Abstract
A simulated neural network has been connected to a simulated mechanical environment. The network is based on a model of the spinal central pattern generator producing rhythmic swimming movements in the lamprey and the model is similar to that used in earlier simulations of fictive swimming. Here, the network has been extended with a model of how motoneuron activity is transformed via the muscles to mechanical forces. Further, these forces are used in a two-dimensional mechanical model including interaction with the surrounding water, giving the movements of the different parts of the body. Finally, these movements are fed back through stretch receptors interacting with the central pattern generator. The combined model provides a platform for various simulation experiments relating the currently known neural properties and connectivity to the behavior of the animalin vivo. By varying a small set of parameters, corresponding to brainstem input to the spinal network, a variety of basic locomotor behaviors, like swimming at different speeds and turning can be produced. This paper describes the combined model and its basic properties.
Similar content being viewed by others
References
Alishenas T (1992) Zur numerischen Behandlung, Stabilisierung durch Projection und Modellierung mechanischer Systeme mit Nebenbedingungen und Invarianten. PhD thesis, Royal Institute of Technology, Stockholm, 1992
Blake RW (1983) Fish locomotion. Cambridge University Press, London
Blevins RD (1984) Applied fluid dynamics handbook. Van Nostrand Reinhold, New York
Bowtell G, Williams T (1991) Anguilliform body dynamics: modellingthe interaction between muscle activation and body curvature. Phil Trans R Soc Lond B 334:385–390
Brodin L, Grillner S, Rovainen CM (1985)N-methyl-d-spartate (NMDA), kainate and quisqualate receptors and the generation of fictive locomotion in the lamprey spinal cord. Brain Res 325:302–306
Brodin L, Tråvén H, Lansner A, Wallén P, Ekeberg Ö, Grillner S (1991) Computer simulations ofN-methyl-d-aspartate (NMDA) receptor induced membrane properties in a neuron model. J Neurophysiol 66:473–484
Buchanan JT (1992) Neural network simulations of coupled locomotor oscillators in the lamprey spinal cord. Biol Cybern 66:367–374
Ekeberg Ö, Stensmo M, Lansner A (1990) SWIM — a simulator for real neural networks. Technical report TRITA-NA-P9014, Dept. of Nmerical Analysis and Computing Science, Royal Institute of Technology, Stockholm, Sweden
Ekeberg Ö, Wallén P, Lansner A, Tråvén H, Brodin L, Grillner S (1991) A computer based model for realistic simulations of neural networks. I. The single neuron and synaptic interaction. Biol Cybern 65:81–90
Gray DE (ed) (1972) American Institute of Physics handbook, 3rd edn. McGraw-Hill, New York
Gray J (1933a) Directional control of fish movement. Proc Roy Soc B 113:115–125
Gray J (1933b) Studies in animal locomotion. I. The movement of fish with special reference to the eel. J Exp Biol 10:88–103
Gray J (1968) Animal locomotion. Weidenfeld and Nicolson, London
Grillner S (1974) On the generation of locomotion in the spinal dogfish. Exp Brain Res 20:459–470
Grillner S (1981) Control of locomotion in bipeds, tetrapods and fish. In: Brooks VB (ed), Handbook of physiology, sect 1, vol 2. American Physiological Society, Bethesda, Md, pp 1179–1236
Grillner S, Kashin S (1976) On the generation and performance of swimming in fish. In: Herman RM, Grillner S, Stein PSG, Stuart DG (eds) Neural control of locomotion. Plenum, New York, pp 181–201
Grillner S, McClellan AD, Perret C (1981a) Entrainment of the spinal pattern generators for swimming by mechanosensitive elements in the lamprey spinal cord in vitro. Brain Res 217:380–386
Grillner S, McClellan AD, Sigvardt K, Wallén P, Wilén M (1981b) Activation of NMDA receptors elicits “fictive locomotion” in lamprey spinel cord in vitro. Acta Physiol Scand 113:549–551
Grillner S, McClellan A, Sigvardt K (1982) Mechanosensitive neurons in the spinal cord of the lamprey. Brain Res 235:169–173
Grillner S, Wallén P, Brodin L, Lansner A (1991) Neuronal network generating locomotor behavior in lamprey: circuitry, transmitters, membrane properties and simulations. Ann Rev Neurosci 4:169–199
Kalveram KT (1991) Controlling the dynamics of a two-joined arm by central patterning and reflex-like processing. Biol Cybern 65:65–71
Lacquaniti F, Soechting JF (1986) Simulation studies on the control of posture and movement in a multi-jointed limb. Biol Cybern 54:367–378
Lighthill J (1969) Hydromechanics of aquatic animal propulsion. Ann Rev Fluid Mech 1:413–445
Matsushima T, Grillner S (1992) Neural mechanisms of intersegmental coordination in lamprey: local excitability changes modify the phase coupling along the spinal cord. J Neurophysiol 67:373–388
Ólafsson Ö, Alishenas T (1992) A comparative study of the numerical integration and velocity stabilization of two mechanical test problems. Technical report TRITA-NA-9210, Dept of Numerical Analysis and Computing Science, Royal Institute of Technology, Stockholm, Sweden
Taga G, Yamaguchi Y, Shimizu H (1991) Self-organized control of bipedal locomotion by neural oscillators in unpredictable environment. Biol Cybern 65:147–159
Tax AAM, Denier van der Gon JJ (1991) A model for neural control of gradation of muscle force. Biol Cybern 65:227–234
Taylor G (1952) Analysis of the swimming of long and narrow animals. Proc Roy Soc Lond A 214:158–183
Viana Di Prisco G, Wallén P, Grillner S (1990) Synaptic effects of intraspinal stretch receptor neurons mediating movement-related feedback during locomotion. Brain Res 530:161–166
Wadden T, Grillner S, Matsushima T, Lansner A (1993) Undulatory locomotion — simulations with realistic segmental oscillator. In: Eeckman FM, Bower JM (eds), Computations & neural systems 92. Kluwer, Boston
Wallén P, Williams T (1984) Fictive locomotion in the lamprey spinal cord in vitro compared with swimming in the intact and spinal animal. J Physiol (Lond) 347:225–239
Wallén P, Ekeberg Ö, Lansner A, Brodin L, Tråvén H, Grillner S (1992) A computer-based model for realistic simulations of neural networks. II. Simulation of the segmental network generating locomotor rhythmicity in the lamprey. J. Neurophysiol 68:1939–1950
Webb PW, Weilhs D (1983) Fish biomechanics. Praeger, New York
Williams TL (1992) Phase coupling by synaptic spread in chains of coupled neuronal oscillators. Science 258:662–665
Williams TL, Grillner S, Smoljaninov VV, Wallén P, Kashin S, Rossignol S (1989) Locomotion in lamprey and trout: the relative timing of activation and movement. J Exp Biol 143:559–566
Wu TY-T (1971) Hydromechanics of swimming propulsion. J Fluid Mech 46:337–355, 521–568
Yates GT (1983) Hydrodynamics of body and caudal fin propulsion. In: Webb PW, Weihs D (eds) Fish biomechanics. Praeger, New York, pp 177–213
Author information
Authors and Affiliations
Rights and permissions
About this article
Cite this article
Ekeberg, Ö. A combined neuronal and mechanical model of fish swimming. Biol. Cybern. 69, 363–374 (1993). https://doi.org/10.1007/BF01185408
Received:
Accepted:
Issue Date:
DOI: https://doi.org/10.1007/BF01185408