Hydrodynamic stress and phenotypic plasticity of the zebrafish regenerating fin.

Understanding how extrinsic factors modulate genetically encoded information to produce a specific phenotype is of prime scientific interest. In particular, the feedback mechanism between abiotic forces and locomotory organs during morphogenesis to achieve efficient movement is a highly relevant example of such modulation. The study of this developmental process can provide unique insights on the transduction of cues at the interface between physics and biology. Here, we take advantage of the natural ability of adult zebrafish to regenerate their amputated fins to assess its morphogenic plasticity upon external modulations. Using a variety of surgical and chemical treatments, we could induce phenotypic responses to the structure of the fin. Through the ablation of specific rays in regenerating caudal fins, we generated artificially narrowed appendages in which the fin cleft depth and the positioning of rays bifurcations were perturbed compared with normal regenerates. To dissect the role of mechanotransduction in this process, we investigated the patterns of hydrodynamic forces acting on the surface of a zebrafish fin during regeneration by using particle tracking velocimetry on a range of biomimetic hydrofoils. This experimental approach enabled us to quantitatively compare hydrodynamic stress distributions over flapping fins of varying sizes and shapes. As a result, viscous shear stress acting on the distal margin of regenerating fins and the resulting internal tension are proposed as suitable signals for guiding the regulation of ray growth dynamics and branching pattern. Our findings suggest that mechanical forces are involved in the fine-tuning of the locomotory organ during fin morphogenesis.

Elephants evolved strategies reducing the biomechanical complexity of their trunk.

The elephant proboscis (trunk), which functions as a muscular hydrostat with a virtually infinite number of degrees of freedom, is a spectacular organ for delicate to heavy object manipulation as well as social and sensory functions. Using high-resolution motion capture and functional morphology analyses, we show here that elephants evolved strategies that reduce the biomechanical complexity of their trunk. Indeed, our behavioral experiments with objects of various shapes, sizes, and weights indicate that (1) complex behaviors emerge from the combination of a finite set of basic movements; (2) curvature, torsion, and strain provide an appropriate kinematic representation, allowing us to extract motion primitives from the trunk trajectories; (3) transport of objects involves the proximal propagation of an inward curvature front initiated at the tip; (4) the trunk can also form pseudo-joints for point-to-point motion; and (5) the trunk tip velocity obeys a power law with its path curvature, similar to human hand drawing movements. We also reveal with unprecedented precision the functional anatomy of the African and Asian elephant trunks using medical imaging and macro-scale serial sectioning, thus drawing strong connections between motion primitives and muscular synergies. Our study is the first combined quantitative analysis of the mechanical performance, kinematic strategies, and functional morphology of the largest animal muscular hydrostat on Earth. It provides data for developing innovative "soft-robotic" manipulators devoid of articulations, replicating the high compliance, flexibility, and strength of the elephant trunk. VIDEO ABSTRACT.

Hydrodynamic stress maps on the surface of a flexible fin-like foil.

We determine the time dependence of pressure and shear stress distributions on the surface of a pitching and deforming hydrofoil from measurements of the three dimensional flow field. Period-averaged stress maps are obtained both in the presence and absence of steady flow around the foil. The velocity vector field is determined via volumetric three-component particle tracking velocimetry and subsequently inserted into the Navier-Stokes equation to calculate the total hydrodynamic stress tensor. In addition, we also present a careful error analysis of such measurements, showing that local evaluations of stress distributions are possible. The consistency of the force time-dependence is verified using a control volume analysis. The flapping foil used in the experiments is designed to allow comparison with a small trapezoidal fish fin, in terms of the scaling laws that govern the oscillatory flow regime. As a complementary approach, unsteady Euler-Bernoulli beam theory is employed to derive instantaneous transversal force distributions on the flexible hydrofoil from its deflection and the results are compared to the spatial distributions of hydrodynamic stresses obtained from the fluid velocity field.

Distribution and Restoration of Serotonin-Immunoreactive Paraneuronal Cells During Caudal Fin Regeneration in Zebrafish.

Aquatic vertebrates possess diverse types of sensory cells in their skin to detect stimuli in the water. In the adult zebrafish, a common model organism, the presence of such cells in fins has only rarely been studied. Here, we identified scattered serotonin (5-HT)-positive cells in the epidermis of the caudal fin. These cells were distinct from keratinocytes as revealed by their low immunoreactivity for cytokeratin and desmosome markers. Instead, they were detected by Calretinin (Calbindin-2) and Synaptic vesicle glycoprotein 2 (SV2) antibodies, indicating a calcium-regulated neurosecretory activity. Consistently, electron microscopy revealed abundant secretory organelles in desmosome-negative cells in the fin epidermis. Based on the markers, 5-HT, Calretinin and SV2, we referred to these cells as HCS-cells. We found that HCS-cells were spread throughout the entire caudal fin at an average density of 140 cells per mm2 on each fin surface. These cells were strongly enriched at ray bifurcations in wild type fins, as well as in elongated fins of another longfin mutant fish. To determine whether hydrodynamics play a role in the distribution of HCS-cells, we used an interdisciplinary approach and performed kinematic analysis. Measurements of particle velocity with a fin model revealed differences in fluid velocities between bifurcated rods and adjacent non-bifurcated regions. Therefore the accumulation of HCS-cells near bone bifurcations may be a biological adaptation for sensing of water parameters. The significance of this HCS-cell pattern is reinforced by the fact, that it is reestablished in the regenerated fin after amputation. Regeneration of HCS-cells was not impaired by the chemical inhibition of serotonin synthesis, suggesting that this neurotransmitter is not essential for the restorative process. In conclusion, our study identified a specific population of solitary paraneurons in the zebrafish fin, whose distribution correlates with fluid dynamics.

Dominant structural defects in amorphous silicon.

The nature of disorder in amorphous silicon (a-Si) is explored by investigating the spatial arrangement and energies of coordination defects in a numerical model. Spatial correlations between structural defects are examined on the basis of a parameter that quantifies the probability for two sites to share a bond. Pentacoordinated atoms are found to be the dominant coordination defects. They show a tendency to cluster, and about 17% of them are linked through three-membered rings. As for tricoordinated sites, they are less numerous, and tend to be distant by at least two bond lengths. Typical local geometries associated to under and overcoordinated atoms are extracted from the model and described using partial bond angle distributions. An estimate of the formation energies of structural defects is provided. Using molecular-dynamics calculations, we simulate the implantation of high-energy atoms in the initial structure in order to study the effect of relaxation on the coordination defects and their environments.

Understanding subtle changes in medium-range order in amorphous silicon.

Based on a detailed study of the radial distribution function (RDF) of a model for amorphous silicon (a-Si), we address the relation between short-range rearrangements and an increase in medium-range order induced by thermal relaxation. Recent experimental measurements have shown that a small peak appears in the RDF around 4.7 Å upon annealing, along with other subtle changes, and this is attributed to ordering among the dihedral angles. We show that, although this is a possible explanation, an increase in short-range order (up to second neighbors) is not only necessary for these changes to occur, but could also be their sole cause. To clarify the nature of disorder in the amorphous system, correlations among dihedral and tetrahedral angles are examined. The bivariate probability distribution of these two variables reveals small correlations between dihedral and tetrahedral angles, associated with the staggered and eclipsed conformations. In the first case, bond angles around 112.5[Formula: see text] are favored versus 120[Formula: see text] in the second case. Bond angles between 95[Formula: see text] and 100[Formula: see text] are less probable in both conformations.