«James Walker 1st year DPhil, Interdisciplinary Bioscience Doctoral Training Partnership Supervised by Prof Fritz Vollrath and Dr Beth Mortimer Oxford ...»
If a spider system is used which does show regeneration, the integration of legs of different sizes into the stepping pattern could be explored using techniques developed in this study in addition investigations into the sensory morphology of regenerated appendages. With additional time, a more sophisticated gait analysis setup could be adopted using a locomotion compensator, such as that developed by Kramer (1975). This would involve attaching the spider above a styrofoam sphere that is rotated by motors to compensate for the subject’s movement keeping it on top of the sphere facilitating an improved data acquisition setup with automated gait analysis.
The regeneration of legs in running spiders provides a system to further investigate the role of sensory information in control as signals are not completely removed but modified peripherally in response to changes in leg morphology. The addition of new sensory information from regenerated legs would presumably result in modifications to the control algorithm providing a useful insight into the mechanism of morphological computation. Whilst moulting and regeneration in spiders is a discontinuous process, inspiration can be gained from this control system to tackle a key problem for soft robotics: the design of control systems for adaptable, compliant robots showing continuous morphological change (Calisti et al., 2011). I conclude my study with the suggestion that the sensory basis of spider walking is likely to provide useful guidelines for the design of embodied controllers that do not require complex central programming but can outsource much of the computation to adaptable peripheral sensors.
ACKNOWLEDGEMENTSI would like to thank Prof Fritz Vollrath for supervising my project, for introducing me to the field of soft robotics and for many interesting discussions and comments throughout project and on earlier versions of this report. I am grateful to Dr Beth Mortimer who helped with the spider identification, experimental setup, statistical techniques and gave comments on several drafts of this manuscript.
Finally, I thank Nick Hawkins for help with the housing and maintenance of spiders. This work was funded by a BBSRC PhD studentship grant.
REFERENCESAMAYA, C. C., KLAWINSKI, P. D. & FORMANOWICZ, D. R. 2001. The effects of leg autotomy on running speed and foraging ability in two species of wolf spider, (Lycosidae). American Midland Naturalist, 145, 201-205.
APONTES, P. & BROWN, C. A. 2005. Between-sex variation in running speed and a potential cost of leg autotomy in the wolf spider Pirata sedentarius. American Midland Naturalist, 154, 115-125.
BARTH, F. G. 2002. A spider’s world: senses and behavior, Springer Science & Business Media.
BENJAMINI, Y. & HOCHBERG, Y. 1995. Controlling the False Discovery Rate - a Practical and Powerful Approach to Multiple Testing. Journal of the Royal Statistical Society Series B-Methodological, 57, 289-300.
BIANCARDI, C. M., FABRICA, C. G., POLERO, P., LOSS, J. F. & MINETTI, A. E. 2011. Biomechanics of octopedal locomotion: kinematic and kinetic analysis of the spider Grammostola mollicoma.
The Journal of experimental biology, 214, 3433-3442.
BROWN, C. A. & FORMANOWICZ, D. R., JR. 2012. The effect of leg autotomy on terrestrial and aquatic locomotion in the wolf spider Pardosa valens (Araneae: Lycosidae). Journal of Arachnology, 40, 234-239.
CALISTI, M., GIORELLI, M., LEVY, G., MAZZOLAI, B., HOCHNER, B., LASCHI, C. & DARIO, P. 2011. An octopus-bioinspired solution to movement and manipulation for soft robots. Bioinspiration & biomimetics, 6, 036002.
DELCOMYN, F. 1985. FACTORS REGULATING INSECT WALKING. Annual Review of Entomology, 30, 239DELCOMYN, F. 1999. Walking robots and the central and peripheral control of locomotion in insects.
Autonomous Robots, 7, 259-270.
FOELIX, R. 2010. Biology of spiders, Oxford University Press.
GÖRNER, P. & CLAAS, B. 1985. Homing behavior and orientation in the funnel-web spider, Agelena labyrinthica Clerck, Springer.
HAUSER, H., IJSPEERT, A. J., FUCHSLIN, R. M., PFEIFER, R. & MAASS, W. 2011. Towards a theoretical foundation for morphological computation with compliant bodies. Biological Cybernetics, 105, 355-370.
KRAMER, E. 1975. Orientation of the male silkmoth to the sex attractant bombycol. Olfaction and Taste. Academic Press, New York.
LAND, M. 1985. The morphology and optics of spider eyes. Neurobiology of arachnids. Springer.
MAYORGA, O., MAZOUCHOVA, N., GOLDMAN, D. I. & SPAGNA, J. C. 2013. Running performance and gait kinematics of a sand-adapted arachnid, Galeodes granti. Integrative and Comparative Biology, 53, E140-E140.
MIKULSKA, I., JACUNSKI, L. & WEYCHERT, K. 1975. regeneration of appendages in Tegenaria atrica CL Koch (Agelenidae, Araneae). ZP Zool Pol.
MOLLER, P. & GORNER, P. 1994. HOMING BY PATH INTEGRATION IN THE SPIDER AGELENALABYRINTHICA CLERCK. Journal of Comparative Physiology a-Sensory Neural and Behavioral Physiology, 174, 221-229.
PFEIFER, R., IIDA, F. & LUNGARELLA, M. 2014. Cognition from the bottom up: on biological inspiration, body morphology, and soft materials. Trends in cognitive sciences.
ROTH, V. D. R., B. M 1984. A review of appendotomy in spiders and other arachnids. Bull. Br. arachnol.
Soc. 6, 137-146.
SEYFARTH, E.-A. 1985. Spider proprioception: receptors, reflexes, and control of locomotion.
Neurobiology of arachnids. Springer.
SPAGNA, J. C., VALDIVIA, E. A. & MOHAN, V. 2011. Gait characteristics of two fast-running spider
species (Hololena adnexa and Hololena curta), including an aerial phase (Araneae:
Agelenidae). Journal of Arachnology, 39, 84-91.
STEFFENSON, M. M., FORMANOWICZ, D. R. & BROWN, C. A. 2014. Autotomy and its Effects on Wolf Spider Foraging Success. Ethology, 120, 1128-1136.
TRIMMER, B. A. 2008. New challenges in biorobotics: Incorporating soft tissue into control systems.
Applied Bionics and Biomechanics, 5, 119-126.
VOLLRATH, F. 1990. Leg regeneration in web spiders and its implications for orb weaver phylogeny.
Bulletin of the British Arachnological Society.
VOLLRATH, F. 1995. Lyriform organs on regenerated spider legs. Bulletin of the British Arachnological Society, 10, 115-118.
WEISSMANN, M. 1987. Web-building and prey capture in two orb weavers. M. Sc., Oxford University.
WETTERGREEN, D. & THORPE, C. Gait generation for legged robots. IEEE International Conference on Intelligent Robots and Systems, 1992.
WILSON, D. M. 1967. Stepping Patterns in Tarantula Spiders. Journal of Experimental Biology, 47, 133WRINN, K. M. & UETZ, G. W. 2007. Impacts of leg loss and regeneration on body condition, growth, and development time in the wolf spider Schizocosa ocreata. Canadian Journal of ZoologyRevue Canadienne De Zoologie, 85, 823-831.
WRINN, K. M. & UETZ, G. W. 2008. Effects of autotomy and regeneration on detection and capture of prey in a generalist predator. Behavioral Ecology, 19, 1282-1288.
ZOLLIKOFER, C. P. E. 1994. Stepping Patterns in Ants.1. Influence of Speed and Curvature. Journal of Experimental Biology, 192, 95-106.