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First published online July 20, 2006
Journal of Experimental Biology 209, 2952-2960 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02350
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Sound production mechanism in carapid fish: first example with a slow sonic muscle

Eric Parmentier1,*, Jean-Paul Lagardère2, Jean-Baptiste Braquegnier3, Pierre Vandewalle1 and Michael L. Fine4

1 Laboratoire de Morphologie Fonctionnelle et Evolutive, Institut de chimie, Université de Liège, B-4000 Liège, Belgium
2 CREMA-L'Houmeau (CNRS-Ifremer), BP 5, 17137 L'Houmeau, France
3 Laboratoire d'Ecophysiologie et Physiologie Animale, Université de Liège, B-4000 Liège, Belgium
4 Department of Biology, Virginia Commonwealth University, Richmond, VA 23284-2012, USA


Figure 1
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  		Fig. 1. Sonic mechanism of Carapus boraborensis. (A) Left lateral view of the first five vertebrae (I-V) with the associated epineural ribs and swimbladder plate. Numbers on the swimbladder plate indicate its thickness, which decreases laterally and toward its posterior margin. (B) Schematic left lateral view of the skeleton (epineural ribs and swimbladder plate removed), sonic muscles and swimbladder. (C) Inset in B illustrates the helical organisation of the myofibrils in a single sonic muscle fiber with straight central myofibrils. SF, swimbladder fenestra; SM, sonic muscle; SP, swimbladder plate, SW, swimbladder.

 

Figure 2
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  		Fig. 2. Contraction of Carapus acus sonic muscle stimulated at 2, 8, 13 and 20 Hz.

 

Figure 3
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  		Fig. 3. Amplitude of sonic muscle contractions at different stimulation frequencies in Carapus acus.

 

Figure 4
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  		Fig. 4. Details of the sonic muscle insertion on the swimbladder in Carapus boraborensis. (A) Micrograph of the anterior face of the swimbladder illustrating the tubercles that anchor the tendon hooks at rest. (B) Micrograph of the complex sonic muscle tendon (anterior is toward the bottom) illustrating the hook and connective tissue, which attach to the swimbladder. (C,D,E) Frontal sections of the anterior part of the swimbladder illustrating the tubercles that anchor the tendon hooks. CT, connective tissue; SWB, swimbladder; Tb, tubercule; black arrows, tissue of the swimbladder fenestra. Scale bars, 50 µm.

 

Figure 5
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  		Fig. 5. (A) Diagram of sound pulse generation progressing in time from top to bottom. Contraction or pulling on the sonic muscle in a fresh specimen extends the anterior bladder and swimbladder fenestra. In the third panel the hook has disengaged from the bladder tubercle, and the system then snaps back to its resting state generating sound. (B) Left lateral view of the anterior part of the swimbladder. Blue arrows represent the amplitude of the displacement of different points of the swimbladder fenestra when stretched.

 

Figure 6
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  		Fig. 6. Voluntary and manually induced sound pulses in Carapus boraborensis. Sonagram of 6 voluntary pulses (Ai) produced by a fish within a sea cucumber and 3 pulses (Aii) produced by 3 pulls on the sonic muscle of a fish with its sonic apparatus exposed. Oscillograms of a single pulse produced voluntarily (B) and by manual manipulation (C). Broken lines indicate termination of the pulse. The color ramp in A shows the scale of the relative dB, from white (lowest) to red (highest).

 

Figure 7
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  		Fig. 7. (A) Periods of successive peaks in the voluntary (open squares) and manually induced (filled circles) sound pulses in Carapus boraborensis. (N=25 in both cases). (B) FFT of voluntary sound in Carapus boraborensis. The peak frequency (black arrow) corresponds to the part of the waveform within the rectangular broken line in A above.

 

Figure 8
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  		Fig. 8. Peak frequency of manually generated sounds (filled circles) and calculated resonant frequency (open squares) in a 5 cm deep vessel of seawater.

 




© The Company of Biologists Ltd 2006