A continuous dynamic beam model for swimming fish

When a fish swims in water, muscle contraction, controlled by the nervous system, interacts with the body tissues and the surrounding fluid to yield the observed movement pattern of the body. A continuous dynamic beam model describing the bending moment balance on the body for such an interaction during swimming has been established. In the model a linear visco-elastic assumption is made for the passive behaviour of internal tissues, skin and backbone, and the unsteady fluid force acting on the swimming body is calculated by the 3D waving plate theory. The body bending moment distribution due to the various components, in isolation and acting together, is analysed. The analysis is based on the saithe (Pollachius virens), a carangiform swimmer. The fluid reaction needs a bending moment of increasing amplitude towards the tail and near-standing wave behaviour on the rear-half of the body. The inertial movement of the fish results from a wave of bending moment with increasing amplitude along the body and a higher propagation speed than that of body bending. In particular, the fluid reaction, mainly designed for propulsion, can provide a considerable force to balance the local momentum change of the body and thereby reduce the power required from the muscle. The wave of passive visco-elastic bending moment, with an amplitude distribution peaking a little before the mid-point of the fish, travels with a speed close to that of body bending. The calculated muscle bending moment from the whole dynamic system has a wave speed almost the same as that observed for EMG-onset and a starting instant close to that of muscle activation, suggesting a consistent matching between the muscle activation pattern and the dynamic response of the system in steady swimming. A faster wave of muscle activation, with a variable phase relation between the strain and activation cycle, appears to be designed to fit the fluid reaction and, to a lesser extent, the body inertia, and is limited by the passive internal tissues. Higher active stress is required from caudal muscle, as predicted from experimental studies on fish muscle. In general, the active force development by muscle does not coincide with the propulsive force generation on the tail. The stiffer backbone may play a role in transmitting force and deformation to maintain and adjust the movement of the body and tail in water.     

Hydrodynamics of burst swimming fish larvae; a conceptual model approach

Burst swimming of fish larvae is analysed from a hydrodynamic point of view. A picture of the expected flow pattern is presented based on information in literature on unsteady-flow patterns around obstacles in the intermediate Reynolds number region. It is shown that the acceleration stage of burst swimming under restricted conditions can be treated as a frictionless impulsive motion. The stream pattern resulting from this motion is presented and the efficiency of locomotion during the acceleration stage is calculated. The flow pattern in the post-acceleration stage is sketched and the origin of an interaction between the viscous and the reactive force contribution to the propulsive force in this stage is discussed. It is explained how this interaction can lead to an increase in propulsive efficiency. A conceptual model is developed describing the three stages in burst swimming locomotion: the acceleration stage, the post-acceleration stage and the gliding stage. Data from literature of the travel distance versus time relation of the common carp larva (Cyprinus carpio) of 5.5-mm length has been used to test the model results. The test appeared remarkably successful, and the model results for larger larvae up to 22 mm length are presented. The gliding distance as a function of larval length resulting from the model has been compared with experimental data from literature.     

Bioenergetic model of planktivorous fish feeding, growth and metabolism: theoretical optimum swimming speed of fish larvae

The feeding activity of an individual fish larva is described by an equation which includes parameters for the area successfully searched, probability of food capture multiplied by the cross-sectional perceptive visual field, larval swimming speed and the time required to consume a unit of food energy. The proportion of ingested food energy used for metabolism increases exponentially with increasing swimming speed. The model predicts that food consumption rate increases asymptotically whereas metabolic rate increases exponentially. This results in a predicted growth rate curve that reaches a maximum at a certain swimming speed and decreases at both higher and lower speeds. The model can be used to predict the influence of type of prey, prey density, water temperature etc. on larval growth. An expression describing how many hours per day fish larvae must forage in order to grow at a certain daily body weight gain allows the limits of environmental conditions for positive, zero and negative growth rate to be set. Results of simulations demonstrated that the optimum swimming speed for maximum growth of coregonid larvae increased with an increase in food density, decrease in water temperature or decrease of prey vulnerability. At optimum ‘theoretical’ swimming speed an increase in water temperature from 5 to 17° C required the food density to be increased from 20 to 80 copepods l^?1 in order to maintain a daily growth increment of 2%. The minimum Artemia density required for maintenance metabolism increased from 10 to 30 items 1¹ over the same temperature increase from 5 to 17° C, and food densities required for 8% growth rates were 26 and 56 Artemia nauplii l^?1 at 5 and 17° C, respectively. Contrary to previous findings, results of the present study suggest that metabolic rates of actively feeding fish larvae may be from 5 to 50 times the standard metabolic rate: earlier studies suggested that a factor of 2–3 may be generally applicable.     

