Average Swimming Velocity Research Articles

Emergence of odd elasticity in a microswimmer using deep reinforcement learning

We use the Deep Q-Network with reinforcement learning to investigate the emergence of odd elasticity in an elastic microswimmer model. For an elastic microswimmer, it is challenging to obtain the optimized dynamics due to the intricate elastohydrodynamic interactions. However, our machine-trained model adopts a transition strategy (the waiting behavior) to optimize the locomotion. For the trained microswimmers, we evaluate the performance of the cycles by the product of the loop area (called ) and the loop frequency and show that the average swimming velocity is proportional to the performance. By calculating the force-displacement correlations, we obtain the effective odd elasticity of the microswimmer to characterize its nonreciprocal dynamics. This emergent odd elasticity is shown to be closely related to the loop frequency of the cyclic deformation. Our work demonstrates the utility of machine learning in achieving optimal dynamics for elastic microswimmers and introduces postanalysis methods to extract crucial physical quantities such as nonreciprocality and odd elasticity. Published by the American Physical Society 2024

Effects of different initial speeds on subsequent glide and underwater undulatory swimming

ABSTRACT This study aimed to investigate the effect of different initial speeds on the performance during underwater undulatory swimming (UUS). The study included 13 female swimmers. Each participant was asked to perform a 15-m maximum UUS, starting with four different push-off speeds. The experiment was recorded using three underwater cameras; subsequently, a two-dimensional motion analysis was conducted. Statistical parametric mapping (SPM) was employed to identify the position where the UUS velocity stabilised. The findings revealed a significant difference in the average swimming velocities during the first cycle of UUS, which was attributed to the variation in initial speed (p < 0.05) while there is no significant difference in the middle and final cycles. The results of SPM analysis suggested that differences in UUS velocity became negligible after approximately 6-m position from the pool wall, regardless of variations in push-off velocity. Furthermore, it was confirmed that swimmers can reach their maximum achievable UUS velocity at approximately 5-m position, even if they fail to execute an effective push-off from the wall. These findings offer valuable insights for future UUS studies, specifically in choosing suitable cycles for analysis.

Locomotory behaviour of euryhaline rotifer Brachionus rotundiformis: The potential influence of probiotics on swimming pattern and speed

AbstractWe evaluated the effect of probiotics on the growth and culture condition of rotifer in medium‐sized culture tanks. The influence of probiotics on the changes in swimming patterns, velocity and sinuosity indices of rotifer was also examined. The rotifers (Brachionus rotundiformis) were cultured for 14 days in nine polycarbonate tanks containing seawater which were divided into three groups: a control group (Chlorella product) and two treatment groups: Chlorella + Probiotic I and Chlorella + Probiotic II. The rotifer population and the water quality parameters were monitored daily. To evaluate the locomotory behaviour of the rotifer, a sub‐sample containing rotifers was obtained from each treatment and placed on a glass‐bottomed dish. A high‐speed camera mounted on a stereo microscope was used to record the rotifer movement. The footage was then analysed using motion analysis software to establish the swimming velocity, swimming pattern and sinuosity indices of the rotifers. Co‐feeding rotifer on microalgae and probiotics resulted in a higher rotifer growth rate (0.51 ± 0.03) and decreased concentration of unionized ammonia (0.14 ± 0.02 mg L−1). In addition, the probiotic‐fed rotifers had 1.59 times higher average swimming velocity (0.97 ± 0.11 mm s−1) and more straight swimming pattern compared to rotifers that fed on microalgae alone whose velocity was slow (0.61 ± 0.14 mm s−1) and more sinuous. From these data, we can postulate that probiotics have the potential to attenuate rotifer culture conditions by decreasing ammonia concentration as well as influence rotifer swimming behaviour which could affect rotifer reproduction and the quality of fish larvae in aquaculture systems.

