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(A) Lateral view of a skipjack tuna. The red line shows the longitudinal extent of the internalised red muscle. The arrow indicates 0.5L, where L is fork length (adapted from Joseph et al., 1988). (B) View of the left half of a skipkjack tuna, seen in transverse section at 0.5L, illustrating the placement of a sonomicrometer crystal in the internalised red muscle (shaded red). The flexible wire leads are anchored by sutures where they exit the skin, near the dorsal midline. Epaxial (upper) and hypaxial (lower) myotomal cones are represented by the concentric rings.
Source publication
Cyclic length changes in the internal red muscle of skipjack tuna (Katsuwonus pelamis) were measured using sonomicrometry while the fish swam in a water tunnel at steady speeds of 1.1-2.3 L s(-)(1), where L is fork length. These data were coupled with simultaneous electromyographic (EMG) recordings. The onset of EMG activity occurred at virtually t...
Context in source publication
Context 1
... Crystals 2 mm in diameter were constructed from piezoelectric ceramic (LTZ-2 Transducer Products Inc.). After soldering to lead wires, the crystals were lensed with a coating of polyester resin, giving a final thickness of 1.5 mm. Crystal pairs were inserted into the internal red muscle of the left side of the body from near the dorsal midline (Fig. 2). A 2 mm incision was made in the skin, and a puncture was made through the underlying white muscle with a 16 g hypodermic needle, precalibrated to the required depth. This provided an acceptable probability of placement into the internal red muscle, above the horizontal septum. Correct crystal alignment was ensured by monitoring the RF ...
Citations
... There have been many publications to describe the fluid movement around a swimming fish; literature by Gray (1933), Lighthill (1969), Webb (1975), andVideler (1993) provide a comprehensive portrayal of fish swimming. Furthermore, other studies (Rosen 1959, Blazka et al 1960, Hertel 1966, Aleyev 1977, Bone 1978, Lindsey 1978, Magnuson 1978, Webb 1978, 1984, Blake 1983, Webb and Weihs 1983, Müller et al 1997, Shadwick et al 1999, Triantafyllou et al 2000, Linden and Turner 2001, Lauder and Drucker 2002, Liao et al 2003, Colgate and Lynch 2004, Lauder and Tytell 2005, Fish and Lauder 2006, Wu 2011, Smits 2019) have reviewed and supplied insight into the kinematics, physiology, and hydrodynamics of fish locomotion as well as providing brief reviews of various aspects of previous studies. ...
Collective motion of organisms is a widespread phenomenon exhibited by many species, most commonly associated with colonial birds and schools of fish. The benefits of schooling behavior vary from defense against predators, increased feeding efficiency, and improved endurance. Schooling motions can be energetically beneficial as schools allow for channeling and vortex-based interactions, creating a less demanding stroke rate to sustain high swimming velocities and increased movement efficiency. Biomimetics is a fast-growing field, and there have been several attempts to quantify the hydrodynamics behind group dynamics and the subsequent benefits of increased maneuverability, which can be applied to unmanned vehicles and devices traveling in a group or swarm-like scenarios. Earlier efforts to understand these phenomena have been composed of physical experimentation and numerical simulations. This literature review examines the existing studies performed to understand the hydrodynamics of group collective motion inspired by schooling habits. Both numerical simulation and physical experimentation are discussed, and the benefits and drawbacks of the two approaches are compared to help future researchers and engineers expand on these models and concepts. This paper also identifies some of the limitations associated with different approaches to studies on fish schooling and suggests potential directions for future work.
... This versatility underlies the role of muscle in steady movement of diverse species including guinea fowl (Daley and Biewener, 2003;Higham and Biewener, 2008), fish (Rome et al., 1993;Shadwick et al., 1999;Young and Rome, 2001), flies (Tu and Dickinson, 1994), frogs (Richards and Biewener, 2007), turkeys (Gabaldón et al., 2004;Roberts et al., 1997), cockroaches (Ahn and Full, 2002;Ahn et al., 2006;Full et al., 1998), mice (James et al., 1995), rats (Ettema, 1996), katydids (Josephson, 1985a), scallops (Cheng and DeMont, 1997;Marsh and Olson, 1994), songbirds (Elemans et al., 2008), moths (George and Daniel, 2011;Sponberg and Daniel, 2012;Tu and Daniel, 2004), humans (Ishikawa and Komi, 2008) and many others. The same muscle can even adopt different functions (Box 1) depending on the strain cycle (see Glossary), phase of activation and other parameters (Ahn et al., 2003;Hedrick et al., 2003;Higham and Biewener, 2008;Josephson, 1985b;Roberts et al., 1997). ...