A review of fish swimming mechanics and behaviour in altered flows

Fishes suspended in water are subject to the complex nature of three-dimensional flows. Often, these flows are the result of abiotic and biotic sources that alter otherwise uniform flows, which then have the potential to perturb the swimming motions of fishes. The goal of this review is to highlight key studies that have contributed to a mechanistic and behavioural understanding of how perturbing flows affect fish. Most of our understanding of fish behaviour in turbulence comes from observations of natural conditions in the field and laboratory studies employing controlled perturbations, such as vortices generated in the wake behind simple geometric objects. Laboratory studies have employed motion analysis, flow visualization, electromyography, respirometry and sensory deprecation techniques to evaluate the mechanisms and physiological costs of swimming in altered flows. Studies show that flows which display chaotic and wide fluctuations in velocity can repel fishes, while flows that have a component of predictability can attract fishes. The ability to maintain stability in three-dimensional flows, either actively with powered movements or passively using the posture and intrinsic compliance of the body and fins, plays a large role in whether fish seek out or avoid turbulence. Fish in schools or current-swept habitats can benefit from altered flows using two distinct though not mutually exclusive mechanisms: flow refuging (exploiting regions of reduced flow relative to the earth frame of reference) and vortex capture (harnessing the energy of environmental vortices). Integrating how the physical environment affects organismal biomechanics with the more complex issue of behavioural choice requires consideration beyond simple body motions or metabolic costs. A fundamental link between these two ways of thinking about animal behaviour is how organisms sense and process information from the environment, which determines when locomotor behaviour is initiated and modulated. New data are presented here which show that behaviour changes in altered flows when either the lateral line or vision is blocked, showing that fish rely on multi-modal sensory inputs to negotiate complex flow environments. Integrating biomechanics and sensory biology to understand how fish swim in turbulent flow at the organismal level is necessary to better address population-level questions in the fields of fisheries management and ecology.     

A histochemical study of the swimming musculature of Antarctic fish

The kiore, once common throughout New Zealand, had disappeared from most of the country by the end of the 19th century, and is now found only on certain offshore islands and in areas of Fiordland where at least one of the three introduced European rodent species is absent. It is usually accepted that the kiore was displaced by ship rats (Rattus r. rattus) and Norway rats (R. norvegicus). However, recent investigations on Stewart Island have revealed kiore, ship rats, and Norway rats living in close association, but in the absence of mice (Mus musculus). In the area studied the kiore seemed to inhabit mainly grassland. Re‐examination of possible reasons for the decline of the kiore strongly suggests that competition from mice has been a major contributing factor. It seems that in New Zealand a niche no longer exists for kiore once mice, ship rats, and Norway rats have all become established.     

Modelling energetic costs of fish swimming

The oxygen consumption rates of two cyprinid fishes, carp (Cyprinus carpio L.) and roach (Rutilus rutilus (L.)), were analysed for a wide range of body mass and swimming speed by computerized intermittent-flow respirometry. Bioenergetic models were derived, based on fish mass (M) and swimming speed (U), to predict the minimal speed and mass-specific active metabolic rate (AMR) in these fishes (AMR=aMbUc). Mass and speed together explained more than 90% of the variance in total swimming costs in both cases. The derived models show that carp consume far more oxygen at a specific speed and body mass, thus being less efficient in energy use during swimming than roach. It was further found that in carp (AMR=0.02M0.8U0.95) the metabolic increment during swimming is more strongly effected by speed, whereas in roach (AMR=0.02M0.93U0.6) it is more strongly effected by body mass. The different swimming traits of carp and roach are suitable for their respective lifestyles and ecological demands.     

A model for swimming unipolar spirilla

A mathematical model employing slender body theory is constructed for a unipolar Spirillum volutans cell with the model cell allowed to move unconstrainedly in the fluid. The results are compared with observation and previous studies and the effects of varying cell dimensions are investigated.     