Development of the technology to measure active drag of swimmers by the method of small perturbations

The research aimed at further improvement of the technology of measuring swimmers’ active drag by the small perturbation method (SPM) at the average maximal swimming velocity (v0max). For better reliability and accuracy of measurements, a standard variant of SPM was developed to measure automatically active drag (Fr(ad)), dimensionless coefficient of hydrodynamic force (Cx) and total external mechanical power Pto in different competitive swimming techniques during a single 50 m trial. The study involved twelve elite swimmers who specialized in front crawl at the distances of 100 and 200 m (780–850 FINA Points). From our measurements, in front crawl the results are following: v0max = 1.909 ± 0.017 m·s−1, Fr(ad) = 106.892 ± 7.276 N, Cx = 0.311 ± 0.028 and Pto = 204.033 ± 13.221 W. For objective estimation of the results, a verification variant of SPM was applied: v0max = 1.911 ± 0.018 m·s−1, Fr(ad) = 107.033 ± 7.232 N, Cx = 0.311 ± 0.030 and Pto = 204.467 ± 12.982 W. The correlation analysis of the results obtained by the standard and verification variants of SPM confirms high accuracy and reliability of the method used (r = 0.988 for v0max; r = 0.979 for Fr(ad); r = 0.988 for Cx(ad); r = 0.979 for Pto). The measurement error, including the method error and the devices error, does not exceed 2.4 %. This method has also a systematic error that reduces the measured data in the same direction from the actual values (min = -2.1 %; max = -2.9 %). This error should be taken into account when analyzing and evaluating the measured hydrodynamic characteristics of swimmers.

Swimming behavior and hydrodynamics of the Chinese cavefish Sinocyclocheilus rhinocerous and a possible role of its head horn structure.

The blind troglobite cavefish Sinocyclocheilus rhinocerous lives in oligotrophic, phreatic subterranean waters and possesses a unique cranial morphology including a pronounced supra-occipital horn. We used a combined approach of laboratory observations and Computational Fluid Dynamics modeling to characterize the swimming behavior and other hydrodynamic aspects, i.e., drag coefficients and lateral line sensing distance of S. rhinocerous. Motion capture and tracking based on an Artificial Neural Network, complemented by a Particle Image Velocimetry system to map out water velocity fields, were utilized to analyze the motion of a live specimen in a laboratory aquarium. Computational Fluid Dynamics simulations on flow fields and pressure fields, based on digital models of S. rhinocerous, were also performed. These simulations were compared to analogous simulations employing models of the sympatric, large-eyed troglophile cavefish S. angustiporus. Features of the cavefish swimming behavior deduced from the both live-specimen experiments and simulations included average swimming velocities and three dimensional trajectories, estimates for drag coefficients and potential lateral line sensing distances, and mapping of the flow field around the fish. As expected, typical S. rhinocerous swimming speeds were relatively slow. The lateral line sensing distance was approximately 0.25 body lengths, which may explain the observation that specimen introduced to a new environment tend to swim parallel and near to the walls. Three-dimensional simulations demonstrate that just upstream from the region under the supra-occipital horn the equipotential of the water pressure and velocity fields are nearly vertical. Results support the hypothesis that the conspicuous cranial horn of S. rhinocerous may lead to greater stimulus of the lateral line compared to fish that do not possess such morphology.

Zeolitic Imidazolate Framework-8 Nanoparticles Exhibit More Severe Toxicity to the Embryo/Larvae of Zebrafish (Danio rerio) When Co-Exposed with Cetylpyridinium Chloride.

The combined application of nanoparticles and surfactants has attracted tremendous attention in basic research and industry. However, knowledge of their combined toxicity remains scarce. In this study, we exposed zebrafish embryos to cetylpyridinium chloride (CPC, a cationic surfactant, at 0 and 20 μg/L), zeolitic imidazolate framework nanoparticles (ZIF-NPs, at 0, 30, and 60 mg/L), and their mixtures until 120 h post-fertilization (hpf). Within the used concentration range, both single and combined exposures exhibited limited effects on the survival and hatching of zebrafish. However, the combined exposure of ZIF-NPs and CPC caused more severe effects on the heart rate at both 48 and 72 hpf. The combined exposure also induced significant hyperactivity (i.e., increasing the average swimming velocity) and oxidative stress in zebrafish larvae (at 120 hpf), although all single exposure treatments exhibited limited impacts. Furthermore, the level of reactive oxygen species (or malondialdehyde) exhibited a significantly positive correlation with the heart rate (or average swimming velocity) of zebrafish, suggesting that oxidative stress plays a role in mediating the combined toxicity of CPC and ZIF-NPs to zebrafish. Our findings suggest that the interaction of CPC and ZIF-NPs should not be ignored when assessing the potential risks of their mixtures.