Muscle function during movement is more than a simple, linear transformation of neural activity into force. The classic work loop technique has pioneered our understanding of muscle, but typically only characterizes function during unperturbed movement cycles, such as those experienced during steady walking, running, swimming and flying. Yet perturbations away from steady movement often place greater demands on muscle structure and function and offer a unique window into muscle's broader capacity. Recently, studies in diverse organisms from cockroaches to humans have started to grapple with muscle function in unsteady (perturbed, transient and fluctuating) conditions, but the vast range of possible parameters and the challenge of connecting in vitro to in vivo experiments are daunting. Here, we review and organize these studies into two broad approaches that extend the classic work loop paradigm. First, in the top-down approach, researchers record length and activation patterns of natural locomotion under perturbed conditions, replay these conditions in isolated muscle work loop experiments to reveal the mechanism by which muscle mediates a change in body dynamics and, finally, generalize across conditions and scale. Second, in the bottom-up approach, researchers start with an isolated muscle work loop and then add structural complexity, simulated loads and neural feedback to ultimately emulate the muscle's neuromechanical context during perturbed movement. In isolation, each of these approaches has several limitations, but new models and experimental methods coupled with the formal language of control theory give several avenues for synthesizing an understanding of muscle function under unsteady conditions.
... The optimum undulation of a solitary thunniform swimmer, shown in figure 2, differs slightly from the reported midline kinematics for non-accelerated solitary thunniform swimmers [62,63]. While in both cases there is no significant head motion, the case presented here shows growing body and tail undulations, while thunniform swimmers in steady motion maintain a relatively straight body and only use their tail for propulsion [63,64]. ...
... The differences are attributed to the effect of acceleration. Indeed, most previous studies [62][63][64][65][66] focus on steady thunniform swimming, with unsteady swimming trends gaining traction only recently. For example, one previous study investigated the acceleration of a solitary thunniform swimmer from rest [20]. ...
Optimal fish array hydrodynamics in accelerating phalanx schools are investigated through a computational framework which combines high fidelity Computational Fluid Dynamics (CFD) simulations with a gradient free surrogate-based optimization algorithm. Critical parameters relevant to a phalanx fish school, such as midline kinematics, separation distance and phase synchronization, are investigated in light of efficient propulsion during an accelerating fish motion. Results show that the optimal midline kinematics in accelerating phalanx schools resemble those of accelerating solitary swimmers. The optimal separation distance in a phalanx school for thunniform biologically-inspired swimmers is shown to be around 2L (where L is the swimmer's total length). Furthermore, separation distance is shown to have a stronger effect, \textit{ceteris paribus}, on the propulsion efficiency of a school when compared to phase synchronization.
... The tendons of the myosepta that form the dorsal and ventral anterior cones between the red muscle fibers are weakly developed in tunas (Westneat and Wainwright, 2001;Long, et al., 2002). These weakly developed tendons within the myosepta uncouples the red muscle from the local body, allowing for more bending in the posterior region of the body and greater opportunity to produce mechanical work via the posterior oblique and great lateral tendons (Westneat et al., 1993;Knower, et al., 1999;Shadwick et al., 1999;Westneat and Wainwright, 2001;Long, 2002). Forces generated by the large muscle mass in the anterior region of the body are transferred via the tendon complex into the caudal fin of the tuna (Westneat et al., 2003;Westneat and Wainwright, 2001;Shadwick, 2003). ...
