The biomechanics of fast-starts during ontogeny in the common carp cyprinus carpio

Common carp Cyprinus carpio L. were reared a constant temperature of 20 degrees C from the larval (7 mm total length) to the juvenile (80 mm) stage. Body morphology and white muscle mass distribution were measured. Fast-start escape responses were recorded using high-speed cinematography from which the velocities, accelerations and hydrodynamic power requirements were estimated. All three measures of fast-start performance increased during development. White muscle contraction regimes were calculated from changes in body shape during the fast-starts and used to predict the muscle force and power production for all longitudinal positions along the body. Scaling arguments predicted that increases in body length would constrain the fish to bend less rapidly because the cross-sectional muscle area, and hence force production, does not increase at the same rate as the inertial mass that resists bending. As predicted, the increases in body length resulted in decreases in muscle shortening velocity, and this coincided with increases in both the force and power produced by the muscles. The hydrodynamic efficiency, which relates the mechanical power produced by the muscles to the inertial power requirements in the direction of travel, showed no significant change during ontogeny. The increasing hydrodynamic power requirements were thus met by increases in the power available from the muscles. The majority of the increases in fast-start swimming performance during ontogeny can be explained by size-dependent increases in muscle power output. For all sizes, there was a decrease in muscle-mass-specific power output and an increase in muscle stress in a posterior direction along the body due to systematic variations in fibre strain. These changing strain regimes result in the central muscle bulk producing the majority of the power requirements during the fast-start, and this power is transmitted to the tail region of the fish and ultimately to the water via muscle in the caudal myotomes.

Body bending during fast-starts in fish can be explained in terms of muscle torque and hydrodynamic resistance

Fish fast-starts are rapid events caused by the simultaneous onset of muscle activity along one side of the body. Spine curvature and the strain and electromyographic activity in white muscle were measured for fast-starts in the common carp Cyprinus carpio. The first bend of the fast-start was powered by muscle on the concave side: muscle fibres on this side were activated and began shortening simultaneously between the length-specific longitudinal sites 0.3L and 0.56L, where L is total body length. However, there was an increasing delay in the timing of the first peak in body curvature and muscle strain along the length of the body. Modelling studies related the rate of body bending to the muscle torque and hydrodynamic resistance of the fish. The muscle torque produced on the spine was greatest in the central region of the trunk, and this acted against the moments of inertia of the fish mass and added mass of water. It was concluded that a wave of body bending can be generated as a result of the hydrodynamic resistance of the fish despite the initiation of that bending being simultaneous along the length of the body.

White muscle strain in the common carp and red to white muscle gearing ratios in fish

White muscle strains were recorded using sonomicrometry techniques for 70 fast-starts in the common carp Cyprinus carpio L. High-speed cine images were recorded simultaneously for 54 of these starts, and muscle strain was calculated independently from the digitized outlines of the fish. Sonomicrometry measurements of superficial muscle strain were not significantly different from the strain as calculated from the theory of simple bending of a homogeneous material: superficial muscle strain thus varied with chordwise distance from the spine. However, white muscle strain across a transverse section of the myotome shows less variation with chordwise position than would be expected from simple bending theory. Muscle strains measured using sonomicrometry thus do not necessarily represent the more uniform strain predicted for the whole section of the fish. White muscle strain can be accurately predicted from the spine curvatures as measured from the cine images if the gearing ratio between the red and white muscle fibres is known. A model for calculating the gearing ratio from the helical muscle fibre geometry was re-evaluated using current data for the kinematics of fast-starting C. carpio. This model predicted a mean gearing ratio of 2.8 for these fast-starts. A quicker, alternative approach to estimating gearing ratio from the position of the centroid of white fibre area is proposed and results in ratios similar to those calculated from the model of helical geometry. White muscle strains in fish can thus be estimated from measurements of spine curvature and muscle distribution alone.

Molecular cloning and developmental expression patterns of the MyoD and MEF2 families of muscle transcription factors in the carp.

cDNA clones encoding the myogenic regulatory factors (MRFs) myogenin, MyoD and myf-5 were isolated by reverse-transcription polymerase chain reaction from larvae and embryos of the common carp (Cyprinus carpio L.). Myocyte-specific enhancer factor 2 (MEF2) cDNAs were identified from a cDNA library from adult carp. Northern blot analysis showed that MyoD, myf-5 and MEF2C transcripts were present in three-somite embryos, whereas myogenin and MEF2A transcripts were not detected until the 15-somite stage. Intense signals of myogenin and MyoD transcripts were observed even in 1-month-old juveniles. Levels of MyoD, myogenin and MEF2A transcripts declined between 1 and 7 months after hatching, and myf-5 gave only a weak signal in the oldest fish. In contrast, levels of MEF2C transcripts were considerably higher in 7-month-old juveniles than in 1-month-old larvae. mRNAs encoding carp myosin heavy chain and -actin were first detected at approximately the time of the first heartbeat, and levels were maximal in juveniles 1 month post-hatching. The relatively high levels of MRF mRNA in juvenile fish probably reflect the recruitment of new muscle fibres from the satellite cell population. It was concluded that the relative importance of the different members of the MyoD and MEF2 families of transcription factors for muscle differentiation changes during ontogeny in the carp.