Burrow ventilation in the tube-dwelling shrimp Callianassa subterranea (Decapoda: thalassinidea). III. Hydrodynamic modelling and the energetics of pleopod pumping.

The process of flow generation with metachronally beating pleopods in a tubiform burrow was studied by designing a hydrodynamic model based on a thrust-drag force balance. The drag of the tube (including the shrimp) comprises components for accelerating the water into the tube entrance, for adjusting a parabolic velocity profile, for accelerating the flow into a constriction due to the shrimp's body and another constriction due to the extended tail-fan, for shear due to separation and for the viscous resistance of all tube parts. The thrust produced by the beating pleopods comprises components for the drag-based thrust and for the added-mass-based thrust. The beating pleopods are approximated by oscillating flat plates with a different area and camber during the power stroke and the recovery stroke and with a phase shift between adjacent pleopod pairs. The added mass is shed during the second half of the power stroke and is minimized during the recovery stroke. A force balance between the pleopod thrust and the tube drag is effected by calculating the mean thrust during one beat cycle at a certain flow velocity in the tube and comparing it with the drag of the tube at that flow velocity. The energetics of the tube and the pump are derived from the forces, and the mechanical efficiency of the system is the ratio of these two. Adjusted to standard Callianassa subterranea values, the model predicts a mean flow velocity in the tube of 1.8 mm s-1. The mean thrust force, equalling the drag, is 36. 8 microN, the work done by the pleopod pump per beat cycle is 0.91 microJ and the energy dissipated by the tube system is 0.066 microJ per cycle. The mechanical efficiency is therefore 7.3 %. Pump characteristics that may be varied by the shrimp are the beat frequency, the phase shift, the amplitude and the difference in pleopod area between the power and recovery strokes. These parameters are varied in the model to evaluate their effects. Furthermore, the moment of added mass shedding, the distance between adjacent pleopods, the number of pleopods and the total tube drag were also varied to evaluate their effects

Burrow ventilation in the tube-dwelling shrimp callianassa subterranea (Decapoda: thalassinidea). II. The flow in the vicinity of the shrimp and the energetic advantages of a laminar non-pulsating ventilation current.

The ventilation flow in the vicinity of the pleopod-pumping thalassinid shrimp Callianassa subterranea in an artificial transparent burrow has been mapped using particle image velocimetry. The flow in the tube in front of the shrimp was unidirectional, laminar and steady, with a parabolic cross-sectional velocity profile. The mean flow velocity was 2.0+/-0.1 mm s-1. The flow passed the thorax of the shrimp along the lateral and ventral sides. Ventral to the abdomen, the flow was dominated by the metachronally oscillating pleopods. The water around a pleopod is accelerated caudally and ventrally during the power stroke, and decelerated to a much lesser extent during the recovery stroke owing to a reduction in pleopod area. On average, the flow ventral to the abdomen converged towards the small opening underneath the telson, simultaneously increasing in velocity. A jet with a core velocity of 18-20 mm s-1 entered the area behind the shrimp from underneath the telson. This caused a separation zone with backflow caudal to the telson. Owing to the high rates of shear, the jet diverged and re-adjusted to a parabolic cross-sectional profile within 1-2 body lengths behind the shrimp, showing no traces of pulsation. The metachronal pleopod movements in combination with the increase in flow velocity at the constriction in the tube caused by the uropods and the telson probably prevented pulsation. The energetic consequences of pulsating and steady flows in combination with several tube configurations were evaluated. The results suggested that, by constricting the tube and keeping the flow steady, C. subterranea saves on ventilation costs by a factor of up to six compared with oscillatory flow in a tube without the tail-fan constriction

Burrow ventilation in the tube-dwelling shrimp Callianassa subterranea (Decapoda Thalassinidea) - I. Morphology and motion of the pleopods, uropods and telson

The morphology of the pleopods, uropods and telson of the tube-dwelling shrimp Callianassa subterranea have been studied using dissection microscopy and scanning electron microscopy. The kinematics of these appendages were examined by motion analysis of macro-video recordings of ventilating shrimps in transparent artificial burrows. The pleopods show the usual crustacean biramous anatomy, but all segments are rostro-caudally flattened. The protopodite bears a triangular medially oriented endopodite and a scoop-shaped exopodite. The contralateral endopodites are linked by the appendix interna, ensuring a perfect phase relationship between contralateral pleopods. The outer rims of the exopodites are fringed with long fern-leaf-like plumose setae bearing flattened setules. These setae have very mobile joints and can be turned caudally. The slits between contralateral endopodites have rims of leaf-like setae as well. Setae of the same leaf-like type fringe the uropods, but these are non-motile. The telson has a narrow fringe of leaf-like setae, locally interrupted by long mechanoreceptory setae. A shrimp, wandering through the burrow or resting, holds its pleopods against the abdomen with the exopodites and their setal fringes retracted medially. The uropods are folded medially as well, probably to reduce the shrimp's drag. During ventilation, the uropods are extended against the tube wall, leaving only a small opening for flow ventral from the telson, and the pleopods beat at a frequency of approximately 1 Hz (0.9+/-0.2 Hz). Fourier analysis of pleopod kinematics showed that the motion pattern of the first flow-generating pleopod pair (PP1) consisted mainly of the first harmonic (75 %) and to a lesser extent of the third harmonic (20 %), resulting in almost perfect sinusoidal motion. The motion patterns of PP2 and PP3 could be modelled by assigning pure sinusoids with a 120 degrees phase shift and a rostro-caudal ranking to the three pairs of pleopods

Does genetic differentiation underlie behavioral divergence in response to migration barriers in sticklebacks?:A common garden experiment

Water management measures in the 1970s in the Netherlands have produced a large number of “resident” populations of three-spined sticklebacks that are no longer able to migrate to the sea. This may be viewed as a replicated field experiment, allowing us to study how the resident populations are coping with human-induced barriers to migration. We have previously shown that residents are smaller, bolder, more exploratory, more active, and more aggressive and exhibited lower shoaling and lower migratory tendencies compared to their ancestral “migrant” counterparts. However, it is not clear if these differences in wild-caught residents and migrants reflect genetic differentiation, rather than different developmental conditions. To investigate this, we raised offspring of four crosses (migrant ♂ × migrant ♀, resident ♂ × resident ♀, migrant ♂ × resident ♀, resident ♂ × migrant ♀) under similar controlled conditions and tested for differences in morphology and behavior as adults. We found that lab-raised resident sticklebacks exhibited lower shoaling and migratory tendencies as compared to lab-raised migrants, retaining the differences in their wild-caught parents. This indicates genetic differentiation of these traits. For all other traits, the lab-raised sticklebacks of the various crosses did not differ significantly, suggesting that the earlier-found contrast between wild-caught fish reflects differences in their environment. Our study shows that barriers to migration can lead to rapid differentiation in behavioral tendencies over contemporary timescales (~ 50 generations) and that part of these differences reflects genetic differentiation