Increase In Swimming Speed Research Articles

Distinct Domains of CheA Confer Unique Functions in Chemotaxis and Cell Length in Azospirillum brasilense Sp7.

Chemotaxis is the movement of cells in response to gradients of diverse chemical cues. Motile bacteria utilize a conserved chemotaxis signal transduction system to bias their motility and navigate through a gradient. A central regulator of chemotaxis is the histidine kinase CheA. This cytoplasmic protein interacts with membrane-bound receptors, which assemble into large polar arrays, to propagate the signal. In the alphaproteobacterium Azospirillum brasilense, Che1 controls transient increases in swimming speed during chemotaxis, but it also biases the cell length at division. However, the exact underlying molecular mechanisms for Che1-dependent control of multiple cellular behaviors are not known. Here, we identify specific domains of the CheA1 histidine kinase implicated in modulating each of these functions. We show that CheA1 is produced in two isoforms: a membrane-anchored isoform produced as a fusion with a conserved seven-transmembrane domain of unknown function (TMX) at the N terminus and a soluble isoform similar to prototypical CheA. Site-directed and deletion mutagenesis combined with behavioral assays confirm the role of CheA1 in chemotaxis and implicate the TMX domain in mediating changes in cell length. Fluorescence microscopy further reveals that the membrane-anchored isoform is distributed around the cell surface while the soluble isoform localizes at the cell poles. Together, the data provide a mechanism for the role of Che1 in controlling multiple unrelated cellular behaviors via acquisition of a new domain in CheA1 and production of distinct functional isoforms.IMPORTANCE Chemotaxis provides a significant competitive advantage to bacteria in the environment, and this function has been transferred laterally multiple times, with evidence of functional divergence in different genomic contexts. The molecular principles that underlie functional diversification of chemotaxis in various genomic contexts are unknown. Here, we provide a molecular mechanism by which a single CheA protein controls two unrelated functions: chemotaxis and cell length. Acquisition of this multifunctionality is seemingly a recent evolutionary event. The findings illustrate a mechanism by which chemotaxis function may be co-opted to regulate additional cellular functions.

Optimizing the restored chemotactic behavior of anticancer agent Salmonella enterica serovar Typhimurium VNP20009

Bacteria, including strains of Salmonella, have been researched and applied as therapeutic cancer agents for centuries. Salmonella are particularly of interest due to their facultative anaerobic nature, facilitating colonization of differentially oxygenated tumor regions. Additionally, Salmonella can be manipulated with relative ease, resulting in the ability to attenuate the pathogen or engineer vectors for drug delivery. It was recently discovered that the anti-cancer Salmonella enterica serovar Typhimurium strain VNP20009 is lacking in chemotactic ability, due to a non-synonymous single nucleotide polymorphism in cheY. Replacing the mutated copy of cheY with the wild-type sequence restored chemotaxis to 70% of the parental strain. We aimed to investigate further if chemotaxis of VNP20009 can be optimized. By restoring the gene msbB in VNP20009 cheY+, which confers attenuation by lipid A modification, we observed a 9% increase in swimming speed, 13% increase in swim plate performance, 19% increase in microfluidic device partitioning towards the attractant at the optimum concentration gradient, and mitigation of a non-motile cell subpopulation. We conclude that chemotaxis can be enhanced further but at the cost of changing one defining characteristic of VNP20009. A less compromised strain might be needed to employ for investigating bacterial chemotaxis in tumor interactions.

