Kc. Nishikawa et al., Morphology and mechanics of tongue movement in the African pig-nosed frog Hemisus marmoratum: A muscular hydrostatic model, J EXP BIOL, 202(7), 1999, pp. 771-780
The goal of this study was to investigate morphological adaptations associa
ted with hydrostatic elongation of the tongue during feeding in the African
pig-nosed frog Hemisus marmoratum. Whereas previous studies had suggested
that the tongue of H, marmoratum elongates hydraulically, the anatomical ob
servations reported here favour a muscular hydrostatic mechanism of tongue
elongation. H. marmoratum possesses a previously undescribed compartment of
the m, genioglossus (m, genioglossus dorsoventralis), which is intrinsic t
o the tongue and whose muscle fibres are oriented perpendicular to the long
axis of the tongue. On the basis of the arrangement and orientation of mus
cle fibres in the m, genioglossus and m, hyoglossus, we propose a muscular
hydrostatic model of tongue movement in which contraction of the m, geniogl
ossus dorsoventralis, together with unfolding of the intrinsic musculature
of the tongue, results in a doubling in tongue length. Electron micrographs
of sarcomeres from resting and elongated tongues show that no special adap
tations of the sarcomeres are necessary to accommodate the observed doublin
g in tongue length during feeding. Rather, the sarcomeres of the m, geniogl
ossus longitudinalis are strikingly similar to those of anuran limb muscles
. The ability to elongate the tongue hydrostatically, conferred by the pres
ence of the m, genioglossus dorsoventralis, is associated with the appearan
ce of several novel aspects of feeding behaviour in H. marmoratum. These in
clude the ability to protract the tongue slowly, thereby increasing capture
success, and the ability to aim the tongue in azimuth and elevation relati
ve to the head. Compared with other frogs, the muscular hydrostatic system
of H, marmoratum allows more precise, localized and diverse tongue movement
s. This may explain why the m, genioglossus of H. marmoratum is composed of
a larger number of motor units than that of other frogs.