The Mechanics of Swimming in Schools: How Fish Coordinate Collective Movement

Fish schooling is a coordinated swimming behavior in which individuals maintain close proximity, align their body orientation, and move as a cohesive group. This article explains the hydrodynamic benefits, social drivers, and sensory mechanisms that enable fish to coordinate collective movement, with practical guidance for observing and assessing schooling behavior in wild and captive settings.

Schooling behavior emerges from local interactions between individual fish instead of from centralized leadership or communication over long distances. Each fish responds to the position, speed, and direction of nearby neighbors using vision and the lateral line system, a sensory network that detects water movement and pressure gradients. The result is a self-organizing group that can change direction rapidly, evade predators, locate food, and reduce the energetic cost of swimming.

Understanding the mechanics of schooling matters for fisheries biologists, aquaculture operators, aquarium managers, and researchers who study fish behavior. The same principles that allow wild fish to coordinate movement also influence how captive fish use space, respond to flow conditions, and cope with stress. This article provides a framework for observing schooling behavior, measuring key variables, and identifying situations that require professional intervention.

Why Fish Form Schools

Fish form schools for several overlapping reasons that vary by species, life stage, and environmental context. The primary benefits fall into three categories: predator defense, feeding efficiency, and energetic savings.

Predator defense is often the most immediate driver of schooling. A group of fish presents a confusing visual target to predators, and the collective vigilance of many individuals increases the likelihood that a threat will be detected early. When a predator attacks, the rapid, coordinated escape maneuvers of a school can disrupt the predator's aim. Research on crucian carp exposed to predation stress found that groups split into subgroups more frequently under predator pressure, suggesting that fragmentation itself may serve an antipredator function 15.

Feeding efficiency improves because fish in schools locate food patches faster than solitary individuals. Once one fish finds food, others can follow using visual cues and lateral line detection of feeding-related water movements. Schooling also allows fish to exploit prey that would be difficult for a solitary individual to capture.

Energetic savings are among the most studied benefits of schooling. Fish swimming in formation can use the vortices shed by neighbors to reduce their own drag and stroke effort. A 2023 review of hydrodynamic studies on fish schooling concluded that schooling motions can be energetically beneficial because schools allow for channeling and vortex-based interactions, creating a less demanding stroke rate to sustain high swimming velocities and increased movement efficiency 4.

The magnitude of these energy savings can be substantial. Research on giant danios swimming in turbulent conditions found that schools reduced their total energy expenditure by 63 to 79 percent compared to solitary fish at high speeds and high turbulence levels 8. Solitary individuals spent approximately 22 percent more kinematic effort, measured as tail beat amplitude multiplied by frequency, to swim in turbulence at higher speeds than in laminar conditions. Schools swimming in turbulence reduced their three-dimensional group volume by 41 to 68 percent and did not alter their kinematic effort compared to laminar conditions.

Hydrodynamic Principles of Schooling

The hydrodynamics of fish schooling involve complex interactions between the wakes of individual fish and the flow field of the surrounding water. Understanding these interactions requires examining how fish generate thrust, how vortices form and propagate, and how neighboring fish position themselves to benefit from or avoid these flow structures.

Vortex Formation and Wake Interactions

Fish swimming generates vortices in their wake as their bodies undulate and their tails beat from side to side. These vortices carry momentum and energy away from the fish. A following fish can potentially extract energy from these vortices if it positions itself correctly relative to the wake of the leader.

The classic model of schooling hydrodynamics, proposed by Weihs, suggested that fish in a diamond formation could benefit from the vortices shed by fish ahead of them. However, more recent integrated analyses combining computational fluid dynamics and experimental data have prompted a reevaluation of this classic model 3. The flow patterns observed in particle image velocimetry experiments with fish schools reveal complex hydrodynamic interactions that do not always match the predictions of simplified models.

Directly measuring hydrodynamic performance in live fish schools poses significant challenges because the flow structures generated by collective moving organisms are three-dimensional, chaotic, and complex 3. Many previous studies have used computational, mechanical, or robotic models to represent live fish. The scarcity of combined approaches that include both computational and experimental studies of the same fish schools has limited understanding of the physical factors involved in fish collective behavior.

Dipole and Vortex Sheet Models

To model the hydrodynamic interactions between swimming fish, researchers have developed mathematical representations of fish as flow-disturbing bodies. Dipole-based models assimilate fish to pairs of vortices and are attractive because of their mathematical tractability. However, validation studies using computational fluid dynamics informed by experimental data have shown that dipole-based models capture key features of the fluid flow but cannot predict the elongated flow streamlines around the fish that are evident in more detailed simulations 5.