Synchronisation through learning for two self-propelled swimmers

The coordinated motion by multiple swimmers is a fundamental component in fish schooling. The flow field induced by the motion of each self-propelled swimmer implies non-linear hydrodynamic interactions among the members of a group. How do swimmers compensate for such hydrodynamic interactions in coordinated patterns? We provide an answer to this riddle though simulations of two, self-propelled, fish-like bodies that employ a learning algorithm to synchronise their swimming patterns. We distinguish between learned motion patterns and the commonly used a-priori specified movements, that are imposed on the swimmers without feedback from their hydrodynamic interactions. First, we demonstrate that two rigid bodies executing pre-specified motions, with an alternating leader and follower, can result in substantial drag-reduction and intermittent thrust generation. In turn, we study two self-propelled swimmers arranged in a leader-follower configuration, with a-priori specified body-deformations. These two self-propelled swimmers do not sustain their tandem configuration. The follower experiences either an increase or decrease in swimming speed, depending on the initial conditions, while the swimming of the leader remains largely unaffected. This indicates that a-priori specified patterns are not sufficient to sustain synchronised swimming. We then examine a tandem of swimmers where the leader has a steady gait and the follower learns to synchronize its motion, to overcome the forces induced by the leader’s vortex wake. The follower employs reinforcement learning to adapt its swimming-kinematics so as to minimize its lateral deviations from the leader’s path. Swimming in such a sustained synchronised tandem yields up to reduction in energy expenditure for the follower, in addition to a increase in its swimming-efficiency. The present results show that two self-propelled swimmers can be synchronised by adapting their motion patterns to compensate for flow-structure interactions. Moreover, swimmers can exploit the vortical structures of their flow field so that synchronised swimming is energetically beneficial.

Fine-scale foraging movements by fish-eating killer whales (Orcinus orca) relate to the vertical distributions and escape responses of salmonid prey (Oncorhynchus spp.)

BackgroundWe sought to quantitatively describe the fine-scale foraging behavior of northern resident killer whales (Orcinus orca), a population of fish-eating killer whales that feeds almost exclusively on Pacific salmon (Oncorhynchus spp.). To reconstruct the underwater movements of these specialist predators, we deployed 34 biologging Dtags on 32 individuals and collected high-resolution, three-dimensional accelerometry and acoustic data. We used the resulting dive paths to compare killer whale foraging behavior to the distributions of different salmonid prey species. Understanding the foraging movements of these threatened predators is important from a conservation standpoint, since prey availability has been identified as a limiting factor in their population dynamics and recovery.ResultsThree-dimensional dive tracks indicated that foraging (N = 701) and non-foraging dives (N = 10,618) were kinematically distinct (Wilks’ lambda: λ16 = 0.321, P < 0.001). While foraging, killer whales dove deeper, remained submerged longer, swam faster, increased their dive path tortuosity, and rolled their bodies to a greater extent than during other activities. Maximum foraging dive depths reflected the deeper vertical distribution of Chinook (compared to other salmonids) and the tendency of Pacific salmon to evade predators by diving steeply. Kinematic characteristics of prey pursuit by resident killer whales also revealed several other escape strategies employed by salmon attempting to avoid predation, including increased swimming speeds and evasive maneuvering.ConclusionsHigh-resolution dive tracks reconstructed using data collected by multi-sensor accelerometer tags found that movements by resident killer whales relate significantly to the vertical distributions and escape responses of their primary prey, Pacific salmon.

Selection on escape performance during ecological speciation driven by predation

Despite the many study systems in which predation has played a major role in phenotypic diversification and speciation, the underlying selective regimes imposed by different predator assemblages have rarely been quantified. We did so for the damselfly genus Enallagma which strongly diverged in antipredator traits when the ancestral species occupying lakes containing fish (hereafter fish lakes) repeatedly invaded fishless lakes with dragonfly larvae as top predators (hereafter dragonfly lakes). In two selection experiments in field enclosures we quantified the selection on two key escape traits of two fish-lake Enallagma species associated with survival selection by fish in the ancestral fish lakes and by dragonfly predators in the invaded fishless, dragonfly lakes. In accordance with the different hunting modes, fish imposed selection for a decreased swimming propensity while dragonfly larvae imposed selection for increased swimming speed in one of the two species. In two complementary quantitative genetic rearing experiments, we found relatively low but significant broad-sense heritabilities for both escape traits. Integrating these estimates for the selection coefficients and the heritabilities suggests that the evolutionary increase in swimming speed associated with the habitat shift may have occurred rapidly. Our study suggests that the phenotypic evolution of ecologically important traits related to habitat shifts may occur at an ecological timescale.