... The tendons of the myosepta that form the dorsal and ventral anterior cones between the red muscle fibers are weakly developed in tunas (Westneat and Wainwright, 2001;Long, et al., 2002). These weakly developed tendons within the myosepta uncouples the red muscle from the local body, allowing for more bending in the posterior region of the body and greater opportunity to produce mechanical work via the posterior oblique and great lateral tendons (Westneat et al., 1993;Knower, et al., 1999;Shadwick et al., 1999;Westneat and Wainwright, 2001;Long, 2002). Forces generated by the large muscle mass in the anterior region of the body are transferred via the tendon complex into the caudal fin of the tuna (Westneat et al., 2003;Westneat and Wainwright, 2001;Shadwick, 2003). ...
Tunas are known for exceptional swimming speeds due to their thunniform, lift-based propulsion, large muscle mass, and rigid fusiform body. A rigid body should restrict maneuverability in regard to turn radius and turn rate. To test if turning maneuvers by the Pacific bluefin tuna (Thunnus orientalis) are constrained by rigidity, captive animals were video recorded overhead as the animals routinely swam around a large circular tank or during feeding bouts. Turning performance was classified into three different types: 1) glide turn, where the tuna uses the caudal fin as a rudder, 2) powered turn, where the animal uses continuous near symmetrical strokes of the caudal fin through the turn, and 3) ratchet turn, where the overall global turn is completed by a series of small local turns by asymmetrical stokes of the caudal fin. Individual points of the rostrum, peduncle, and tip of the caudal fin were tracked and analyzed. Frame-by-frame analysis showed that the ratchet turn had the fastest turn rate for all points with a maximum of 302 deg s-1. During the ratchet turn, the rostrum exhibited a minimum global 0.38L turn radius. The local turn radii were only 18.6% of the global ratchet turn. The minimum turn radii ranged from 0.4L to 1.7L. Compared to the performance of other swimmers, the increased flexion of the peduncle and tail and the mechanics of turning behaviors used by tuna overcomes any constraints to turning performance from the rigidity of the anterior body morphology.
... Furthermore, sonomicrometry has demonstrated that the anterior part of the body bends dorsally like a beam during suction feeding , analogous to beam-like lateral body bending during body-caudal fin swimming in fishes (e.g. Shadwick et al., 1999;Wakeling and Johnston, 1999). Beam-like dorsal bending imposes a dorsoventral gradient of longitudinal strain in the epaxial muscle mass that is similar, but orthogonal, to the mediolateral strain gradient generated by lateral bending in locomotion. ...
Suction feeding in ray-finned fishes requires substantial muscle power for fast and forceful prey capture. The axial musculature located immediately behind the head has been long known to contribute some power for suction feeding, but recent XROMM and fluoromicrometry studies found nearly all the axial musculature (over 80%) provides effectively all (90-99%) of the power for high-performance suction feeding. The dominance of axial power suggests a new framework for studying the musculoskeletal biomechanics of fishes: the form and function of axial muscles and bones should be analysed for power production in feeding (or at least as a compromise between swimming and feeding), and cranial muscles and bones should be analysed for their role in transmitting axial power and coordinating buccal expansion. This new framework is already yielding novel insights, as demonstrated in four species for which suction power has now been measured. Interspecific comparisons suggest high suction power can be achieved in different ways: increasing the magnitude of suction pressure or the rate of buccal volume change, or both (as observed in the most powerful of these species). Our framework suggests that mechanical and evolutionary interactions between the head and the body, and between the swimming and feeding roles of axial structures, may be fruitful areas for continued study.
... We currently have little understanding of how complicated hydrodynamics at river confluences affect the fish locomotion and spatial distribution in the local basin, in contrast to the extensive literature available for studies of fish swimming in steady flow or turbulence with relatively simple structures [105][106][107][108][109][110][111][112][113][114][115] . The flow complexity metrics which can calculate the velocity gradient, kinetic energy change and eddy intensity of local water flow were widely used to describe the different behaviors of fish and possible habitats [116][117] . ...
Confluences act as critical nodes in a river system. They affect hydrodynamics, sediment transport, bed morphology, and eco-hydraulics of the river system. Convergence of streams produces the complex mechanism of flow momentum and mass mixing which may affect the aquatic environment locally and even lasting for a long
distance downstream. The confluence creates a hotspot for the river system’s ecological change, which usually leads to changes in water temperature, suspended-sediment load, bed material, nutrient concentrations, water chemistry, and organic-matter content. Hence, the dynamics of river confluences are very complex and have critical effects on river system’s water environment and ecology. For this reason, a review summarizing turbulent flow, sediment transport, morphological-dynamics, mixing processes, and their effects on the ecology of the aquatic environment at river confluences is in order. A future research agenda and opportunities pertinent to river confluence are vitally emphasized as a multidisciplinary research topic.
