Swimming Velocity Research Articles

The concentration of dissolved organic matter impacts the neurobehavior in zebrafish larvae exposed to cyclophosphamide.

Dissolved organic matter (DOM) occurs ubiquitously in various water matrices and affects the chemical speciation and toxicity of emerging contaminants, such as cyclophosphamide (CP). However, the effects of CP in aquatic organisms with the presence of DOM have been relatively less addressed. In this study, zebrafish eggs < 4h post fertilization (hpf) were exposed to CP (0 and 50μg/L) and humic acid (HA, a main component of DOM, 0, 3, 10, and 30mg-C/L) until 7days post fertilization, and its toxicity was evaluated by behavioral approaches and transcription of nervous-related genes. An increase in swimming velocity and anxiety was noticed in zebrafish larvae exposed to CP. The related genes of neurotransmitter (drd1, mao, thp1b, and gad2), neurodevelopment (gli2b, nrd, and gfap), and neuroinflammation (thfα, casp3, and il-6) were upregulated by CP. In the presence of HA (3mg-C/L), the behaviors and gene transcripts of zebrafish larvae were enhanced, while at 10mg-C/L, they were mitigated. This study has demonstrated that DOM at low concentration increases the toxicity of CP and at high concentration alleviates its toxicity. This study highlights the importance of emerging contaminant exposure with the presence of DOM on their toxicities in aquatic organisms.

Study on the effect of geometric shape on microswimmer upstream motion

The upstream motility of three microswimmer shapes (circular squirmer, squirmer rod, and elliptical squirmer) at the center of a Poiseuille flow is numerically investigated using the lattice Boltzmann method. Based on the stability and upstream ability, the swimming velocities and four motion states (stable motion, progressively unstable motion, unstable motion, and upstream failure) are summarized. The results show that the circular squirmer and squirmer rod are more stable than the elliptical squirmer; however, the elliptical squirmer has the greatest advantage in velocity and can swim up to twice as fast as the circular squirmer under the same conditions. The swimming type is also the key to influencing the motion state, which is reflected differently in the distinct microswimmer shapes. The increase in the Reynolds number (Re) and self-propelled strength (α) aggravates the motion instability; however, for elongated microswimmers, the aspect ratio (ε) plays a role in velocity rather than the motion state. Moreover, the upstream velocity of the pusher is always better than that of the puller, especially when Re increases. Notably, all microswimmers can maintain stable swimming when the preset velocity is twice the maximum velocity of the flow field. These findings can provide guidelines for the selection of design parameters and the appearance of microswimmers that resist complex incoming flows.

Alleviation of bovine serum albumin on the neurotoxicity of silver nanoparticles in zebrafish (Danio rerio) larvae

Silver nanoparticles (AgNPs) are ubiquitous in the aquatic environment and have attracted extensive attention to their toxic effects on aquatic species. However, responses of the nervous system to AgNPs are little known, especially co-existing with the ubiquitous natural organic matter (NOM), which is critical for the ability to act in aquatic species. Here, this study investigated the neurotoxicology of environmentally relevant AgNPs with or without bovine serum albumin (BSA; a classical NOM) to zebrafish (Danio rerio) using visualized transgenic zebrafish. Exposure to AgNPs reduced the locomotor behavior of zebrafish by 28%–45%, including swimming distance and velocity, exhibiting obvious behavioral inhibition. The visualized transgenic zebrafish treated with AgNPs showed developmental retardation in the early development of the heart, central nervous, and motor nerve, as well as the related neurodevelopment genes, which may be responsible for the lowered locomotor behavior. In addition, AgNPs can specifically interfere with the cholinergic system and affect neuronal signaling, ultimately leading to behavioral abnormalities. However, the co-existing BSA alleviated the neurotoxicity of AgNPs in zebrafish, which may partially be attributed to the increased size and electronegativity of AgNPs caused by BSA, thus reducing the direct interaction of AgNPs with cells. The interaction between BSA and the released Ag+ from AgNPs may also be responsible for the alleviation of the neurodevelopment dysfunction in zebrafish. These findings provide valuable insights into the toxicity and risks of AgNPs in natural aquatic environments.