Effective Propulsion in Swimming: Grasping the Hydrodynamics of Hand and Arm Movements.

In this paper, a literature review is presented regarding the hydrodynamic effects of different hand and arm movements during swimming with the aim to identify lacunae in current methods and knowledge, and to distil practical guidelines for coaches and swimmers seeking to increase swimming speed. Experimental and numerical studies are discussed, examining the effects of hand orientation, thumb position, finger spread, sculling movements, and hand accelerations during swimming, as well as unsteady properties of vortices due to changes in hand orientation. Collectively, the findings indicate that swimming speed may be increased by avoiding excessive sculling movements and by spreading the fingers slightly. In addition, it appears that accelerating the hands rather than moving them at constant speed may be beneficial, and that (in front crawl swimming) the thumb should be abducted during entry, catch, and upsweep, and adducted during the pull phase. Further experimental and numerical research is required to confirm these suggestions and to elucidate their hydrodynamic underpinnings and identify optimal propulsion techniques. To this end, it is necessary that the dynamical motion and resulting unsteady effects are accounted for, and that flow visualization techniques, force measurements, and simulations are combined in studying those effects.

Effect of Turbidity on the Swimming Performance of the Golden Shiner,Notemigonus crysoleucas

Of the many environmental factors that could affect swimming performance and critical swimming speed (Ucrit), turbidity has received relatively little attention. The purpose of this study was to examine the effects of turbidity on Golden Shiner (Notemigonus crysoleucas) swimming performance through the analysis of critical swimming speed. It was found that turbidity had an effect on critical swimming speeds; critical swimming speeds increased significantly with rising turbidity values. At the highest turbidity levels reported in this paper, Golden Shiner experienced a 56.5% increase in critical swimming speed over controls swimming in clear water. We likewise observed an increase in swimming efficiency and cost of transport with increasing turbidity. We suggest that a hydrodynamic effect may be the most likely explanation for the increase in performance and swimming efficiency observed in this study.

Not too slow, not too fast: water currents affect group structure, aggression and welfare in post-smolt Atlantic salmon Salmo salar

Increased swimming speed of Atlantic salmon is generally considered an improvement to welfare under aquaculture settings, as group structure is improved and agonistic behaviour re- duced. As such, establishing fish farms in exposed areas with fast water current velocities should be favourable. However, at some locations, velocities exceed what is known as preferable for salmonids, and this may compromise fish welfare. In this study, behaviour and fin erosion were observed on post-smolt salmon stocked at 39 kg m �3 in raceways at 3 water current velocities: fast (1.5 body lengths (BL) s �1 ), moderate (0.8 BL s �1 ) and slow (0.2 BL s �1 ). Movements that affect group structure and interactions between individuals varied by up to 20-fold between velocities. A behavioural change occurred directly after velocities were set. Severe fin erosion decreased over time in all groups, but new injuries increased almost 3-fold in the faster-velocity group. Our results suggest that moderate velocity is ideal from a welfare perspective. At slow velocity, higher frequency of structural movements and between-individual interactions could be stressful for the fish. At faster velocity, the fish have to focus on swimming, which could increase unintentional collisions with obstacles and other individuals and result in new fin erosion. Our results suggest that management of water currents may be an effective way of controlling behaviour and may thereby improve welfare.

Ontogenetic changes in larval swimming and orientation of pre-competent sea urchin Arbacia punctulata in turbulence.