An alternative model replaces each vortex in the pair with a sheet along the fish length. Using a pair of vortex sheets that span approximately 80 percent of the fish body length with a separation distance of approximately 50 percent of the body width, this model successfully predicts the fluid flow around a swimming fish for a range of background flow speeds and channel widths 5. The vortex sheet model shows improved accuracy at the cost of mildly increased computational effort.

Turbulence Sheltering

One of the most significant recent findings in schooling hydrodynamics is the turbulence sheltering hypothesis. This hypothesis proposes that collective movements of fish schools in turbulent flow can reduce the total energetic cost of locomotion by shielding individuals from the perturbation of chaotic turbulent eddies 8.

The mechanism works because fish within a school create a more stable internal hydrodynamic environment. The bodies of neighboring fish block and dissipate turbulent eddies before they can disrupt an individual's swimming gait. This sheltering effect is particularly important at high swimming speeds and high turbulence levels, where solitary fish must expend considerable energy to maintain their trajectory.

The practical implication is that schooling provides a buffer against environmental flow variability. Fish in schools can maintain stable swimming kinematics in conditions that would force solitary fish to increase their tail beat frequency and amplitude.

Sensory Mechanisms for Coordination

Fish coordinate schooling through multiple sensory modalities, with vision and the lateral line system playing the primary roles. The relative importance of each sensory system varies by species, environmental conditions, and the specific behavioral context.

The Lateral Line System

The lateral line is a mechanosensory system unique to fish and some amphibians. It consists of sensory receptors called neuromasts distributed across the body surface and within canals in the head. These receptors detect water movement, pressure gradients, and low-frequency vibrations.

The cephalic lateral line system of schooling fish is often highly developed. A study of the pelagic schooling fish Normanichthys crockeri found a complex arrangement of pores and canals in the head, including five preopercular pores, five mandibular pores, three supraorbital pores, eight infraorbital pores, three postorbital pores, three supratemporal pores, and one coronary pore 18. The high number of pores distributed along the anterior, middle, and posterior zones of the orbit makes this a quite sensitive area. These pores connect with eight narrow, continuous canals that respond maximally to water flow velocity along the axis of the canal.

The lateral line allows fish to detect the water movements generated by neighboring fish swimming nearby. This information helps fish maintain appropriate distances and adjust their swimming to match the movements of the group. The lateral line also detects the vortices shed by other fish, which may inform positioning decisions within the school.

Vision

Vision provides fish with information about the position, orientation, and movement of neighbors over longer distances than the lateral line can detect. Fish use visual cues to maintain alignment with neighbors, avoid collisions, and respond to changes in group direction.

Research on giant danios has examined how schooling changes when fish are unable to see or unable to use their lateral lines. The study found that giant danios were able to school normally without their lateral lines but did not school in darkness 16. Surprisingly, giant danios in darkness had the same attraction and alignment properties as fish in light, potentially indicating that they do not feel as much risk in darkness.

These findings suggest that vision is necessary for schooling in giant danios, while the lateral line is not essential when vision is available. However, the relative importance of these sensory systems likely varies among species depending on their natural predators and environment 16.

Multimodal Sensory Integration

Recent perspectives emphasize that fish integrate information from multiple sensory modalities during schooling. The acoustic hypothesis proposes that fish mainly use acoustic signals, including pressure waves and other water movements produced by their own swimming, to achieve synchronization and maintain appropriate distance from neighbors 19.

Swimming together causes overlapping and complex acoustic signals that may give the school significant survival advantages. Schooling prey fish may confuse predator hearing and lateral line systems with these overlapping signals. The acoustic hypothesis also suggests that the regular and predictable sounds produced by swimming fish could serve as communication signals within the school, conveying spatial and temporal information that enables synchronized locomotion 19.

A three-dimensional simulation study integrating physiological characteristics, movement patterns, and multi-source perception introduced a fluid dynamics based lateral line perception field and dynamic visual area calculation to model fish schooling 17. This approach quantified the process of energy recovery from wake vortices by clusters during swimming, demonstrating that fish likely combine lateral line and visual information to position themselves advantageously within the school.

Social Rules and Individual Interactions

The coordinated movement of fish schools emerges from local interaction rules that each fish follows relative to its neighbors. These rules have been described abstractly as attraction to neighbors that are far away, repulsion from neighbors that are too close, and alignment with neighbors at the correct distance 16.