Puzzle feeder as a cognitive enrichment strategy determines long-term welfare improvements in a teleost fish

Providing access to cognitive tasks, such as puzzle feeders, holds promise as enrichment strategy to enhance the welfare of captive animals. Despite the substantial number of teleost fish in captivity, the effects of cognitive enrichments in this group remain relatively unexplored. To address this gap in knowledge, we investigated the response of zebrafish to a puzzle feeder. In the first experiment of this study, zebrafish were presented with the option of a puzzle feeder or freely available food. While the subjects exhibited a preference for the latter option, they demonstrated some interest in the puzzle feeder, continuing to use it even though there was easily accessible food nearby. In a subsequent experiment, zebrafish groups were maintained in either a cognitive enrichment condition, where food was consistently provided through puzzle feeders, or a control condition with no cognitive enrichments. Welfare effects of the treatments were then assessed using standard behavioural indicators in the open-field test (i.e., time spent in the centre of the apparatus; number of entrances in the central sector; swim velocity; erratic swimming). Results unveiled an improvement in three welfare indicators due to cognitive enrichment, becoming apparent approximately 20 days after the treatment. Our study underscores the significance of puzzle feeders as cognitive enrichments in captive fish welfare, although consideration of other welfare indicators and different types of puzzles that elicit a more pronounced attraction is required.

The Load-Velocity Slope Is an Indicator of the Active Drag in all Competitive Swimming Strokes.

Active drag in swimming is a critical variable that affects swimmers' performance as well as the physiological load, but it is challenging for practitioners to assess this variable. This study aimed to assess if the load-velocity profiling method can be used as an indicator of active drag. A total of 419 swimmers performed three semi-tethered swimming trials in their speciality among the four competitive strokes with different external loads. Linear regression between external load and swimming velocity, as well as the external load relative to the body mass and swimming velocity, were established. The active drag and drag coefficient of each swimmer were calculated using a velocity perturbation method. There were significant correlations of the active drag with the absolute slope (r ≥ 0.713, p < 0.001) and relative slope (r ≥ 0.538, p < 0.001) in all four strokes and both sexes. A multiple regression analysis exhibited that the primary determinant of these relationships was the drag coefficient (semi-partial correlation ≥0.404, p < 0.001). The effects of the height and body mass index (BMI) on the relationship between the drag and the absolute slope were small (0.195 ≤ semi-partial correlation ≤0.248, p < 0.001) in both cases, which became either non-significant (height: p ≥ 0.282) or trivial (BMI: -0.099 ≤ semi-partial correlation ≤ -0.073, p ≤ 0.009). These results indicate that the absolute load-velocity slope is a strong indicator of the active drag, and the relative slope is useful when indirectly assessing the drag coefficient.

Efficient navigation of cargo-towing microswimmer in non-uniform flow fields

The vision of deploying miniature vehicles within the human body for intricate tasks holds tremendous promise across engineering and medical domains. Herein, optimal navigation of a cargo-towing swimmer under an applied zig-zag flow is studied by employing direct numerical simulations coupled with a deep reinforcement learning algorithm. Tasks include navigation in flow and shear-gradient directions. We initially explore combinations of state inputs, finding that optimal navigation necessitates swimmers to perceive hydrodynamics and alignment, surpassing reliance solely on hydrodynamic signals while considering their memories. Next, we study combinations of action spaces, allowing dynamic changes in swimming and/or rotational velocities by tuning B1 and C1 parameters of the squirmer model, respectively. By keeping both parameters fixed, cargo-towing swimmers demonstrate superior performance in the flow direction compared to swimmers without load due to tumbling movements influenced by shear flow. In the shear-gradient direction, swimmers without load outperform cargo-towing swimmers, with performance decreasing as load length increases. Across the combination of allowing B1 and C1 to change, the policies from solely dynamic B1 actions demonstrate superior navigation. The policies are then used as a showcase against naive cargo-towing and inert colloidal chains. A t-distributed stochastic neighbor embedding analysis reveals the complex interplay between perceived hydrodynamic signals and swimmer position. In the flow direction, swimmers align effectively with regions of maximum velocity, while in the shear-gradient direction, periodic transitions from minimum to maximum state values occur. Comparing pullers, pushers, and neutral swimmers, cargo-towing swimmers show a reversal in swimming velocity trends, with pullers outpacing neutral and pusher swimmers, irrespective of load lengths. Published by the American Physical Society 2024

Modeling and predicting heat transfer performance in bioconvection flow around a circular cylinder using an artificial neural network approach