ABSTRACTMany marine organisms have complex life histories, having sessile adults and relying on the planktonic larvae for dispersal. Larvae swim and disperse in a complex fluid environment and the effect of ambient flow on larval behavior could in turn impact their survival and transport. However, to date, most studies on larvae–flow interactions have focused on competent larvae near settlement. We examined the importance of flow on early larval stages by studying how local flow and ontogeny influence swimming behavior in pre-competent larval sea urchins, Arbacia punctulata. We exposed larval urchins to grid-stirred turbulence and recorded their behavior at two stages (4- and 6-armed plutei) in three turbulence regimes. Using particle image velocimetry to quantify and subtract local flow, we tested the hypothesis that larvae respond to turbulence by increasing swimming speed, and that the increase varies with ontogeny. Swimming speed increased with turbulence for both 4- and 6-armed larvae, but their responses differed in terms of vertical swimming velocity. 4-Armed larvae swam most strongly upward in the unforced flow regime, while 6-armed larvae swam most strongly upward in weakly forced flow. Increased turbulence intensity also decreased the relative time that larvae spent in their typical upright orientation. 6-Armed larvae were tilted more frequently in turbulence compared with 4-armed larvae. This observation suggests that as larvae increase in size and add pairs of arms, they are more likely to be passively re-oriented by moving water, rather than being stabilized (by mechanisms associated with increased mass), potentially leading to differential transport. The positive relationship between swimming speed and larval orientation angle suggests that there was also an active response to tilting in turbulence. Our results highlight the importance of turbulence to planktonic larvae, not just during settlement but also in earlier stages through morphology–flow interactions.

Behavioural responses to sound exposure in captivity by two fish species with different hearing ability

Underwater sound generated by human activities is increasing in, on and near aquatic environments. Such anthropogenic noise can induce artificially elevated ambient sound levels and cause various detrimental effects in fish, such as temporary or permanent hearing loss, masking of relevant acoustic signals and cues or behavioural changes that may have fitness consequences. Also, captive fish are often exposed to noisy conditions, which may have consequences for production in aquaculture, biases in scientific results in laboratories or welfare in hobby aquaria. However, we still have limited insight into how fish cope with artificial sound exposure and how species differ in sensitivity. Here, we compared zebrafish, Danio rerio, and Lake Victoria cichlids, Haplochromis piceatus, the former being sensitive to lower absolute thresholds and wider spectral ranges than the latter. Experimental sound exposure induced a significant reduction in swimming speed in the first minute of exposure for both species in captive conditions. Furthermore, zebrafish showed clear startle response behaviour with the onset of the sound exposure leading to an initial, brief increase in swimming speed, which was not found for the cichlids. Neither species showed spatial shifts away from the active speaker in the horizontal plane, but cichlids shifted downwards to spend more time in the bottom area of the fish tank after the onset of sound exposure, while zebrafish retained their average swimming height during the same exposure levels. Our results show that sound exposure can cause both similar and species-specific responses in two fish species and that these responses are not obviously related to differences in their hearing ability.

Azospirillum brasilense Chemotaxis Depends on Two Signaling Pathways Regulating Distinct Motility Parameters.

The genomes of most motile bacteria encode two or more chemotaxis (Che) systems, but their functions have been characterized in only a few model systems. Azospirillum brasilense is a motile soil alphaproteobacterium able to colonize the rhizosphere of cereals. In response to an attractant, motile A. brasilense cells transiently increase swimming speed and suppress reversals. The Che1 chemotaxis pathway was previously shown to regulate changes in the swimming speed, but it has a minor role in chemotaxis and root surface colonization. Here, we show that a second chemotaxis system, named Che4, regulates the probability of swimming reversals and is the major signaling pathway for chemotaxis and wheat root surface colonization. Experimental evidence indicates that Che1 and Che4 are functionally linked to coordinate changes in the swimming motility pattern in response to attractants. The effect of Che1 on swimming speed is shown to enhance the aerotactic response of A. brasilense in gradients, likely providing the cells with a competitive advantage in the rhizosphere. Together, the results illustrate a novel mechanism by which motile bacteria utilize two chemotaxis pathways regulating distinct motility parameters to alter movement in gradients and enhance the chemotactic advantage. Chemotaxis provides motile bacteria with a competitive advantage in the colonization of diverse niches and is a function enriched in rhizosphere bacterial communities, with most species possessing at least two chemotaxis systems. Here, we identify the mechanism by which cells may derive a significant chemotactic advantage using two chemotaxis pathways that ultimately regulate distinct motility parameters.

