The Mechanics of Animal Locomotion: From Walking to Flying

Animal locomotion is the study of how animals move through their environments, encompassing walking, running, jumping, swimming, crawling, and flying. This field integrates biomechanics, neuroscience, evolutionary biology, and robotics to explain how neural circuits generate rhythmic movement patterns, how muscles and skeletons convert neural signals into motion, and how different species have evolved specialized structures for particular habitats. For students, researchers, and life-science professionals, understanding locomotion mechanics provides insight into animal behavior, welfare assessment, and bioinspired engineering. This article explains the major modes of animal locomotion, the neural and biomechanical principles that govern them, and how locomotion research translates into practical applications in animal agriculture and veterinary medicine.

What Is Animal Locomotion

Animal locomotion refers to the self-propelled movement of an animal from one location to another. It requires coordinated activity across multiple biological systems: the nervous system generates and patterns motor commands, the musculoskeletal system executes those commands through muscle contraction and joint movement, and sensory systems provide feedback about body position, external forces, and environmental conditions.

Locomotion serves fundamental biological functions. Animals move to find food, escape predators, locate mates, migrate to favorable environments, and explore territory. The NCBI Literature Resources database catalogs extensive research on these topics, and PubMed provides access to peer-reviewed studies on locomotion physiology, biomechanics, and neural control.

The study of locomotion spans multiple scales of analysis. At the molecular level, researchers investigate how ion channels and signaling proteins within neurons regulate muscle activation. At the systems level, scientists examine how central pattern generators in the spinal cord and brainstem produce rhythmic motor output. At the whole-organism level, biomechanists measure ground reaction forces, joint angles, and energy expenditure during movement. Each scale contributes to a complete understanding of how animals move.

Neural Control of Locomotion

Central Pattern Generators

Central pattern generators (CPGs) are neural circuits that produce rhythmic, patterned output without requiring rhythmic external input. These circuits are fundamental to locomotion across animal groups. Research on insect locomotion demonstrates that CPGs generate the motor patterns underlying flying, walking, swimming, and crawling, with sensory information playing a critical role in establishing functional motor output and coordinating different body parts (Central pattern generating networks in insect locomotion).

CPGs operate through networks of interconnected neurons whose connectivity determines the timing and sequence of muscle activation. In insects, the topology and structure of CPGs vary across locomotion forms, reflecting behavior-specific adaptations. Sensory feedback continuously adjusts CPG output to accommodate changes in terrain, load, and speed.

The practical significance of CPG research extends to robotics. Engineers have developed quadruped robots with locomotion rhythm generators using pulse-type hardware neural networks, demonstrating how biological principles of rhythmic pattern generation can be implemented in artificial systems (Development of quadruped robot with locomotion rhythm generator using pulse-type hardware neural networks).

Brainstem Circuits and Action Diversification

The brainstem serves as a critical interface between higher motor centers involved in action planning and spinal cord circuits that execute body movements. Research using genetic and viral tracing methods has revealed functionally dedicated brainstem populations that control skilled forelimb behavior, orofacial movements, and locomotion. These populations are frequently spatially intermingled but serve distinct functions, and they integrate synaptic inputs from the basal ganglia and cortex to regulate behavioral function in different contexts (Brainstem Circuits Controlling Action Diversification).

The mesopontine tegmentum of the brainstem contains neural mechanisms for initiating locomotion and regulating postural muscle tone. This region receives inhibitory input from the basal ganglia and excitatory input from the limbic-hypothalamic system and neocortex. Functional gating mechanisms in the mesopontine tegmentum determine whether an animal initiates volitionally guided or emotionally triggered locomotion depending on behavioral context (Forebrain control of locomotor behaviors).

Descending Control of Movement

Purposeful locomotion requires descending commands from the brain that modulate low-level rhythmic patterns generated by spinal and brainstem circuits. Research on steering in walking fruit flies identified descending neurons whose activity predicts steering maneuvers. Two specific descending cell types evoke distinct limb gestures: one lengthens strides on the outside of a turn, while the other attenuates strides on the inside of a turn. A single descending neuron can have opposite effects during different locomotor rhythm phases, with networks positioned to implement phase-specific gating (Fine-grained descending control of steering in walking Drosophila).

This research demonstrates that purposeful locomotion emerges from specific, coordinated modulations of low-level movement patterns. The brain does not generate every detail of limb movement, instead, it sends commands that bias and shape the output of lower-level pattern generators.