Attraction, Repulsion, and Alignment

Attraction keeps the school cohesive. When a fish finds itself too far from its neighbors, it moves toward them. Repulsion prevents collisions. When a neighbor comes too close, the fish moves away. Alignment ensures that fish swim in the same direction at similar speeds.

These rules operate over different spatial scales. Repulsion dominates at very short distances, alignment operates at intermediate distances, and attraction becomes important at longer distances. The specific distances at which these rules switch depend on the species, body size, and swimming speed.

Speed Matching

Speed matching is a critical component of collective motion. Fish must adjust their swimming speed to match that of their neighbors to maintain school cohesion. This process is not always straightforward, especially in groups where individuals have different intrinsic movement characteristics.

Research on mixed-species schools of rosy barbs and tiger barbs found that these species readily form mixed-species schools despite striking differences in their intrinsic movement patterns. Tiger barbs exhibit both slow and fast swimming, whereas rosy barbs display fast swimming only. In mixed-species groups, the slow swimming speed of tiger barbs disappears, and the collective motion is dominated by a single fast-swimming mode 11.

An individual-based model incorporating local interactions involving speed matching demonstrated that bimodal speed in conspecific schools of tiger barbs is an emergent property that is lost in mixed-species groups. Despite high cohesion, the two species showed spatial sorting within the mixed-species groups, which the model explained through differences in inter- and intra-specific interactions 11.

Topological vs. Metric Interactions

Fish do not interact with all neighbors equally. Research suggests that fish interact with a finite number of topologically defined neighbors instead of all fish within a fixed distance. A coupled framework for simulating fish schooling integrated social interactions through a self-propelled particle model in which fish interact with a finite number of topologically defined neighbors 9.

In simulations of a minimal two-fish group, the trailing fish achieved stable locomotion through a slight side-slip, an emergent behavior not explicitly encoded in the model input. The model was further extended to large schools, demonstrating that group efficiency increases with the Reynolds number because of more favorable hydrodynamic interactions 9.

At a Glance: Key Factors Influencing School Formation

The following table summarizes the primary factors that influence whether and how fish form schools. These factors should be considered when observing fish in the wild or managing fish in captivity.

Observing and Measuring Schooling Behavior

Systematic observation of schooling behavior requires attention to specific variables that can be measured consistently across observation sessions. The following steps provide a practical framework for assessing schooling behavior in wild or captive fish.

Step 1: Define the Observation Context

Record the environmental conditions at the time of observation, including water temperature, flow velocity, light levels, time of day, and the presence of potential predators or stressors. These variables can significantly influence schooling behavior and must be documented to interpret observations correctly.

Step 2: Measure Group Structure

Quantify the spatial structure of the school by measuring nearest neighbor distance and mean pairwise distance between individuals. These metrics describe how tightly packed the school is and how evenly fish are spaced. Research on Leptobotia elongata found that nearest neighbor distance showed no significant differences across flow conditions, indicating stable schooling cohesion, while mean pairwise distance increased under higher flow heterogeneity 14.

Step 3: Assess Group Polarization

Polarization describes the degree to which fish in a school are oriented in the same direction. A highly polarized school has all fish pointing the same way, while a low-polarization group has individuals oriented randomly. Polarization can be estimated visually or measured from video recordings using image analysis software.

Step 4: Record Swimming Speed and Kinematics

Measure the swimming speed of individual fish and the group as a whole. Note tail beat frequency and amplitude, as these kinematic variables indicate the energetic cost of swimming. Fish schools swimming in turbulence did not alter their kinematic effort compared to laminar conditions, while solitary fish increased their effort 8.

Step 5: Document Behavioral Events

Record specific behavioral events such as group splitting, rejoining, direction changes, and responses to stimuli. The splitting number of fish groups was higher in predator stress treatments compared to control treatments across three ecological contexts 15.

Step 6: Compare Across Conditions

To understand what factors influence schooling in your study system, compare observations across different conditions. For example, compare school structure in laminar versus turbulent flow, in light versus darkness, or in the presence versus absence of predators.

Records and Measurements for Schooling Studies

Maintaining consistent records is essential for tracking schooling behavior over time and detecting changes that may indicate welfare problems or environmental issues. The following measurements provide a useful baseline for any schooling study.

Common Failure Patterns in Schooling Observations

Several recurring problems can compromise the validity of schooling observations. Recognizing these patterns helps observers correct their methods and interpret data accurately.