Sound exposure changes European seabass behaviour in a large outdoor floating pen: Effects of temporal structure and a ramp-up procedure

Underwater sound from human activities may affect fish behaviour negatively and threaten the stability of fish stocks. However, some fundamental understanding is still lacking for adequate impact assessments and potential mitigation strategies. For example, little is known about the potential contribution of the temporal features of sound, the efficacy of ramp-up procedures, and the generalisability of results from indoor studies to the outdoors. Using a semi-natural set-up, we exposed European seabass in an outdoor pen to four treatments: 1) continuous sound, 2) intermittent sound with a regular repetition interval, 3) irregular repetition intervals and 4) a regular repetition interval with amplitude ‘ramp-up’. Upon sound exposure, the fish increased swimming speed and depth, and swam away from the sound source. The behavioural readouts were generally consistent with earlier indoor experiments, but the changes and recovery were more variable and were not significantly influenced by sound intermittency and interval regularity. In addition, the ‘ramp-up’ procedure elicited immediate diving response, similar to the onset of treatment without a ‘ramp-up’, but the fish did not swim away from the sound source as expected. Our findings suggest that while sound impact studies outdoors increase ecological and behavioural validity, the inherently higher variability also reduces resolution that may be counteracted by increasing sample size or looking into different individual coping styles. Our results also question the efficacy of ‘ramp-up’ in deterring marine animals, which warrants more investigation.

An animal-borne active acoustic tag for minimally invasive behavioral response studies on marine mammals

There is a variety of evidence that increased anthropogenic noise (e.g., shipping, explosions, sonar) has a measureable effect on marine mammal species. Observed impacts range in severity from brief interruptions of basic life functions to physiological changes, acute injury, and even death. New research tools are needed to better measure and understand the potential effects of anthropogenic noise on marine mammals. Current behavioral response studies typically utilize ship-based sound sources to study potential acute behavioral responses in tagged animals experimentally exposed to noise. Integrating the sound source within animal-mounted passive acoustic and motion-sensing tags provides a novel tool for conducting additional highly controlled response studies. We developed and conducted pilot field trials of a prototype tag on five juvenile northern elephant seals, Mirounga angustirostris, using experimental exposures to both natural and anthropogenic noise stimuli. Results indicate behavioral responses were elicited in tagged individuals. However, no pattern was found in the occurrence and types of response compared to stimulus type. Responses during the ascending dive phase consisted of a dive inversion, or sustained reversal from ascending to descending (8 of 9 exposures). Dive inversions following exposure were 4–11 times larger than non-exposure inversions. Exposures received during the descending dive phase resulted in increased descent rates in 9 of 10 exposures. All 8 exposures during dives in which maximum dive depth was limited by bathymetry were characterized by increased flow noise in the audio recordings following exposure, indicating increased swim speed. Results of this study demonstrate the ability of an animal-mounted sound source to elicit behavioral responses in free-ranging individuals. Behavioral responses varied by seal, dive state at time of exposure, and bathymetry, but followed an overall trend of diving deeper and steeper and swimming faster. Responses did not consistently differ based on stimulus type, which may be attributable to the unique exposure context of the very close proximity of the sound source. Further technological development and focused field efforts are needed to advance and apply these tools and methods in subsequent behavioral response studies to address specific questions.