Neuromodulation and Locomotion

Locomotion is associated with specific patterns of neuromodulator release in the brain. A 2022 study using two-photon fluorescence lifetime imaging measured protein kinase A (PKA) activity within individual spiny projection neurons of the mouse dorsolateral striatum during locomotion. The results showed that dopamine activated PKA activity in direct pathway neurons through the dopamine D1 receptor, while indirect pathway neurons exhibited a greater increase in PKA activity that was largely abolished through blockade of adenosine A2A receptors. Extracellular adenosine increased acutely during locomotion, suggesting that adenosine accumulation interplays with dopamine release to orchestrate PKA activity and proper striatal function during animal locomotion (Locomotion activates PKA through dopamine and adenosine in striatal neurons).

Sensory Integration During Movement

Vision is an active process shaped by actions, intentions, and expectations. During locomotion, internal signals must be integrated with visual information from the retina. Research in mice has advanced understanding of how contextual signals related to locomotion, behavioral relevance, and predictions shape sensory representations in the visual cortex (Contextual signals in visual cortex).

Locomotion alters cortical computation by changing effective synaptic connectivity. Studies imaging four cell types in mouse primary visual cortex found that locomotion activates VIP cells, inhibits SST cells, and disinhibits pyramidal cells in darkness. However, when visual stimuli were present, locomotion increased SST cell responses to large stimuli and VIP cell responses to small stimuli. A recurrent network model successfully predicted each cell type's activity, suggesting that locomotion may alter cortical computation by changing effective synaptic connectivity (Vision and Locomotion Shape the Interactions between Neuron Types in Mouse Visual Cortex).

Sensory cell types in brainstem locomotion centers also respond to environmental stimuli. Fiber photometry recordings in mice found that glutamatergic and GABAergic cells in the cuneiform and pedunculopontine nuclei responded to sound, visual looming, and air puffs, with cell-type specific patterns. Air puffs triggered high-speed locomotion, whereas visual looming and sound stimuli evoked low-speed locomotion. During air puff-evoked locomotion, cuneiform glutamatergic neuron activity was higher than in trials without locomotion, while during locomotion evoked by visual looming or sound, pedunculopontine glutamatergic neuron activity was higher (Cell-type specific sensory and motor activity in the cuneiform nucleus and pedunculopontine nucleus in mice).

Biomechanics of Terrestrial Locomotion

Walking and Running Mechanics

Walking and running involve cyclic movements of the limbs that generate forward propulsion while maintaining stability. The mechanics of these gaits differ fundamentally. In walking, at least one foot remains in contact with the ground at all times, and the body's center of mass rises and falls in an inverted pendulum pattern. In running, there are periods when both feet leave the ground, and the body's center of mass follows a spring-mass pattern with elastic energy storage and return.

Musculoskeletal models provide a bridge between motor neuron activity and joint movements. A 2025 study introduced the first three-dimensional, data-driven musculoskeletal model of Drosophila legs, incorporating Hill-type muscle representations based on high-resolution X-ray scans. Simulations of muscle activity across walking and grooming behaviors predicted coordinated muscle synergies that can be tested experimentally. Training imitation learning policies in simulation showed that damping and stiffness facilitate learning of naturalistic locomotion (Musculoskeletal simulation of limb movement biomechanics in Drosophila melanogaster).

Limb Coordination and Gait Patterns

Terrestrial animals use a variety of gait patterns depending on speed, body size, and environmental conditions. Quadrupedal gaits include walk, trot, and gallop, each characterized by distinct footfall sequences and inter-limb coordination. The choice of gait affects energy expenditure, stability, and maneuverability.

Research on insect locomotion has revealed how CPGs coordinate different body parts during walking. Sensory information is essential for establishing functional motor output, and behavior-specific adaptations reflect the demands of different ecological niches (Central pattern generating networks in insect locomotion).

Locomotion on Complex Substrates

Animals frequently move across substrates that deform or shift under their weight. Research on animal and robotic locomotion on wet granular media examines how movement strategies must adapt to substrates with variable mechanical properties (Animal and Robotic Locomotion on Wet Granular Media). These studies inform both biological understanding and robotic design for challenging environments.

Aquatic Locomotion

Swimming Mechanics

Swimming encompasses diverse mechanisms across aquatic animals. Fish use undulatory body movements, oscillatory fin movements, or jet propulsion depending on species and ecological niche. The mechanical principles governing swimming include drag reduction, thrust generation, and maneuverability in three-dimensional space.