Confusing Shoaling with Schooling

Shoaling refers to fish simply being together in a group, while schooling specifically involves synchronized, polarized swimming. Fish may shoal without schooling, particularly when resting or foraging. Observers must distinguish between these behaviors and record them separately.

Observer Disturbance

The presence of an observer can alter fish behavior, particularly in captive settings. Fish may school more tightly or more loosely in response to human presence. Minimize disturbance by observing from a distance, using one-way glass, or allowing fish to acclimate before recording data.

Inadequate Sample Size

Schooling behavior is variable, and observations of a single group or a single time point may not represent typical behavior. Collect data from multiple groups and multiple time points to account for natural variability.

Ignoring Environmental Variables

Schooling behavior changes with environmental conditions. Failing to record flow velocity, temperature, light levels, and other variables makes it impossible to interpret differences in schooling behavior across observation sessions.

Misidentifying Individual Fish

In large schools, tracking individual fish is challenging. Use natural markings, fin clips, or tagging methods when individual identification is necessary. Alternatively, use group-level metrics that do not require individual identification.

Schooling in Turbulent and Heterogeneous Flow

Fish in natural environments rarely encounter uniform flow conditions. Spatially heterogeneous turbulent flow, which refers to nonuniform flow with coexisting multiple flow velocities, is widely distributed in fish natural and husbandry environments 10.

Behavioral Responses to Flow Heterogeneity

Research on Schizothorax prenanti examined volitional swimming behavior in a complex hydrodynamic environment with three flow velocity regions coexisting in an open-channel flume. The swimming speed of individual fish during upstream migration was significantly higher than that of fish schools. The swimming trajectories of fish schooling showed that they spent more time synchronously exploring the flow environment during upstream migration compared with individual fish 10.

By superimposing fish swimming trajectories on the environmental flow field, researchers quantified the range of hydrodynamic environments preferred by fish in complex flow fields. This approach provides a theoretical reference for the restoration of fish natural habitats or flow enrichment of husbandry environments 10.

Schooling Enhances Upstream Migration

Schooling behavior can significantly enhance upstream migration efficiency. Research on the endangered species Leptobotia elongata found that schooling fish arrived at a target area on average 8.93 seconds earlier than solitary individuals, while flow condition alone had no detectable effect on arrival time 14.

The fish consistently preferred low-velocity zones between 0.20 and 0.50 meters per second and avoided high-velocity regions between 0.75 and 1.25 meters per second, with meandering upstream trajectories predominating. These findings have direct implications for fishway design and habitat restoration 14.

Obstacle Environments

Fish schools navigating obstacle environments exhibit specific collective behaviors and upstream tactics. Research on collective behavior and upstream tactics of schooling fish in obstacle environments has documented how groups coordinate their movements to navigate around barriers while maintaining cohesion 20.

Rheotaxis and Flow Sensing

Rheotaxis is the directed movement resulting from fluid velocity gradients, long studied in fish, aquatic invertebrates, and spermatozoa 6. Fish use rheotaxis to orient themselves relative to water flow, swimming upstream against the current or holding position in flowing water.

The lateral line system plays a critical role in rheotaxis by detecting water flow over the body surface. Fish use this information to maintain their position in the water column and to orient themselves relative to the current. Rheotaxis is distinct from schooling behavior but interacts with it, as fish schools must coordinate their rheotactic responses to maintain group cohesion in flowing water.

Research on bacterial rheotaxis has shown that this behavior can be a purely physical phenomenon resulting from the interplay between velocity gradients and the helical shape of flagella 6. Fish rheotaxis, in contrast, involves active sensory processing and behavioral responses.

Escape Responses and Predator Avoidance

The c-start escape response is one of the most studied unsteady locomotor behaviors exhibited by fishes 7. During a c-start, the fish bends its body into a C shape and then rapidly straightens, propelling itself away from a threat.

Hydrodynamic studies of c-start escape responses have revealed that escape responses generate three distinct vortex rings, each with central orthogonal jet flows. Jet number one is formed by the tail during stage one and moves in the same direction as stage two movement of the fish, thereby reducing final escape velocity but also rotating the fish. Jet number two moves approximately opposite to the final direction of the fish's motion and contains the bulk of the total fluid momentum powering the escape response. Jet number three forms during stage two in the mid-body region and moves in a direction approximately perpendicular to jets one and two 7.

In schooling contexts, escape responses must be coordinated across individuals to be effective. The rapid, synchronized escape of a school can create complex hydrodynamic signals that confuse predators. The overlapping acoustic signals produced by schooling fish may also disrupt predator sensory systems 19.

