title: Positive force feedback in development of substrate grip in the stick insect tarsus.
abstract: The mechanics of substrate adhesion has recently been intensively studied in insects but less is known about the sensorimotor control of substrate engagement. We characterized the responses and motor effects of tarsal campaniform sensilla in stick insects to understand how sensory signals of force could contribute to substrate grip. The tarsi consist of a chain of segments linked by highly flexible articulations. Morphological studies showed that one to four campaniform sensilla are located on the distal end of each segment. Activities of the receptors were recorded neurographically and sensilla were identified by stimulation and ablation of their cuticular caps. Responses were characterized to bending forces and axial loads, muscle contractions and to forces applied to the retractor apodeme (tendon). The tarsal sensilla effectively encoded both the rate and amplitude of loads and muscle forces, but only when movement was resisted. Mechanical stimulation of the receptors produced activation of motor neurons in the retractor unguis and tibial flexor muscles. These findings indicate that campaniform sensilla can provide information about the effectiveness of the leg muscles in generating substrate adherence. They can also produce positive force feedback that could contribute to the development of substrate grip and stabilization of the tarsal chain.
authors.affiliation: Department of Anatomy and Pathology, Joan C. Edwards School of Medicine, Marshall University, Huntington, WV 25704, USA. Electronic address: sensillum@aol.com.
authors.forename: Sasha N
authors.initials: SN
authors.lastname: Zill
authors.affiliation: Department of Anatomy and Pathology, Joan C. Edwards School of Medicine, Marshall University, Huntington, WV 25704, USA.
authors.forename: Sumaiya
authors.initials: S
authors.lastname: Chaudhry
authors.affiliation: Department of Biological Cybernetics, University of Bielefeld, 33501 Bielefeld, Germany.
authors.forename: Annelie
authors.initials: A
authors.lastname: Exter
authors.affiliation: Department of Animal Physiology, Zoological Institute, Biocenter Cologne, University of Cologne, 50923 Cologne, Germany.
authors.forename: Ansgar
authors.initials: A
authors.lastname: Büschges
authors.affiliation: Department of Biological Cybernetics, University of Bielefeld, 33501 Bielefeld, Germany.
authors.forename: Josef
authors.initials: J
authors.lastname: Schmitz
mesh_terms: D000818:Animals; D001696:Biomechanical Phenomena; D005121:Extremities / Q000502:physiology; D005260:Female; D007313:Insecta / Q000502:physiology*; D009043:Motor Activity; D009046:Motor Neurons / Q000502:physiology; D009119:Muscle Contraction; D011434:Proprioception; D058618:Sensilla / Q000502:physiology
keywords: Encoding; Force; Grip; Insects