The current research aims to study the heat transfer rate in mixed convection flow of viscous fluid along a vertical cylinder containing swimming microorganisms and velocity slip effects. The thermal and solutal processes are examined using gyrotactic microorganisms, chemical reaction and slip conditions. The non-linear (PDEs) of the governing flow are transformed into set of highly nonlinear (ODEs) by employing appropriate similarity transformations, which are then resolved computationally by Runge-Kutta-Fehlberg 4th order toward with shooting approach. The predicted solution is derived using the Levenberg-Marquardt scheme combined with a backpropagation neural network (LMS-BPNN). The labelled data sheet is divided into three parts: 80% data is used for training, 10% for validation and 10% for testing. The performance of the proposed AI-based LMS-BPNN is examined in term of MSE for considered scenarios. An optimal solution is achieved at 675, 397, 727, 647, and 711, epochs, while performance in terms of MSE for these epochs is to be 1.06×E−9, 8.47×E−10, 9.20×10−9, 9.94×E−10,and 9.09×10−10. The assigned computational valuations and ANN valuations are in excellent alignment than those existing studies on numerous special cases. The predicted solution with AI-based technique LMS-BPNN is reliable, well-organized, and easy to handle the thermal and solutal transport analysis of flow across a stretchable cylinder. The validity of proposed LMS-BPNN algorithm is examined with absolute error analysis and error is found to be approximately 10−4.

HCN channels are essential for the escape response of Paramecium.

When mechanical stimulation was applied to free swimming Paramecium, forward swimming velocity transiently increased due to activation of the posterior mechanosensory channels. The behavior response, known as "escape response," requires membrane hyperpolarization and the activation of K-channel type adenylate cyclases. Our hypothesis is that this escape response also involves activation of hyperpolarization-activated cyclic nucleotide-gated (HCN) channels. HCN channels are activated by hyperpolarization and are modulated by cyclic nucleotides such as cAMP and cGMP. They play a critical role in many excitable cells in higher animals. If HCN channels act in Paramecium, this should help to enhance and prolong hyperpolarization, thereby increasing the swimming speed of Paramecium. This study used RNAi to examine the role of the HCN channel 1 in the escape responses by generating hcn1-gene knockdown cells (hcn1-KD). These cells showed reduced mechanically-stimulated escape responses and a lack of cGMP-dependent increases in swimming speed. Electrophysiological experiments demonstrated reduced hyperpolarization upon injection of large negative currents in hcn1-KD cells. This is consistent with a decrease in HCN1 channel activity and changes in the escape response. These findings suggest that HCN1 channels are K+ channels that regulate the escape response of Paramecium by amplifying the hyperpolarizations elicited by posterior mechanical stimulation.

The turbulent soundscape of intertidal oyster reefs.

Turbulence and sound are important cues for oyster reef larval recruitment. Numerous studies have found a relationship between turbulence intensity and swimming behaviors of marine larvae, while others have documented the importance of sounds in enhancing larval recruitment to oyster reefs. However, the relationship between turbulence and the reef soundscape is not well understood. In this study we made side-by-side acoustic Doppler velocimeter turbulence measurements and hydrophone soundscape recordings over 2 intertidal oyster reefs (1 natural and 1 restored) and 1 adjacent bare mudflat as a reference. Sound pressure levels (SPL) were similar across all three sites, although SPL > 2000 Hz was highest at the restored reef, likely due to its larger area that contained a greater number of sound-producing organisms. Flow noise (FN), defined as the mean of pressure fluctuations recorded by the hydrophone at f < 100 Hz, was significantly related to mean flow speed, turbulent kinetic energy, and turbulence dissipation rate (ε), agreeing with theoretical calculations for turbulence. Our results also show a similar relationship between ε and FN to what has been previously reported for ε vs. downward larval swimming velocity (w b ), with both FN and w b demonstrating rapid growth at ε > 0.1 cm2 s-3. These results suggest that reef turbulence and sounds may attract oyster larvae in complementary and synergistic ways.

