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First published online January 27, 2004
Journal of Experimental Biology 207, 827-839 (2004)
Published by The Company of Biologists 2004
doi: 10.1242/jeb.00819
A three-dimensional kinematic analysis of tongue flicking in Python molurus
1 Section Evolutionary Morphology, Institute of Biology (IBL), Leiden
University, PO Box 9516, 2300 RA Leiden, The Netherlands
2 Experimental Zoology Group, Wageningen Institute of Animal Sciences
(WIAS), Wageningen University, Marijkeweg 40, 6709 PG, Wageningen, The
Netherlands
* Author for correspondence (e-mail: j.h.de_groot{at}lumc.nl)
Accepted 3 December 2003
The forked snake tongue is a muscular organ without hard skeletal support.
A functional interpretation of the variable arrangement of the intrinsic
muscles along the tongue requires a quantitative analysis of the motion
performance during tongue protrusion and flicking. Therefore, high-speed
fluoroscopy and high-speed stereo photogrammetry were used to analyse the
three-dimensional shape changes of the tongue in Python molurus
bivittatus (Boidae). The posterior protruding part of the tongue
elongated up to 130% while the flicking anterior portion elongated maximally
60%. The differences in tongue strains relate to the absence or presence,
respectively, of longitudinal muscle fibres in the peripheral tongue. Maximum
overall protrusion velocity (4.3 m s1) occurred initially
when the tongue tip left the mouth. Maximum tongue length of
0.01 body
length (20 mm) was reached during the first tongue flick. These observations
are discussed within the scope of the biomechanical constraints of hydrostatic
tongue protrusion: a negative forward pressure gradient, longitudinal tongue
compliance and axial tongue stiffness. The three-dimensional deformation
varied along the tongue with a mean curvature of 0.06 mm1
and a maximum value of 0.5 mm1. At the basis of the anterior
forked portion of the tongue tips, extreme curvatures up to 2.0
mm1 were observed. These quantitative results support
previously proposed inferences about a hydrostatic elongation mechanism and
may serve to evaluate future dynamic models of tongue flicking.
Key words: snake, tongue, tongue sheet, muscular hydrostat, flicking, curvature, 3-D kinematics, high-speed fluoroscopy, X-ray
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