... Virtually every component of the locomotor system in fishes has some degree of flexibility and undergoes both bending and longitudinal strain during swimming [96][97][98][99][100][101][102]. Undulatory motion of the fish body results in an obvious wave of bending that progresses from head to tail during forward locomotion [103][104][105]. ...
One of the emerging themes of fish-inspired robotics is flexibility. Adding flexibility to the body, joints, or fins of fish-inspired robots can significantly improve thrust and/or efficiency during locomotion. However, the optimal stiffness depends on variables such as swimming speed, so there is no one ‘best’ stiffness that maximizes efficiency in all conditions. Fish are thought to solve this problem by using muscular activity to tune their body and fin stiffness in real-time. Inspired by fish, some recent robots sport polymer actuators, adjustable leaf springs, or artificial tendons that tune stiffness mechanically. Models and water channel tests are providing a theoretical framework for stiffness-tuning strategies that devices can implement. The strategies can be thought of as analogous to car transmissions, which allow users to improve efficiency by tuning gear ratio with driving speed. We provide an overview of the latest discoveries about (1) the propulsive benefits of flexibility, particularly tunable flexibility, and (2) the mechanisms and strategies that fish and fish-inspired robots use to tune stiffness while swimming.
... Many studies historically provide a good amount of knowledge on fish locomotion in still water or in flows with relatively simple structures (e.g., Gray, 1933;Lauder & Drucker, 2002;Müller et al., 1997;Shadwick et al., 1999;Webb, 1975). These studies mainly focused on how fish consumed energy and produced thrust to swim forward against the resistance offered by water. ...
Knowledge of locomotion of fish near river confluences is important for prediction of fish distribution in a river network. The flow separation zone near the confluence of a river network is a favorite habitat and feeding place for silver carp, which is one of the four major species of Chinese carp and usually provides positive rheotaxis to water flow. In the current study, a series of laboratory experiments were done to determine the behavioral responses of juvenile silver carp to the hydrodynamic forces near the separation zone of a channel confluence. The locomotion and trajectory of juvenile silver carp were recorded by infrared thermal imaging, while the flow velocity field near the separation zone was measured by a Particle Image Velocimetry (PIV) system. A total of 60 juvenile silver carp were released near the separation zone among which 40 carp swam in the upstream direction. Amongst them, 24 carp swam to the tributary and the remaining 16 swam into the main channel. Almost all these 24 carp travelled initially along the boundary of the separation zone near the corner, where flow shear was strongest, and then swam to the tributary. Instead of avoiding zones of strong vorticity, they chose and followed a trajectory along which the flow vorticity was large. On encountering these vortical flows, they increased the tail-beat frequency and decreased the tail-beat amplitude to maintain body stability. These observations provide important knowledge on locomotion of fish near river confluences and are beneficial for the fish habitat protection.
... These results on the intermuscular distribution of elements are consistent with other studies of tuna for both THg (Ando et al., 2008;Bosch et al., 2016;Itano et al., 1977;Vieira et al., 2017) and Se (Itano et al., 1977;Ralston et al., 2019). The obtained differences in THg and Se concentrations between red and white muscles are likely the consequence of differences in muscle composition (Crawford, 1972) and function (Moyle and Cech, 1996;Patterson and Goldspink, 1972;Shadwick et al., 1999). Red muscles have a more abundant blood supply and substantially higher haemoglobin and myoglobin levels than white muscles (Crawford, 1972), and hence more sulfhydryl-containing molecules such as cysteine and glutathione that have a high affinity for inorganic and organic forms of Hg (>90% of MeHg in the blood is bound to haemoglobin in the red blood cells; Berglund et al. (2005)). ...