Exposure to thimerosal induces behavioral abnormality in the early life stages of zebrafish via altering amino acid homeostasis

Thimerosal (THI) has become a significant source of organic mercury pollutants in aquatic ecosystems, but there is limited information regarding its adverse effects on fish. In this study, zebrafish embryos were exposed to THI at 0 (control), 5.0, and 50 ng/L from 0–5 days post fertilization (dpf), and variations in their survival, development, behavior, free amino acid contents, and the biochemical responses involved in monoaminergic systems were examined. Although THI exposure did not significantly affect the survival, heart rate, or hatching time of zebrafish embryos, it substantially increased swimming velocity (136–154 % of the control) and reduced exploratory behavior (141–142 % of the control) in zebrafish larvae at 5 dpf. Exposure also significantly altered the amino acid contents (51–209 % of the control) and monoamine levels (70–154 % of the control) in zebrafish larvae, some of which displayed significant correlations with behavioral traits. THI significantly elevated dopamine receptor gene expression and monoamine oxidase activity in zebrafish larvae. Adding extra phenylalanine or tryptophan to the E3 medium facilitates the recovery of zebrafish larvae from the abnormal behaviors induced by THI. These findings reveal for the first time that THI exposure at the level of ng/L is sufficient to induce neurobehavioral toxic effects in the early life stages of zebrafish, and disrupting amino acid homeostasis is a critical underlying mechanism. This study provides valuable insights into the toxicity of THI to fish and highlights the importance of assessing its potential risks to aquatic ecosystems.

Agreement between an automated video-based system and tethered system to measure instantaneous swimming velocity

ABSTRACT Successful performance in competitive swimming requires a swimmer to maximise propulsion and minimise drag, which can be assessed using instantaneous swimming velocity. Many systems exist to quantify velocity, and therefore, it is important to understand the agreement between systems. This study examined the agreement between an automated video-based system and a tethered system to measure instantaneous velocity. Twenty-two competitive swimmers (state level or higher) completed 25 m of each stroke at maximal intensity. The tethered speedometer was attached to the swimmer’s waist, while videos of each trial were recorded. The swimmer’s head was then automatically tracked using proprietary software, and instantaneous velocity was determined from each system. Bland-Altman plots showed good agreement between the two systems in backstroke (95% Limits of Agreement (LOA): −0.24–0.26 m.s−1) and freestyle (95% LOA: −0.36–0.38 m.s−1) but poorer agreement in butterfly (95% LOA: −0.51–0.53 m.s−1) and breaststroke (95% LOA: −0.88–0.92 m.s−1). The root mean square error was higher in butterfly (0.27 m.s−1) and breaststroke (0.46 m.s−1) compared to backstroke (0.13 m.s−1) and freestyle (0.19 m.s−1). Results demonstrated that the two systems are comparable for measuring instantaneous swimming velocity; however, larger discrepancies are evident for butterfly and breaststroke.

Sex, drugs, and zebrafish: Acute exposure to anxiety-modulating compounds in a modified novel tank dive test

This study investigated the effects of anxiogenic and anxiolytic drugs on zebrafish (Danio rerio) behaviour using a modified novel tank dive test with higher walls and a narrower depth. Zebrafish were administered chondroitin sulfate, beta-carboline, delta-9-tetrahydrocannabinol (THC), ethanol, and beta-caryophyllene, and their behaviours were evaluated for geotaxis, swimming velocity, and immobility. Both anxiogenic and anxiolytic compounds generally increased bottom-dwelling behaviour, suggesting that the tank's modified dimensions significantly influence zebrafish responses. EC50 values for ethanol showed a lower threshold for velocity reduction compared to zone preference. Chondroitin sulfate uniquely caused a sex-specific increase in male swimming velocity, whereas no other sex-differences were observed with any compound. Interestingly, the presence of drug-treated fish did not alter the behaviour of observer fish, suggesting limited social buffering effects. The findings underscore the complexity of zebrafish behavioural phenotypes and highlight the need for considering tank dimensions and multiple behavioural parameters to accurately assess the effects of anxiety-modulating drugs. This study demonstrates the utility of the modified novel tank dive test in providing nuanced insights into the behavioural effects of different pharmacological agents in zebrafish.

Swimming Velocity Analysis Using Wearable Inertial Sensors and Speedometer: A Comparative Study.

