Power output from a flight muscle of the bumblebee Bombus terrestris. II. Characterization of the parameters affecting power output
1. Length-tension relationships and work output were investigated in the intact, dorso-ventral flight muscle of the bumblebee Bombus terrestris. The muscle is an asynchronous muscle. Like other asynchronous flight muscles, it has high resting stiffness and produces relatively low active force in response to tetanic stimulation. 2. The muscle shows shortening deactivation and stretch activation, properties that result in delayed force changes in response to step changes in length, a phase lag between force and length during imposed sinusoidal strain and, under appropriate conditions, positive work output during oscillatory length change. 3. Work loops were used to quantify work output by the muscle during imposed sinusoidal oscillation. The curves relating net work per cycle with muscle length, oscillatory strain and oscillatory frequency were all roughly bell-shaped. The work-length curve was narrow. The optimum strain for net work per cycle was approximately 3 %, which is probably somewhat greater than the strain experienced by the muscle in an intact, flying bumblebee. The optimum frequency for net work output per cycle was 63 Hz (30 &deg;C). The optimum frequency for power output was 73 Hz, which agrees well with the normal wing stroke frequency if allowance is made for the elevated temperature (approximately 40 &deg;C) in the thorax of a flying bumblebee. The optimal strain for work output was not strongly dependent on oscillation frequency. 4. Resilience (that is the work output during shortening/work input during lengthening) for unstimulated muscle and dynamic stiffness (=stress/strain) for both stimulated and unstimulated muscles were determined using the strain (3 %) and oscillation frequency (64 Hz) which maximized work output in stimulated muscles. Unstimulated muscle is a good energy storage device. Its resilience increased with increasing muscle length (and increasing resting force) to reach values of over 90 %. The dynamic stiffness of both stimulated and unstimulated muscles increased with muscle length, but the increase was relatively greater in unstimulated muscle, and at long muscle lengths the stiffness of unstimulated muscle exceeded that of stimulated muscle. Effectively, dynamic stiffness is reduced by stimulation! This is taken as indicating that part of the stiffness in an unstimulated muscle reflects structures, possibly attached cross bridges, whose properties change upon stimulation.
Josephson
R