Anthropogenic stressors and the marine environment: From sources and impacts to solutions and mitigation.

Human-released contaminants are often poorly understood wholistically in marine ecosystems. This review examines the sources, pathways, impacts on marine animals, and mitigation strategies of five pollutants (plastics, per- and polyfluoroalkyl substances, bisphenol compounds, ethynylestradiol, and petroleum hydrocarbons). Both abiotic and biotic mechanisms contribute to all five contaminants' movement. These pollutants cause short- and long-term effects on many biological processes genetically, molecularly, neurologically, physiologically, reproductively, and developmentally. We explore the extension of adverse outcome pathways to ecosystem effects by considering known inter-generational and trophic relations resulting in large-scale direct and indirect impacts. In doing so, we develop an understanding of their roles as environmental stressors in marine environments for targeted mitigation and future work. Ecosystems are interconnected and so international collaboration, standards, measures preceding mass production, and citizen involvement are required to protect and conserve marine life.

Flexibility is a hidden axis of biomechanical diversity in fishes.

Nearly all fish have flexible bodies that bend as a result of internal muscular forces and external fluid forces that are dynamically coupled with the mechanical properties of the body. Swimming is therefore strongly influenced by the body's flexibility, yet we do not know how fish species vary in their flexibility and in their ability to modulate flexibility with muscle activity. A more fundamental problem is our lack of knowledge about how any of these differences in flexibility translate into swimming performance. Thus, flexibility represents a hidden axis of diversity among fishes that may have substantial impacts on swimming performance. Although engineers have made substantial progress in understanding these fluid-structure interactions using physical and computational models, the last biological review of these interactions and how they give rise to fish swimming was carried out more than 20 years ago. In this Review, we summarize work on passive and active body mechanics in fish, physical models of fish and bioinspired robots. We also revisit some of the first studies to explore flexural stiffness and discuss their relevance in the context of more recent work. Finally, we pose questions and suggest future directions that may help reveal important links between flexibility and swimming performance.

The Hydrodynamics of Jellyfish Swimming.

Jellyfish have provided insight into important components of animal propulsion, such as suction thrust, passive energy recapture, vortex wall effects, and the rotational mechanics of turning. These traits are critically important to jellyfish because they must propel themselves despite severe limitations on force production imposed by rudimentary cnidarian muscular structures. Consequently, jellyfish swimming can occur only by careful orchestration of fluid interactions. Yet these mechanics may be more broadly instructive because they also characterize processes shared with other animal swimmers, whose structural and neurological complexity can obscure these interactions. In comparison with other animal models, the structural simplicity, comparative energetic efficiency, and ease of use in laboratory experimentation allow jellyfish to serve as favorable test subjects for exploration of the hydrodynamic bases of animal propulsion. These same attributes also make jellyfish valuable models for insight into biomimetic or bioinspired engineeringof swimming vehicles. Here, we review advances in understanding of propulsive mechanics derived from jellyfish models as a pathway toward the application of animal mechanics to vehicle designs.

Airfoil-like mechanics generate thrust on the anterior body of swimming fishes.

The anterior body of many fishes is shaped like an airfoil turned on its side. With an oscillating angle to the swimming direction, such an airfoil experiences negative pressure due to both its shape and pitching movements. This negative pressure acts as thrust forces on the anterior body. Here, we apply a high-resolution, pressure-based approach to describe how two fishes, bluegill sunfish (Lepomis macrochirus Rafinesque) and brook trout (Salvelinus fontinalis Mitchill), swimming in the carangiform mode, the most common fish swimming mode, generate thrust on their anterior bodies using leading-edge suction mechanics, much like an airfoil. These mechanics contrast with those previously reported in lampreys-anguilliform swimmers-which produce thrust with negative pressure but do so through undulatory mechanics. The thrust produced on the anterior bodies of these carangiform swimmers through negative pressure comprises 28% of the total thrust produced over the body and caudal fin, substantially decreasing the net drag on the anterior body. On the posterior region, subtle differences in body shape and kinematics allow trout to produce more thrust than bluegill, suggesting that they may swim more effectively. Despite the large phylogenetic distance between these species, and differences near the tail, the pressure profiles around the anterior body are similar. We suggest that such airfoil-like mechanics are highly efficient, because they require very little movement and therefore relatively little active muscular energy, and may be used by a wide range of fishes since many species have appropriately shaped bodies.

A pressure-based force and torque prediction technique for the study of fish-like swimming.