The speedometer is widely used to evaluate swimming velocity but has some constraints. With the constant development of inertial units (IMUs), it is expected that they will become a good alternative to the speedometer. This study aimed to compare the data retrieved by an IMU and a speedometer when breaststroke is performed at maximum speed. Sixteen swimmers, nine males and seven females (20.3 ± 3.3 vs. 18.7 ± 1.1 years old, 65.8 ± 11.2 vs. 57.7 ± 9.1 kg of body mass and 1.75 ± 0.07 vs. 1.61 ± 0.10 m of height, respectively), performed 4 × 25 m of breaststroke sprint. They were equipped with an IMU fixed to the sacrum and with the line of an electromechanical speedometer (acquisition frequency of 50 Hz) fixed at the central point in the lumbar region. Statistical parametric mapping was used to compare the velocity curves, IBM SPSS was used for descriptive statistics and Bland-Altman plots were used for agreement of measurements. The results show that the IMU and speedometer do not show similar patterns, and the velocity values measured by the IMU are lower (p < 0.001). Bland-Altman plots presented a larger bias in terms of coefficient of variation and intracycle velocity variation. It can be concluded that IMUs and speedometers are not substitutes for each other as methods for evaluating intracycle velocity variations.

Identifying fish and estimating abundance and swim velocities of migrating Pacific salmon using adaptive resolution imaging sonar in mobile surveys

Abstract Mobile acoustic sounding is an effective survey method for fish abundance residing or migrating in large riverine basins and marine areas. A long-standing challenge in acoustic fish surveys with conventional sonar is the uncertainty in identifying fish targets from acquired echo data. Identification errors of fish targets can significantly bias estimates of fish abundances and negatively impact the management of fisheries. In contrast to conventional sonar, adaptive resolution imaging sonar (ARIS), if deployed properly with appropriate settings, can yield high-quality images of fish targets. ARIS images acquired with adequate frame rates can form video recordings to allow for confident identification of fish targets from recorded morphological features, sizes, direction of movements, and speeds. In this paper, we present an approach of using ARIS sonar for mobile surveys of fish passage in a riverine environment. Applications of this approach are demonstrated for its practical values with results from an ARIS-based mobile survey of upstream migrating salmon at an acoustically challenging fish-counting site on the lower Fraser River in British Columbia, Canada.

Non-native amphipods reach higher maximum swimming velocities than natives

The invasion of non-native amphipods often leads to severe changes in the composition of aquatic communities and may also result in the local replacement of native species. Particularly, a lower risk of being preyed upon resulting from high swimming velocities can be an advantage in interspecific competition. Furthermore, as swimming velocities are ecologically important, they are often used in ecotoxicological studies to estimate effects of different stressors. However, knowledge on swimming velocities of native and non-native amphipods is still rather limited. We experimentally investigated the maximum swimming velocities of three native and three non-native amphipods via video analyses in the laboratory. Results showed that non-native species reach higher maximum swimming velocities compared to natives probably leading to a higher predation success and reduced risk of being preyed upon. Additionally, body length correlates positively with swimming velocities, except for the invader Dikerogammarus villosus. As D. villosus can be cannibalistic, the high swimming velocities of the small individuals may reduce the intraspecific predation and may increase the survival rates of smaller specimen. Hence, knowledge about the swimming velocities of species contribute to the understanding of interspecific competition among species and might support explanation approaches for the success of invasive species. Furthermore, it provides baselines for ecotoxicological studies of stressor impacts.

Agouti-Induced Anxiety-Like Behavior Is Mediated by Central Serotonergic Pathways in Zebrafish.

Overexpression of the agouti-signaling protein (asip1), an endogenous melanocortin antagonist, under the control of a constitutive promoter in zebrafish [Tg(Xla.Eef1a1:Cau.Asip1]iim4] (asip1-Tg) increases food intake by reducing sensitivity of the central satiety systems and abolish circadian activity rhythms. The phenotype also shows increased linear growth and body weight, yet no enhanced aggressiveness in dyadic fights is observed. In fact, asip1-Tg animals choose to flee to safer areas rather than face a potential threat, thus suggesting a potential anxiety-like behavior (ALB). Standard behavioral tests, i.e., the open field test (OFT), the novel object test (NOT), and the novel tank dive test (NTDT), were used to investigate thigmotaxis and ALB in male and female zebrafish. Results showed that the asip1-Tg strain exhibited severe ALB in every test, mainly characterized by pronounced freezing behavior and increased linear and angular swimming velocities. asip1-Tg animals exhibited low central serotonin (5-HT) and dopamine (DA) levels and high turnover rates, thus suggesting that central monoaminergic pathways might mediate melanocortin antagonist-induced ALB. Accordingly, the treatment of asip1-Tg animals with fluoxetine, a selective serotonin reuptake inhibitor (SSRI), reversed the ALB phenotype in NTDT as well as 5-HT turnover. Genomic and anatomical data further supported neuronal interaction between melanocortinergic and serotonergic systems. These results suggest that inhibition of the melanocortin system by ubiquitous overexpression of endogenous antagonist has an anxiogenic effect mediated by serotonergic transmission.