Energetic advantages of burst swimming of fish

It is shown theoretically that fish can swim more efficiently by alternating periods of accelerated motion and powerless gliding. Analysis of the mechanics of swimming shows that large savings of over 50% in the energy required to traverse a given distance can be obtained by such means. In calculations based upon measured data for salmon and haddock, the possibility of range increases of up to three times the range at constant speed are demonstrated.     

Systems-level modeling of neuronal circuits for leech swimming

This paper describes a mathematical model of the neuronal central pattern generator (CPG) that controls the rhythmic body motion of the swimming leech. The systems approach is employed to capture the neuronal dynamics essential for generating coordinated oscillations of cell membrane potentials by a simple CPG architecture with a minimal number of parameters. Based on input/output data from physiological experiments, dynamical components (neurons and synaptic interactions) are first modeled individually and then integrated into a chain of nonlinear oscillators to form a CPG. We show through numerical simulations that the values of a few parameters can be estimated within physiologically reasonable ranges to achieve good fit of the data with respect to the phase, amplitude, and period. This parameter estimation leads to predictions regarding the synaptic coupling strength and intrinsic period gradient along the nerve cord, the latter of which agrees qualitatively with experimental observations.     

Light-oriented swimming of schooling fish in continuous channels

Synopsis Fish groups were tested both in a circular and in a figure eight-shaped channel. In both cases fish showed a long lasting, constant direction swimming provided that illumination was maintained at a constant angle around the channel. In the circular channel, fish did not reverse direction, as would be expected, when light angle was shifted from one side to the other in the channel. However, direction reversals did occur when these illumination shifts were performed on the eight-shaped channel. We suggest that constant-oriented swimming reflects a sun-compass oriented behavior, but swimming at a constant angle in the circular channel produces an irreversible disarrangement of the inertial-orientation system, which does not occur in the eight-shaped channel due to the geometrical relationship between the light and the shape of the channel.     

Wave energy and swimming performance shape coral reef fish assemblages

Physical factors often have an overriding influence on the distribution patterns of organisms, and can ultimately shape the long-term structure of communities. Although distribution patterns in sessile marine organisms have frequently been attributed to functional characteristics interacting with wave-induced water motion, similar evidence for mobile organisms is lacking. Links between fin morphology and swimming performance were examined in three diverse coral reef fish families from two major evolutionary lineages. Among-habitat variation in morphology and performance was directly compared with quantitative values of wave-induced water motion from seven coral reef habitats of different depth and wave exposure on the Great Barrier Reef. Fin morphology was strongly correlated with both field and experimental swimming speeds in all three families. The range of observed swimming speeds coincided closely with the magnitude of water velocities commonly found on coral reefs. Distribution patterns in all three families displayed highly congruent relationships between fin morphology and wave-induced water motion. Our findings indicate a general functional relationship between fin morphology and swimming performance in labriform-swimming fishes, and provide quantitative evidence that wave energy may directly influence the assemblage structure of coral reef fishes through interactions with morphology and swimming performance.     

A compact cartilaginous fish model genome

The genomes of several vertebrates, including six mammals, the chicken, Xenopus and four ray-finned fishes have been sequenced or are currently being sequenced to provide a better understanding of the human genome through comparative analysis. However, this list does not include cartilaginous fishes, which are the most basal living jawed vertebrates [1]. The genomes of the current ‘popular’ cartilaginous fishes such as the nurse shark, dogfish, and horn shark are larger than the human genome (∼3800 Mb to 7000 Mb) [2], and are not attractive for whole-genome sequencing. Here, we report the characterization of the relatively small genome (1200 Mb) of a cartilaginous fish, the elephant fish (Callorhinchus milii), and propose it as a model for whole-genome sequencing.     

A community model of mutually learning neuronal nets

A model of a community is suggested whose members are formal neuron nets interacting by signals exchange. As a signal each net can emit an image formed by it when recognising the preceding signal. The emitted signal comes to the inputs of other nets and is used as their initial state for the recognition process. The collective dynamics of such model is discussed for the case of non-learning nets. Possible algorithm of mutual learning of the nets in them course of signals exchange is considered.