Air-Breathing Synchrony and Physiological Constraints

Some fish species that breathe air must periodically surface to obtain oxygen. This physiological requirement creates a potential conflict with schooling behavior, as individuals with different oxygen needs may need to surface at different times.

Research on juvenile Arapaima gigas found that individuals differ consistently in surfacing rhythms when alone, yet in a large shoal of about 200 same-aged individuals, a substantial portion of the group surfaces within the same second 13. This synchrony emerges despite individual variation in surfacing rhythms through a simple social interaction rule.

The model developed to explain this behavior suggests that assortative social responsiveness, termed cluster synchrony, can buffer internal constraints, enabling coordination without overriding individual physiological limitations 13. This finding demonstrates that schooling coordination can accommodate individual physiological needs through flexible social interactions.

Robotics and Experimental Approaches

Robotic fish have become valuable tools for studying schooling behavior because they enable repeatable experiments, systematic variation of body and control parameters, and direct measurement of otherwise inaccessible quantities such as internal forces or energy use 12.

A literature analysis reveals that only a minority of robot-fish studies contribute to biological understanding, with most focusing on engineering design. Among biology-oriented studies, closed-loop robotic systems capable of real-time adaptation remain underrepresented but are essential for probing sensorimotor and social feedback mechanisms 12.

Future directions combining robotics, simulations, and emerging experimental technologies aim to unravel the multi-scale feedback loops that shape fish locomotion and schooling 12.

Limitations of Current Research

Understanding the hydrodynamics of fish schooling faces significant challenges. Directly measuring hydrodynamic performance in live fish schools is difficult because the flow structures generated by collective moving organisms are three-dimensional, chaotic, and complex 3.

Numerous previous simulations and experiments have utilized computational, mechanical, or robotic models to represent live fish. Existing studies of live fish schools have contributed significantly to dissecting the complexities of fish schooling, but the scarcity of combined approaches that include both computational and experimental studies of the same fish schools has limited understanding of the physical factors involved in fish collective behavior 3.

An integrated method that combines experiments on live fish schools with computational fluid dynamics simulations represents an innovative method of studying the hydrodynamics of fish schooling. Computational fluid dynamics techniques can deliver accurate performance measurements and high-fidelity flow characteristics for comprehensive analysis. Concurrently, experimental approaches can capture the precise locomotor kinematics of fish and offer additional flow information through particle image velocimetry measurements 3.

A comprehensive review of hydrodynamic studies on fish schooling identified limitations associated with different approaches and suggested potential directions for future work 4. Both numerical simulation and physical experimentation have benefits and drawbacks that researchers must consider when designing studies.

Welfare and Safety Considerations

Observing and managing schooling fish requires attention to welfare considerations. Fish that cannot school may experience chronic stress, particularly species that naturally form large schools. Providing appropriate group sizes and environmental conditions that support natural schooling behavior is important for fish welfare in captivity.

When designing observation studies, minimize disturbance to fish and avoid procedures that cause unnecessary stress. Handle fish gently, maintain appropriate water quality, and provide adequate space for natural schooling behavior.

For researchers working with live fish, follow institutional animal care guidelines and obtain appropriate approvals before conducting experiments. Some schooling studies involve predator exposure or sensory manipulation that may cause stress to fish. These procedures should be justified by the scientific value of the research and conducted with attention to minimizing harm.

Professional Escalation Criteria

Certain observations may indicate problems that require professional intervention. Consult a fish health specialist, aquatic veterinarian, or experienced fisheries biologist if you observe any of the following:

• Schooling behavior suddenly ceases in a species that normally schools, particularly if accompanied by other signs of stress or disease
• Fish within a school show erratic swimming, loss of coordination, or inability to maintain position
• School cohesion breaks down persistently, with fish unable to maintain appropriate distances from neighbors
• Fish show signs of injury, fin damage, or skin lesions that may result from collisions or aggressive interactions within the school
• Schooling behavior changes dramatically following environmental changes such as water quality deterioration, temperature shifts, or flow alterations
• Fish schools fail to respond to predator presence or show abnormal escape responses

Frequently Asked Questions

Why do fish swim in schools?

Fish swim in schools primarily for predator defense, feeding efficiency, and energetic savings. Schools present a confusing target to predators, allow fish to locate food faster, and reduce the energetic cost of swimming through hydrodynamic interactions with neighbors 4. Research on giant danios found that schools reduced their total energy expenditure by 63 to 79 percent compared to solitary fish in turbulent conditions 8.