... Red muscles have a more abundant blood supply and substantially higher haemoglobin and myoglobin levels than white muscles (Crawford, 1972), and hence more sulfhydryl-containing molecules such as cysteine and glutathione that have a high affinity for inorganic and organic forms of Hg (>90% of MeHg in the blood is bound to haemoglobin in the red blood cells; Berglund et al. (2005)). Alternatively, Bosch et al. (2016) suggested that, since Hg is continuously accumulated in fish by binding to protein sites, differences in inorganic mercury (IHg) and THg concentrations between red and white muscles of tuna result from differences in the composition and fiber development due to higher activity of red muscle (Shadwick et al., 1999). ...
This study examined total mercury (THg) and selenium (Se) levels in archive samples (white and red muscles, liver, gills) of the wild Atlantic bluefin tuna (ABFT) (Thunnus thynnus) (n = 18) captured in the central Adriatic Sea. The influence of fish size, age, and tissue type on element distribution was examined. There were significant differences in THg and Se levels, and Se:THg molar ratios among tissues. THg levels were highest in liver and lowest in gills (liver > red muscle > white muscle > gills), while Se levels were also highest in liver but lowest in white muscle (liver > red muscle > gills > white muscle). Se:THg molar ratios were highest in gills (22–82), intermediate in liver (11–29) and red muscle (7–36), and lowest in white muscle (1.7–7.6). Concentrations of THg in all tissues and Se in liver and caudal muscle were positively correlated with tuna age and size, while the Se:THg molar ratio in gills and all white muscles was negatively correlated with tuna age and size, indicating that the protective role of Se against THg is reduced in older specimens. The selenium health benefit values (HBVSe) were above zero in all tissues, indicating a small excess of Se after Hg sequestration. However, since the obtained HBVSe for edible tissues were near zero (0.01–0.04), and more than 70% of white muscle samples and all red muscle samples exceeded the EU regulatory limit for THg in fish muscle, it would be advisable to limit their intake in adults to one meal per month.
... Due to the complexity of musculoskeletal dynamics much of what is understood about skeletal muscle function during human locomotion has been indirectly inferred from correlations between anatomical classification, inverse dynamics analysis and electromyographic activity (EMG) analyses; however, there is a difficulty in developing causal relationships between muscle excitation and task performance during locomotion from these correlations alone (Kautz and Neptune, 2002). However, the implementation of mathematical arguments used to drive models have enabled researchers to gain valuable insight in those parameters and phenomena (Zajac, 1989;Winters, 1990) previously difficult to elucidate when solely relying on experimental data (Biewener et al., 1998;Shadwick et al., 1999). Using cycling as a paradigm to weave experimental data into model-driven simulations has provided insight into how forces interact between the leg segments and the external environment (Kautz et al., 1994;Raasch et al., 1997;Rankin and Neptune, 2008). ...
Optimisation of movement strategies during cycling is an area which has gathered a lot of attention over the past decade. Resolutions to augment performance have involved manipulations of bicycle mechanics, including chainring geometries. Elliptical chainrings are proposed to provide a greater effective diameter during the downstroke, manipulating mechanical leverage and resulting in greater power production during this period.
A review of the literature indicates that there is a pervasive gap in our understanding of how the theoretical underpinnings of elliptical chainrings might be translated to practical use. Despite reasonable theory of how these chainrings might enforce a variation in crank angular velocity and consequently alter force production, performance-based analyses have struggled to present evidence of this.
The purpose of this thesis was to provide a novel approach to this problem by combining experimental data with musculoskeletal modelling and evaluating how elliptical chainrings might influence crank reactive forces, joint kinematics, muscle-tendon unit behaviour and muscle activation. One main study was proposed to execute this analysis, and an anatomically constrained model was subsequently used to determine the joint kinematics and muscle-tendon unit behaviour. Bespoke elliptical chainrings were designed for this study and as such, different levels of chainring eccentricity (i.e. ratio of major to minor axis) and positioning against the crank were presented whilst controlling the influence of other variables known to affect the neuromuscular system such as cadence and load.
Findings presented in this thesis makes a new and major contribution in our understanding of the neuromusculoskeletal adaptations which occur when using elliptical chainrings, showing alterations in crank reaction force, muscle-tendon unit velocities, joint kinematics and muscle excitation over a range of cadences and loads, and provides direction for where the future of this research might be best applied.
Keywords: Elliptical chainrings; Cycling; Musculoskeletal modelling; Principal Component Analysis; Electromyography