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

Hydrodynamic behavior of inertial elongated microswimmers in a horizontal channel

In the current study, the lattice Boltzmann method was used to explore the motion of an elongated microswimmer in a horizontal channel with finite fluid inertia. By employing an extended squirmer rod model, the swimming velocity, hydrodynamic efficiency, and interaction with the channel wall of the capsule-shaped squirmer rod were simulated. It was found that the aspect ratio α and the swimming Reynolds number Res of the squirmer rod significantly affect its swimming velocity and efficiency. Specifically, as the Reynolds number increases, the pusher rod's velocity increases, whereas the puller rod's velocity decreases. Moreover, compared with the puller rod, the pusher rod has a higher efficiency with the same power consumption. With the increase of the aspect ratio α, the velocity of the squirmer rod increases gradually, the power consumption of the pusher rod and the puller rod decreases gradually, and the efficiency increases gradually, showing the characteristics of lower energy consumption and higher efficiency. During the interaction of the squirmer rod with the wall, four distinct motion modes were identified, namely, steady linear motion, motion away from the wall, damped swinging motion, and wall-attraction oscillation. The emergence of these motion modes and their transitions could be associated with the pressure distribution formed between the squirmer rod and the wall. The results provide another perspective and theoretical basis for the design of bioinspired microswimming devices and microrobots, especially in medical applications such as precision drug delivery systems.

Hydrodynamic performance analysis of swimming processes in self-propelled manta rays

To fill the research gap regarding the whole process (steady-state and nonsteady-state phases) of median and/or paired fin (MPF) mode swimming in underwater organisms, a two-degree-of-freedom self-propelled coupling method of motion and hydrodynamics based on user-defined functions of Fluent software was established, and numerical simulations were carried out for the startup, acceleration, and steady-state phases of manta rays. The interaction mechanism among the hydrodynamic characteristics, vortex evolution, and pressure distribution was investigated in the mentioned phases. We concluded that the negative pressure zone generated by the leading edge vortex and the shear layer contributes to thrust generation and changes in swimming velocity dominate the hydrodynamic characteristics by affecting the evolution of the shear layer and the leading edge vortex, with a 17.54% increase in forward average velocity in the fourth cycle compared to the third cycle and a consequent 9.5% increase in average thrust. In the end, the relationship between the formation of trailing edge vortex rings and changes in thrust was revealed. The vortex ring contributes to the increase in thrust, but the formation of the vortex ring comes at the cost of the loss of the leading edge vortex negative pressure zone, which greatly affects thrust, decreasing to 38.3% of its peak. The swimming mechanism revealed in this study provides a reference for the study of MPF-driven biodynamics and a new simulation strategy for the prediction of bionic navigator motions.

Tubulin glycylation controls ciliary motility through modulation of outer-arm dyneins.

Tubulins undergo several kinds of posttranslational modifications (PTMs) including glutamylation and glycylation. The contribution of these PTMs to the motilities of cilia and flagella is still unclear. Here, we investigated the role of tubulin glycylation by examining a novel Chlamydomonas mutant lacking TTLL3, an enzyme responsible for initiating glycylation. Immunostaining of cells and flagella revealed that glycylation is only restricted to the axonemal tubulin composing the outer-doublet but not the central-pair microtubules. Furthermore, the flagellar localization of TTLL3 was found to be dependent on intraflagellar transport. The mutant, ttll3(ex5), completely lacks glycylation and consequently exhibits slower swimming velocity compared with the wild-type strain. By combining the ttll3(ex5) mutation with multiple axonemal dynein-deficient mutants, we found that the lack of glycylation does not affect the motility of the outer-arm dynein lacking mutations. Sliding disintegration assay using isolated axonemes revealed that the lack of glycylation decreases microtubule sliding velocity in the normal axoneme but not in the axoneme lacking the outerarm dyneins. Based on our recent study that glycylation occurs exclusively on β-tubulin in Chlamydomonas, these findings suggest that tubulin glycylation controls flagellar motility through modulating outer-arm dyneins, presumably by neutralizing the negative charges of glutamate residues at the C-terminus region of β-tubulin.