SUMMARYMany arthropod olfactory appendages bear arrays of hair-like chemosensory sensillae. Odor molecules in the fluid around the animal must reach the surfaces of those hairs to be sensed. We used the lateral flagellum of the olfactory antennule of the spiny lobster, Panulirus argus, as a system to study how the morphology, orientation, and motion of sensilla-bearing appendages affects the small-scale water flow within the hair array. We tested whether antennule flicking enables lobsters to take discrete odor samples by measuring flow fields through an aesthetasc array on a dynamically scaled physical model of a P. argus antennule. Particle image velocimetry revealed that the magnitude and duration of velocity through the aesthetasc array during the rapid flick downstroke is just enough to allow complete replacement of the fluid entrained within the hair array. The complex zig-zag arrangement of aesthetascs hairs, combined with their offset orientation along the antennule, generates flow velocities that are uniform along the length of the hairs. This increases fluid exchange during the flick and reduces the boundary layer thickness surrounding the hairs. The return stroke occurs at about a quarter the speed of the flick, but the velocity of the fluid between the aesthetascs is approximately 25 times slower. The retained fluid during the return stroke remains virtually unstirred and sufficient time occurs for odor molecules to diffuse to aesthetasc surfaces.

Muscles not only generate force. They may act as springs, providing energy storage to drive locomotion. Although extensible myofilaments are implicated as sites of energy storage, we show that intramuscular temperature gradients may enable molecular motors (cross-bridges) to store elastic strain energy. Using time-resolved small-angle X-ray diffraction paired with in situ measurements of mechanical energy exchange in flight muscle of Manduca sexta we produced high-speed movies of X-ray equatorial reflections indicating cross-bridge association with myofilaments. A temperature gradient within the flight muscle leads to lower cross-bridge cycling in the cooler regions. Those cross-bridges could elastically return energy at the extrema of muscle lengthening and shortening, helping drive cyclic wing motions. These results suggest cross-bridges can perform functions other than contraction, acting as molecular links for elastic energy storage.

Given the notable Q 10 of the physiological properties of muscle and the presence of thermal gradients, temperature itself likely produces significant functional differences within a single muscle. Thus, although muscles are classically thought to function solely as a motor, spring, brake or strut, in some cases they may actually concurrently operate with an array of functions as a consequence of an internal temperature gradient (Altringham et al., 1993; Full Accepted 16 November 2011 SUMMARY A temperature gradient throughout the dominant flight muscle (dorsolongitudinal muscle, DLM 1 ) of the hawkmoth Manduca sexta, together with temperature-dependent muscle contractile rates, demonstrates that significant spatial variation in power production is possible within a single muscle. Using in situ work-loop analyses under varying muscle temperatures and phases of activation, we show that regional differences in muscle temperature will induce a spatial gradient in the mechanical power output throughout the DLM 1 . Indeed, we note that this power gradient spans from positive to negative values across the predicted temperature range. Warm ventral subunits produce positive power at their in vivo operating temperatures, and therefore act as motors. Concurrently, as muscle temperature decreases dorsally, the subunits produce approximately zero mechanical power output, acting as an elastic energy storage source, and negative power output, behaving as a damper. Adjusting the phase of activation further influences the temperature sensitivity of power output, significantly affecting the mechanical power output gradient that is expressed. Additionally, the separate subregions of the DLM 1 did not appear to employ significant physiological compensation for the temperature-induced differences in power output. Thus, although the components of a muscle are commonly thought to operate uniformly, a significant within-muscle temperature gradient has the potential to induce a mechanical power gradient, whereby subunits within a muscle operate with separate and distinct functional roles.

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