Clozapine modulation of zebrafish swimming behavior and gene expression as a case study to investigate effects of atypical drugs on aquatic organisms

Mental illnesses affect more than 150 million people in Europe and lead to an increasing consumption of neuroactive drugs during the last twenty years. The antipsychotic compound, clozapine, is one of the most used psychotropic drugs worldwide, with potentially negative consequences for the aquatic environment. Hence, the objectives of the study presented here were the quantification of clozapine induced changes in swimming behavior of exposed Danio rerio embryos and the elucidation of the molecular effects on the serotonergic and dopaminergic systems. Yolk-sac larvae were exposed to different concentrations (0.2 mg/L, 0.4 mg/L, 0.8 mg/L, 1.6 mg/L, 3.2 mg/L and 6.4 mg/L) of clozapine for 116 h post-fertilization, and changes in the swimming behavior of the larvae were assessed. Further, quantitative real-time PCR was performed to analyze the expression of selected genes. The qualitative evaluation of changes in the swimming behavior of D. rerio larvae revealed a significant decrease of the average swimming distance and velocity in the light-dark transition test, with more than a 36% reduction at the highest exposure concentration of 6.4 mg/L. Furthermore, the total larval body length was reduced at the highest concentration. An in-depth analysis based on expression of selected target genes of the serotonin (slc6a4a) and dopamine (drd2a) system showed an upregulation at a concentration of 1.6 mg/L and above. In addition, a lower increase in expression was detected for biomarkers of general stress (adra1a and cyp1a2). Our data show that exposure to clozapine during development inhibits swimming activity of zebrafish larvae, which could, in part, be due to disruption of the serotonin- and dopamine system.

Effects of Toluene on the Development of the Inner Ear and Lateral Line Sensory System of Zebrafish

ObjectiveThe aim of this study was to explore the ototoxicity of toluene in the early development of zebrafish embryos/larvae. MethodsZebrafish were utilized to explore the ototoxicity of toluene. Locomotion analysis, immunofluorescence, and qPCR were used to understand the phenotypes and molecular mechanisms of toluene ototoxicity. ResultsThe results demonstrated that at 2 mmol/L, toluene induced zebrafish larvae death at 120 hours post fertilization (hpf) at a rate of 25.79% and inhibited the rate of hatching at 72 hpf. Furthermore, toluene exposure inhibited the distance travelled and average swimming velocity of zebrafish larvae while increasing the frequency of movements. As shown by fluorescence staining of hair cells, toluene inhibited the formation of lateral line neuromasts and middle line 1 (Ml1) neuromasts in 3 days post fertilization larvae in a concentration-dependent manner. Toluene altered the expression level of genes involved in ear development/function in zebrafish, among which the mRNA levels of cd164l2, tekt3, and pcsk5a were upregulated, while the level of otofb was downregulated, according to the qPCR results. ConclusionThis study indicated that toluene may affect the development of both the inner ear and lateral line systems in zebrafish, while the lateral line system may be more sensitive to toluene than the inner ear.

Effects of temperature on hatching and growth performance of embryos and yolk-sac larvae of a threatened estuarine fish: Longfin smelt (Spirinchus thaleichthys)

The abundance of longfin smelt (Spirinchus thaleichthys) in the San Francisco Estuary (SFE) has declined to less than 1% of historic numbers, contributing to their listing as “threatened” under the California Endangered Species Act. To prevent local extirpation, a captive culture program is being developed for the longfin smelt at the Fish Conservation Culture Laboratory at UC Davis. To begin systematically addressing current issues with larviculture methods, we tested the effects of temperature on longfin smelt embryos from 7 clutches with different parentages. We reared embryos and newly hatched non-feeding yolk-sac larvae in freshwater at 9, 12, or 15 °C until the age when these fish underwent mass mortality (>90% mortality) during the experiment (3, 4, or 5 days post-hatch (dph), depending on rearing temperature). We measured hatch success, hatch morphometrics (notochord length, dry mass, yolk volume, and oil volume), length and dry mass growth rates, yolk and oil resorption rates, routine oxygen consumption rate, and locomotor activity (percent of time spent moving, total distance travelled, and average swimming velocity). Embryos and larvae reared at 15 °C performed poorly, experiencing reduced hatch success, hatch notochord lengths, growth rates, and earlier mass mortality. Embryos reared at 9 and 12 °C had similar hatch success but at hatch, larvae were longer at 9 compared to 12 °C. However, larvae grew faster in 12 compared to 9 °C resulting in both temperature groups having similar lengths prior to mass mortality (4 dph for 12 °C and 5 dph for 9 °C). Temperature had no effect on routine oxygen consumption rate among temperature groups but locomotor activity was elevated at 9 and 15 °C groups compared to 12 °C, suggesting that locomotor activity consumed a higher proportion of the energy budget, leaving a smaller fraction for growth. Finally, larval swimming speeds, one day prior to each temperature's respective day of mass mortality, were higher at 12 °C (3 dph) compared to 9 (4 dph) and 15 °C (2 dph), which could lead to higher foraging success. Interclutch variation was high, however, suggesting that parentage effects may result in brood-specific variation in the responses of larvae to water temperature. Taken together, these data suggest that rearing longfin smelt at 15 °C would have detrimental effects and we suggest that cooler temperatures such as 9 or 12 °C may improve the culturing of the earliest life stages of longfin smelt.