How do fish coordinate their movements in a school?

Fish coordinate movements through local interactions with neighbors using vision and the lateral line system. Vision provides information about neighbor position and orientation over longer distances, while the lateral line detects water movements and pressure gradients generated by nearby fish 16. Each fish follows simple rules of attraction, repulsion, and alignment relative to its neighbors.

What is the lateral line and how does it help fish school?

The lateral line is a mechanosensory system that detects water movement, pressure gradients, and low-frequency vibrations. It consists of sensory receptors called neuromasts distributed across the body surface and within canals in the head 18. The lateral line allows fish to detect the water movements generated by neighboring fish and to sense flow conditions in their environment.

Can fish school in the dark?

Some fish can school in darkness, but others cannot. Research on giant danios found that they did not school in darkness, indicating that vision is necessary for schooling in this species 16. The ability to school in darkness likely depends on the species and the availability of alternative sensory information.

Do fish from different species school together?

Yes, fish from different species can form mixed-species schools. Research on rosy barbs and tiger barbs found that these species readily form mixed-species schools despite differences in their intrinsic movement patterns 11. In mixed-species groups, the collective motion may be dominated by a single swimming mode, and species may show spatial sorting within the group.

What fish swim under sharks?

Some small fish species swim in association with sharks, using them as mobile shelter or feeding opportunities. These fish may position themselves near or under sharks to avoid predators or to feed on scraps. The specific species that associate with sharks include pilot fish and some species of remoras, though the exact behaviors vary by species and location.

What fish swim with sharks?

Pilot fish and remoras are well-known species that swim with sharks. These associations can provide the smaller fish with protection from predators and access to food resources. The hydrodynamic interactions in these mixed-s

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Combining Computational Fluid Dynamics and Experimental Data to Understand Fish Schooling Behavior.. Integrative and comparative biology, 2024.
• A comprehensive review of hydrodynamic studies on fish schooling.. Bioinspiration & biomimetics, 2023.
• Dipole- and vortex sheet-based models of fish swimming.. Journal of theoretical biology, 2023.
• Bacterial rheotaxis.. Proceedings of the National Academy of Sciences of the United States of America, 2012.
• Hydrodynamics of C-Start Escape Responses of Fish as Studied with Simple Physical Models.. Integrative and comparative biology, 2015.
• Collective movement of schooling fish reduces the costs of locomotion in turbulent conditions.. PLoS biology, 2024.
• A coupled framework for modeling fish schooling.. Bioinspiration & biomimetics, 2026.
• Differences in the Natural Swimming Behavior of Schizothorax prenanti Individual and Schooling in Spatially Heterogeneous Turbulent Flows.. Animals : an open access journal from MDPI, 2023.
• Speed Synchrony Promotes Collective Motion in Mixed-Species Fish Schools. 2026.
• Swimming with robots: investigating fish locomotion, sensing, and schooling behavior with robotic swimmers.. 2026.
• Air-breathing synchrony in juvenile Arapaima gigas reveals collective coordination under individual physiological constraints.. 2026.
• Experimental Study on the Upstream Migration Behavior of Adult &lt,i&gt,Leptobotia elongata&lt,/i&gt, Under Flow Heterogeneity and Schooling in a Controlled Flume System.. 2026.
• The effects of predation stress and contexts on the collective behaviour of crucian carp (Carassius auratus).. 2026.
• The Role of Vision and Lateral Line Sensing for Schooling in Giant Danios (Devario Aequipinnatus). bioRxiv, 2023.
• Three-Dimensional Dynamics Simulation of Fish Schooling Integrating Physiological Characteristics, Movement Patterns, and Multi-Source Perception. Journal of Innovation and Development, 2026.
• The cephalic lateral line system of the pelagic fish Normanichthys crockeri (Clark, 1937) (Pisces: Normanichthyidae) from southern Chile. Latin American Journal of Aquatic Research, 2019.
• Schooling Fish from a New, Multimodal Sensory Perspective. Animals, 2024.
• Collective behavior and upstream tactics of schooling fish in an obstacle environment. Environmental Biology of Fishes, 2025.
• Problems and Prospects of Studying Schooling Behavior of Fish. Journal of Ichthyology, 2023.
• Biophysical Simulations Support Schooling Behavior of Fish Larvae Throughout Ontogeny. Frontiers in Marine Science, 2018.

This article is educational and does not replace institutional policy, professional advice, or applicable safety and regulatory requirements.