Modulatory multiplicity in the feeding mechanism in cichlid fishes, as exemplified by the invertebrate pickers of Lake Tanganyika

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Abstract
Electromyography, motion analysis, osteology, myology and feeding behaviour of a morphologically specialized monophyletic lineage of cichlids in Lake Tanganyika have revealed that Eretmodus, Spathodus and Tanganicodus possess a feeding apparatus with a more extensive functional repertoire than that of any other teleost studied to date. When collecting a wide range of foods by inertial suction this trophic group can employ two strategies, a preprogrammed cyclical, energy saving pattern, or a modulated mode effected by extensive overlap of the firing patterns of multiple muscles resulting in a precise control of the magnitude and direction of suction. When dislodging sessile prey from the substrate the complexity of electromyographic and kinematic patterns increases. Because upper jaw protrusion can be effected and controlled independently from the complex couplings causing mouth opening and movements of the suspensory apparatus, a new decoupled model of upper jaw protrusion is proposed. The decoupled model predicts that upper jaw protrusion can be effected directly by contraction of epaxial muscles that raise the neurocranium, causing the premaxillae to slide anteroventrally. Upper jaw protrusion can be modulated continuously and directly by balanced cocontractions of antagonistic muscle sets giving the decoupled model an improved function over a very extensive range. The morphologically symmetrical muscular apparatus can function asymmetrically. Very pronounced asymmetrical firings of multiple muscles produce a continuously modulated jaw mechanism with an extensive repertoire.