Comparison of performance of various leg-kicking techniques in fin swimming in terms of achieving the different goals of underwater activities.

The aim of this study was to compare underwater fin swimming performance using dolphin, flutter and breaststroke kicks with and without diving gear. Performance was evaluated in terms of average swimming velocity. The parameters of spatiotemporal structure of the stroke reflecting to the swimming economy were employed. Conscious modifications in propulsion technique were considered here with the aim of controlling swimming performance. A total of ten professional scuba divers swam at maximal speed underwater for 50m using each of three techniques: dolphin, flutter, or breaststroke kicks. Swimmers’ performance was compared between holding their breath and using breathing apparatus. Two cameras recorded their movements in sagittal and transverse planes. The average swimming velocity (vav), stroke length (SL), stroke rate (SR), index of variation of intracycle velocity (VIVIndex) and stroke index (SI) were estimated. Relative to the other techniques, the dolphin kick without a diving gear demonstrated the highest vav and low SI and VIVIndex values, which reflects the most advantageous economy of propulsion at given velocity. Given the lack of statistical differences, using the breaststroke kick and flutter kick when swimming with a diving gear seems to be comparable to dolphin-kick in terms of average velocity and parameters reflecting the economy of propulsion. Thus, a search for fin swimming techniques with the aim of achieving specific goals seemed reasonable. The results suggest, that performance achieved while using various fin swimming techniques was probably controlled by different strategies of leg movements. These strategies revealed differences in a spatiotemporal (SR-SL) structure of the stroke and they were closely associated in terms of the velocity variation decrease.

Psychophysiological respond in a swimming ultra-endurance event.

Ultra-endurance events under critical environmental conditions represent unique stress, resulting in acute marked adaptations to the cardiovascular, respiratory, metabolic and neuromuscular systems of the organism. In line with this, no studies in swimming events were found thus far. This research aimed to analyze the psychophysiological response of swimmers in an ultra-endurance swimming event. Changes in the rate of perceived exertion, blood lactate concentration, 1000m swimming time and cortical arousal were measured in 19 male volunteer swimmers (28.0±5.6 years; 175.0±7.9 cm; 70.1±7.8 kg) in 3 moments during a 24h swimming event. We found a significant increase in lactate concentration after series 1, decreasing the values in series 2 and series 3. Cortical arousal decreased significantly in series 3 respect the basal sample. Rate of perceived exertion significantly increased in the swimming ultra-endurance event and 1000m swimming time was maintained during the series analyzed. An ultra-endurance swimming event produced an increase in blood lactate concentration and rated of perceived exertion and a decrease in cortical arousal, not affecting average swimming velocity along the event.