Long-Term Cognitive Deficits After Subarachnoid Hemorrhage in Rats

Cognitive dysfunction can be a long-term complication following subarachnoid hemorrhage (SAH). Preclinical models have been variously characterized to emulate this disorder. This study was designed to directly compare long-term cognitive deficits in the context of similar levels of insult severity in the cisterna magna double-blood (DB) injection versus prechiasmatic blood (PB) injection SAH models. Pilot work identified blood injectate volumes necessary to provide similar mortality rates (20-25%). Rats were then randomly assigned to DB or PB insults. Saline injection and naïve rats were used as controls. Functional and cognitive outcome was assessed over 35days. DB and PB caused similar transient rotarod deficits. PB rats exhibited decreased anxiety behavior on the elevated plus maze, while anxiety was increased in DB. DB and PB caused differential deficits in the novel object recognition and novel object location tasks. Morris water maze performance was similarly altered in both models (decreased escape latency and increased swimming speed). SAH caused histologic damage in the medial prefrontal cortex, perirhinal cortex, and hippocampal CA1, although severity of injury in the respective regions differed between DB and PB. Both SAH models caused long-term cognitive deficits in the context of similar insult severity. Cognitive deficits differed between the two models, as did distribution of histologic injury. Each model offers unique properties and both models may be useful for study of SAH-induced cognitive deficits.

Swimming in a two-dimensional Brinkman fluid: Computational modeling and regularized solutions

The incompressible Brinkman equation represents the homogenized fluid flow past obstacles that comprise a small volume fraction. In nondimensional form, the Brinkman equation can be characterized by a single parameter that represents the friction or resistance due to the obstacles. In this work, we derive an exact fundamental solution for 2D Brinkman flow driven by a regularized point force and describe the numerical method to use it in practice. To test our solution and method, we compare numerical results with an analytic solution of a stationary cylinder in a uniform Brinkman flow. Our method is also compared to asymptotic theory; for an infinite-length, undulating sheet of small amplitude, we recover an increasing swimming speed as the resistance is increased. With this computational framework, we study a model swimmer of finite length and observe an enhancement in propulsion and efficiency for small to moderate resistance. Finally, we study the interaction of two swimmers where attraction does not occur when the initial separation distance is larger than the screening length.

Partitioning the metabolic scope: the importance of anaerobic metabolism and implications for the oxygen- and capacity-limited thermal tolerance (OCLTT) hypothesis.

Ongoing climate change is predicted to affect the distribution and abundance of aquatic ectotherms owing to increasing constraints on organismal physiology, in particular involving the metabolic scope (MS) available for performance and fitness. The oxygen- and capacity-limited thermal tolerance (OCLTT) hypothesis prescribes MS as an overarching benchmark for fitness-related performance and assumes that any anaerobic contribution within the MS is insignificant. The MS is typically derived from respirometry by subtracting standard metabolic rate from the maximal metabolic rate; however, the methodology rarely accounts for anaerobic metabolism within the MS. Using gilthead sea bream (Sparus aurata) and Trinidadian guppy (Poecilia reticulata), this study tested for trade-offs (i) between aerobic and anaerobic components of locomotor performance; and (ii) between the corresponding components of the MS. Data collection involved measuring oxygen consumption rate at increasing swimming speeds, using the gait transition from steady to unsteady (burst-assisted) swimming to detect the onset of anaerobic metabolism. Results provided evidence of the locomotor performance trade-off, but only in S. aurata. In contrast, both species revealed significant negative correlations between aerobic and anaerobic components of the MS, indicating a trade-off where both components of the MS cannot be optimized simultaneously. Importantly, the fraction of the MS influenced by anaerobic metabolism was on average 24.3 and 26.1% in S. aurata and P. reticulata, respectively. These data highlight the importance of taking anaerobic metabolism into account when assessing effects of environmental variation on the MS, because the fraction where anaerobic metabolism occurs is a poor indicator of sustainable aerobic performance. Our results suggest that without accounting for anaerobic metabolism within the MS, studies involving the OCLTT hypothesis could overestimate the metabolic scope available for sustainable activities and the ability of individuals and species to cope with climate change.

Taylor line swimming in microchannels and cubic lattices of obstacles.

