in the tortoise (Testudo graeca andT. hermanni). Neurosci Lett Suppl 1:104Google Scholar
  37. Steffens H, Schomburg ED, Koehler WJ (1979) Zur Frage des gekreuzten Streckreflexes bei Landschildkröten (Testudo graeca, T. hermanni). Verh Dtsch Zool Ges 307Google Scholar
  38. Stein PSG (1978) Swimming movements elicited by electrical stimulation of the turtle spinal cord: The high spinal preparation. J Comp Physiol 124:203–210Google Scholar
  39. Stein PSG (1980) Neural control of locomotion in the turtle. Proc Int Union Physiol Sci, XXVIII Int Congress, vol XIV. Budapest, p 252Google Scholar
  40. Stein PSG, Grossman ML (1980) Central program for scratch reflex in turtle. J Comp Physiol 140:287–294Google Scholar
  41. Stein PSG, Robertson GA, Keifer J, Grossman ML, Berenbein JA, Lennard PR (1982) Motor neuron synaptic potentials during fictive scratch reflex in turtle. J Comp Physiol 146:401–409Google Scholar
  42. Ten Cate J (1937) Reaktionen der Schildkröte vor und nach der Durchtrennung des Rückenmarks. Arch Néerl Physiol 22:76–83Google Scholar
  43. Valk-Fai T, Crowe A (1978) Analyses of reflex movements in the hind limbs of the terrapinPseudemys scripta elegans. J Comp Physiol 125:351–357Google Scholar
  44. Valk-Fai T, Crowe A (1979) Further analyses of reflex movements in the hind limbs of the terrapin,Pseudemys scripta elegans. J Comp Physiol 130:241–249Google Scholar
  45. Walker WF (1973) The locomotor apparatus of testudines. In: Gans C (ed) Biology of the reptilia, vol 4, Morphology D. Academic Press, New York, pp 1–100Google Scholar
  46. Willis WD, Tate GW, Ashworth RD, Willis JC (1966) Monosynaptic excitation of motoneurons of individual forelimb muscles. J Neurophysiol 29:410–424Google Scholar

Copyright information

© Springer-Verlag 1985

Authors and Affiliations

  • H. Steffens
    • 1
  • E. D. Schomburg
    • 1
  • W. J. Koehler
    • 1
  1. 1.Physiologisches Institut der UniversitätGöttingenFederal Republic of Germany

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