Many outstanding questions about the evolution and function of fish morphology are linked to swimming dynamics, and a detailed knowledge of time-varying forces and torques along the animal's body is a key component in answering many of these questions. Yet, quantifying these forces and torques experimentally represents a major challenge that to date prevents a full understanding of fish-like swimming. Here, we develop a method for obtaining these force and torque data non-invasively using standard 2D digital particle image velocimetry in conjunction with a pressure field algorithm. We use a mechanical flapping foil apparatus to model fish-like swimming and measure forces and torques directly with a load cell, and compare these measured values to those estimated simultaneously using our pressure-based approach. We demonstrate that, when out-of-plane flows are relatively small compared to the planar flow, and when pressure effects sufficiently dominate shear effects, this technique is able to accurately reproduce the shape, magnitude, and timing of locomotor forces and torques experienced by a fish-like swimmer. We conclude by exploring of the limits of this approach and its feasibility in the study of freely-swimming fishes.

Performance variation due to stiffness in a tuna-inspired flexible foil model.

Tuna are fast, economical swimmers in part due to their stiff, high aspect ratio caudal fins and streamlined bodies. Previous studies using passive caudal fin models have suggested that while high aspect ratio tail shapes such as a tuna's generally perform well, tail performance cannot be determined from shape alone. In this study, we analyzed the swimming performance of tuna-tail-shaped hydrofoils of a wide range of stiffnesses, heave amplitudes, and frequencies to determine how stiffness and kinematics affect multiple swimming performance parameters for a single foil shape. We then compared the foil models' kinematics with published data from a live swimming tuna to determine how well the hydrofoil models could mimic fish kinematics. Foil kinematics over a wide range of motion programs generally showed a minimum lateral displacement at the narrowest part of the foil, and, immediately anterior to that, a local area of large lateral body displacement. These two kinematic patterns may enhance thrust in foils of intermediate stiffness. Stiffness and kinematics exhibited subtle interacting effects on hydrodynamic efficiency, with no one stiffness maximizing both thrust and efficiency. Foils of intermediate stiffnesses typically had the greatest coefficients of thrust at the highest heave amplitudes and frequencies. The comparison of foil kinematics with tuna kinematics showed that tuna motion is better approximated by a zero angle of attack foil motion program than by programs that do not incorporate pitch. These results indicate that open questions in biomechanics may be well served by foil models, given appropriate choice of model characteristics and control programs. Accurate replication of biological movements will require refinement of motion control programs and physical models, including the creation of models of variable stiffness.

Effects of non-uniform stiffness on the swimming performance of a passively-flexing, fish-like foil model.

Simple mechanical models emulating fish have been used recently to enable targeted study of individual factors contributing to swimming locomotion without the confounding complexity of the whole fish body. Yet, unlike these uniform models, the fish body is notable for its non-uniform material properties. In particular, flexural stiffness decreases along the fish's anterior-posterior axis. To identify the role of non-uniform bending stiffness during fish-like propulsion, we studied four foil model configurations made by adhering layers of plastic sheets to produce discrete regions of high (5.5 × 10(-5) Nm(2)) and low (1.9 × 10(-5) Nm(2)) flexural stiffness of biologically-relevant magnitudes. This resulted in two uniform control foils and two foils with anterior regions of high stiffness and posterior regions of low stiffness. With a mechanical flapping foil controller, we measured forces and torques in three directions and quantified swimming performance under both heaving (no pitch) and constant 0° angle of attack programs. Foils self-propelled at Reynolds number 21 000-115 000 and Strouhal number ∼0.20-0.25, values characteristic of fish locomotion. Although previous models have emphasized uniform distributions and heaving motions, the combination of non-uniform stiffness distributions and 0° angle of attack pitching program was better able to reproduce the kinematics of freely-swimming fish. This combination was likewise crucial in maximizing swimming performance and resulted in high self-propelled speeds at low costs of transport and large thrust coefficients at relatively high efficiency. Because these metrics were not all maximized together, selection of the 'best' stiffness distribution will depend on actuation constraints and performance goals. These improved models enable more detailed, accurate analyses of fish-like swimming.

Bending rules for animal propulsion.

Animal propulsors such as wings and fins bend during motion and these bending patterns are believed to contribute to the high efficiency of animal movements compared with those of man-made designs. However, efforts to implement flexible designs have been met with contradictory performance results. Consequently, there is no clear understanding of the role played by propulsor flexibility or, more fundamentally, how flexible propulsors should be designed for optimal performance. Here we demonstrate that during steady-state motion by a wide range of animals, from fruit flies to humpback whales, operating in either air or water, natural propulsors bend in similar ways within a highly predictable range of characteristic motions. By providing empirical design criteria derived from natural propulsors that have convergently arrived at a limited design space, these results provide a new framework from which to understand and design flexible propulsors.