Biologically inspired swimming robotic frog based on pneumatic soft actuators

Research on soft robots and swimming robots has been widely reported and demonstrated. However, none of these soft swimming robots can swim flexibly and efficiently using legs, just like a frog. This paper demonstrates a self-contained, untethered swimming robotic frog actuated by 12 pneumatic soft actuators, which can swim in the water for dozens of minutes by mimicking the paddling gait of the natural frog. We designed two kinds of pneumatic soft actuators as the joints on the robotic frog’s legs, which allows the legs to be lighter and more compact. It is found that such soft actuators have great potential in developing amphibious bionic robots, because they are fast-responding, inherently watertight and simple in structural design. The kinematic analysis in swimming locomotion was conducted for the prototype robotic frog, and the locomotion trajectory of each leg was planned based on the analysis of the paddling gait of frogs. Combined with the deformation model of the soft actuators, the robotic frog’s legs are controlled by coordinating the air pressure of each joint actuator. The robotic frog’s body is compact and the total mass is 1.29 kg. Different paddling gaits were tested to investigate swimming performance. The results show that the robotic frog has agile swimming ability and high environmental adaptability. The robotic frog can swim forward more than 0.6 m (3.4 times the body length) in one paddling gait cycle(6 s), whose average swimming velocity is about 0.1 m s−1, and the minimum turning radius is about 0.15 m (less than 1 body length).

Swimming performance of Arctic grayling (Thymallus arcticus Pallas) in an open-channel flume

Installing effective fish passage structures that provide connectivity for Arctic grayling is a promising conservation strategy for imperiled populations. The swimming abilities and behaviour of age 1+ adfluvial grayling from Montana were examined in an open-channel flume to provide design information for passage structures. Swimming behaviours and distance of ascent (Dmax) in a 12.25 m section was measured at four velocities (0.49, 1.04, 1.43, 2.26 m/s) at an average temperature of 11.3 °C (SD = 0.7); effects of fish length and time of day were also examined. Median Dmax was equal to 12.25 m for all treatments except the 2.26 m/s treatment, where it dropped to 4.5 m. Average fish swimming velocities increased from 0.85 to 2.97 m/s from the lowest to the highest velocity treatments and the maximum velocity observed was 4.03 m/s. Survival model parameterization allowed prediction of the proportion passing relative to distance for covariate values within the range tested. There was strong evidence Dmax was related to water velocity (p < 0.001) and an 82% decrease in Dmax is estimated for every 1 m/s increase in water velocity. There was no evidence for relationships among distance of ascent and fish length (p = 0.91) or time of day (p = 0.81).

A three-sphere microswimmer in a structured fluid

We discuss the locomotion of a three-sphere microswimmer in a viscoelastic structured fluid characterized by typical length and time scales. We derive a general expression to link the average swimming velocity to the sphere mobilities. In this relationship, a viscous contribution exists when the time-reversal symmetry is broken, whereas an elastic contribution is present when the structural symmetry of the microswimmer is broken. As an example of a structured fluid, we consider a polymer gel, which is described by a “two-fluid” model. We demonstrate in detail that the competition between the swimmer size and the polymer mesh size gives rise to the rich dynamics of a three-sphere microswimmer.

Three-disk microswimmer in a supported fluid membrane.

A model of three-disk micromachine swimming in a quasi-two-dimensional supported membrane is proposed. We calculate the average swimming velocity as a function of the disk size and the arm length. Due to the presence of the hydrodynamic screening length in the quasi-two-dimensional fluid, the geometric factor appearing in the average velocity exhibits three different asymptotic behaviors depending on the microswimmer size and the hydrodynamic screening length. This is in sharp contrast with a microswimmer in a three-dimensional bulk fluid that shows only a single scaling behavior. We also find that the maximum velocity is obtained when the disks are equal-sized, whereas it is minimized when the average arm lengths are identical. The intrinsic drag of the disks on the substrate does not alter the scaling behaviors of the geometric factor.

Habitat orientation alters the outcome of interspecific competition: A microcosm study with zooplankton grazers.

Habitat orientation has recently been demonstrated to affect the foraging behavior, growth, and production of plankton grazers. Because the orientation effect may vary with species, we hypothesize that habitat orientation may alter interspecific interactions between animal species. We experimentally investigated how habitat orientation (placing cuboid chambers in three orientations with long, medium, and small side as the chamber height) affected the interaction between two common cladoceran species, Daphnia magna and Moina micrura, which competitively exploited green algae of Chlorella pyrenoidosa at two volume scales (64 and 512 ml). Results show that chamber orientation and volume additively affected the behavior and species performance of the grazers. Specifically, both grazer species generally decreased their average swimming velocity, grazing rate (on algal cells), body size, and survival and reproduction rates with increasing chamber height for both chamber volumes and with decreasing chamber volume regardless of chamber orientation. Nevertheless, the decrease magnitude was greater for M. micrura with increasing chamber height but was greater for D. magna with decreasing chamber volume. Correspondingly, when cocultured, the density ratio of D. magna to M. micrura increased with increasing chamber height but decreased with decreasing chamber volume. At the end of the experiment, none of D. magna individuals survived in the small and short (large‐based) chambers, and few M. micrura individuals survived in large and tall (small‐based) chambers. These results indicate that both habitat orientation and size affect the outcome of interspecific competition between grazer species. We suggest that variation in habitat orientation may improve community coexistence and species diversity in nature.