Microorganisms naturally move in microstructured fluids. Using the simulation method of multi-particle collision dynamics, we study in two dimensions an undulatory Taylor line swimming in a microchannel and in a cubic lattice of obstacles, which represent simple forms of a microstructured environment. In the microchannel the Taylor line swims at an acute angle along a channel wall with a clearly enhanced swimming speed due to hydrodynamic interactions with the bounding wall. While in a dilute obstacle lattice swimming speed is also enhanced, a dense obstacle lattice gives rise to geometric swimming. This new type of swimming is characterized by a drastically increased swimming speed. Since the Taylor line has to fit into the free space of the obstacle lattice, the swimming speed is close to the phase velocity of the bending wave traveling along the Taylor line. While adjusting its swimming motion within the lattice, the Taylor line chooses a specific swimming direction, which we classify by a lattice vector. When plotting the swimming velocity versus the magnitude of the lattice vector, all our data collapse on a single master curve. Finally, we also report more complex trajectories within the obstacle lattice.

Enhancement of Swimming Speed Leads to a More-Efficient Chemotactic Response to Repellent.

Negative chemotaxis refers to the motion of microorganisms away from regions with high concentrations of chemorepellents. In this study, we set controlled gradients of NiCl2, a chemorepellent, in microchannels to quantify the motion of Escherichia coli over a broad range of concentrations. The experimental technique measured the motion of the bacteria in space and time and further related the motion to the local concentration profile of the repellent. Results show that the swimming speed of bacteria increases with an increasing concentration of repellent, which in turn enhances the drift velocity. The contribution of the increased swimming speed to the total drift velocity was in the range of 20 to 40%, with the remaining contribution coming from the modulation of the tumble frequency. A simple model that incorporates receptor dynamics, including adaptation, intracellular signaling, and swimming speed variation, was able to qualitatively capture the observed trend in drift velocity.

Heart rate and muscle performance during forced swimming of jack mackerel Trachurus japonicus at different temperatures

The physiological condition of jack mackerel [18.3 ± 1.2 cm fork length (FL), n = 24] was simultaneously monitored using electrocardiography (ECG) and electromyography (EMG) techniques during swimming exercise in a flume tank at different temperatures of 10, 15 and 22 °C. To monitor the effect of temperature on the threshold level of the maximum sustainable speed, EMG demonstrated that the initiation of ordinary muscle activity at the lowest temperature (10 °C) occurred at swimming speeds of 3.2–3.5 FL/s, which was slower than the activation speeds at 15 and 22 °C of 4.0–5.4 FL/s. Ordinary muscle became active with increasing swimming speed which subsequently resulted in the frequency, amplitude and duration of burst discharge. ECG and EMG monitoring showed that the heart rate started to increase at the sustained speed level before the threshold and markedly increased up to the maximum level of heart rate at the higher speed. The index of ordinary muscle power output at the lowest temperature of 10 °C was lower than at higher temperatures of 15 and 22 °C. The heart rate increased with the increasing power output of ordinary muscle for all temperatures.

How body torque and Strouhal number change with swimming speed and developmental stage in larval zebrafish.

Small undulatory swimmers such as larval zebrafish experience both inertial and viscous forces, the relative importance of which is indicated by the Reynolds number (Re). Re is proportional to swimming speed (vswim) and body length; faster swimming reduces the relative effect of viscous forces. Compared with adults, larval fish experience relatively high (mainly viscous) drag during cyclic swimming. To enhance thrust to an equally high level, they must employ a high product of tail-beat frequency and (peak-to-peak) amplitude fAtail, resulting in a relatively high fAtail/vswim ratio (Strouhal number, St), and implying relatively high lateral momentum shedding and low propulsive efficiency. Using kinematic and inverse-dynamics analyses, we studied cyclic swimming of larval zebrafish aged 2-5 days post-fertilization (dpf). Larvae at 4-5 dpf reach higher f (95 Hz) and Atail (2.4 mm) than at 2 dpf (80 Hz, 1.8 mm), increasing swimming speed and Re, indicating increasing muscle powers. As Re increases (60 → 1400), St (2.5 → 0.72) decreases nonlinearly towards values of large swimmers (0.2-0.6), indicating increased propulsive efficiency with vswim and age. Swimming at high St is associated with high-amplitude body torques and rotations. Low propulsive efficiencies and large yawing amplitudes are unavoidable physical constraints for small undulatory swimmers.