Simulations of microscopic propulsion of soft elastic bodies.

Using simulations that realistically model both hydrodynamic and elastic behavior, we study the motion of a microscopic, driven elastic sphere immersed in water. We first confirm the "jittery" relaxation recently predicted theoretically for an externally driven elastic sphere. The sphere is then divided in two and each section is driven internally with the two sections 180° out of phase. With periodic and perfectly symmetric driving, the elastic sphere spontaneously breaks symmetry and can attain macroscopic average swimming velocities to the right or left, the direction depending only on the initial state. With asymmetric driving the elastic sphere swims in one direction and the maximum speed is obtained with a 1/3:2/3 split. At high drive frequencies close to elastic resonances of the sphere, the motion can be quite efficient. At low drive frequencies the propulsion speed becomes independent of the elastic constants of the sphere and less efficient, but still substantial. Inertia is found to be an important driver of the behavior despite the small size of the spheres. As we model the full three-dimensional elasticity and compressible hydrodynamics, our simulations give not just qualitative indications but quantitative predictions for the motion.

Pacing the phasing of leg and arm movements in breaststroke swimming to minimize intra-cyclic velocity fluctuations.

In swimming propelling efficiency is partly determined by intra-cyclic velocity fluctuations. The higher these fluctuations are at a given average swimming velocity, the less efficient is the propulsion. This study explored whether the leg-arm coordination (i.e. phase relation ϕ) within the breaststroke cycle can be influenced with acoustic pacing, and whether the so induced changes are accompanied by changes in intra-cyclic velocity fluctuations. Twenty-six participants were asked to couple their propulsive leg and arm movements to a double-tone metronome beat and to keep their average swimming velocity constant over trials. The metronome imposed five different phase relations ϕi (90, 135, 180, 225 and 270°) of leg-arm coordination. Swimmers adjusted their technique under the influence of the metronome, but failed to comply to the velocity requirement for ϕ = 90 and 135°. For imposed ϕ = 180, 225 and 270°, the intra-cyclic velocity fluctuations increased with increasing ϕ, while average swimming velocity did not differ. This suggests that acoustic pacing may be used to adjust ϕ and thereby performance of breaststroke swimming given the dependence of propelling efficiency on ϕ.

Elastic Three-Sphere Microswimmer in a Viscous Fluid

We discuss the dynamics of a generalized three-sphere microswimmer in which the spheres are connected by two elastic springs. The natural length of each spring is assumed to undergo a prescribed cyclic change. We analytically obtain the average swimming velocity as a function of the frequency of the cyclic change in the natural length. In the low-frequency region, the swimming velocity increases with the frequency and its expression reduces to that of the original three-sphere model by Najafi and Golestanian. In the high-frequency region, conversely, the average velocity decreases with increasing the frequency. Such a behavior originates from the intrinsic spring relaxation dynamics of an elastic swimmer moving in a viscous fluid.

Age and serotonin effects on locomotion in marine trematode cercariae

The effect of serotonin solutions on the locomotor activity of cercariae with different swimming patterns, Cryptocotyle lingua and Himasthla elongate, was investigated during their lifespan using motion track analysis and a frame-by-frame count of behavioral elements. Serotonin caused a noticeable alteration of several locomotion parameters in both species studied, while the activity per se decreased significantly with age. In intermittently swimming C. lingua cercariae, serotonin induced an increase in the frequency of active swimming phases (spurts) without significant changes in the duration and velocity of the spurts themselves. In continuously swimming H. elongata cercariae, serotonin increased the number of stops during the first hour after the emission from the host mollusk under a constant average swimming velocity. Immunocytochemical visualization of serotonin revealed a decline in its level with age (time after emission) and different dynamics of this process in neurons located in the tail and body of C. lingua cercariae. The data are interpreted in the light of the modulatory role of serotonin known for other animals.