Aquatic locomotion presents unique challenges compared to terrestrial movement. Water is denser and more viscous than air, requiring different strategies for propulsion and energy conservation. Buoyancy reduces the gravitational load on the musculoskeletal system, allowing for different body plans and movement patterns.

Locomotion in Marine Invertebrates

Marine invertebrates exhibit remarkable diversity in locomotion mechanisms. Ctenophores, a pre-bilaterian lineage positioned as the sister group to all other animals, possess a gravity-sensing organ called the aboral statocyst that is analogous to the vertebrate vestibular system. Research on the ctenophore Mnemiopsis leidyi found that the PIEZO ortholog is a mechanically gated ion channel expressed in the statocyst's mechanosensory balancer cells. Chemical modulation with Yoda1 disrupted balancer ciliary beating, impairing postural control and oriented locomotion. Transgenic expression of ctenophore PIEZO in Drosophila piezo mutants rescued mechanosensory phenotypes, establishing functional conservation of PIEZO across ancient animal lineages (PIEZO provides an ancient molecular framework for animal heading control).

Aerial Locomotion

Flight Mechanics

Flight represents one of the most energetically demanding forms of locomotion. Birds, bats, and insects have evolved distinct wing structures and flight muscles adapted to their body sizes and ecological niches. The biomechanics of flight involve generating lift and thrust through wing movements, managing aerodynamic forces, and controlling stability and maneuverability.

Insects utilize flight as one of several locomotion forms during their life cycle. The CPGs underlying insect flight generate rhythmic motor patterns that coordinate wing movements, with sensory feedback adjusting wing kinematics in response to aerodynamic conditions (Central pattern generating networks in insect locomotion).

Gliding and Soaring

Some animals achieve aerial locomotion through gliding instead of powered flight. Gliding involves descending through the air with wings or other aerodynamic surfaces extended, using gravity to maintain forward motion. Soaring uses rising air currents to maintain or gain altitude with minimal energy expenditure. These strategies are energetically efficient but require specific environmental conditions.

Locomotion in Arthropods

Arthropods represent the largest animal phylum and occupy terrestrial, aquatic, arboreal, and subterranean niches. Their evolutionary success depends on specific morphological and biomechanical adaptations related to their materials and structures. Modern research methods including imaging techniques, mechanical testing, movement capture, and numerical modeling have advanced understanding of arthropod flight, locomotion, and attachment mechanisms (Functional morphology and biomechanics of arthropods).

The exoskeleton of arthropods provides both support and attachment points for muscles. Joint structure and muscle arrangement determine the range and direction of limb movements. The diversity of arthropod locomotion strategies reflects adaptations to different substrates, from the complex three-dimensional environment of vegetation to the open surfaces of desert sands.

Locomotion Assessment in Animal Agriculture

Locomotion Scoring in Dairy Cattle

Locomotion assessment is a critical component of dairy herd management. Lameness is a common issue on dairy farms with serious implications for economy and animal welfare. Affected animals may be overlooked until their condition becomes severe, making improved detection methods necessary.

Research using inertial measurement units attached to predefined locations on cows' upper body and limbs has characterized kinematic changes in dairy cows with induced, mild to moderate hindlimb lameness. Analysis of over 3,000 stride cycles from 41 straight-line walk measurements found significant differences between baseline and lameness conditions in 23 of 31 kinematic parameters. Lameness induction was associated with decreased maximum protraction and retraction angles of the distal portion of the induced and non-induced limbs respectively (Kinematic changes in dairy cows with induced hindlimb lameness: transferring methodology from the field of equine biomechanics).

Genetic Basis of Locomotion Traits

Locomotion traits in dairy cattle have a genetic component that can be improved through selective breeding. Research on Swiss Holstein and Brown Swiss cattle estimated heritabilities ranging from 0.10 to 0.28 for locomotion-related traits including bone structure, heel depth, foot angle, and rear leg conformation. Genome-wide association studies detected significant associated regions for bone structure and locomotion in Holstein cattle and for heel depth and hock quality in Brown Swiss cattle. A nonsense variant in the HYAL1 gene showed a significant recessive effect on bone structure in Holstein cattle, with adverse effects on six-year survival (Genomic dissection of hoof and leg conformation in Swiss dairy cattle populations reveals polygenic architecture and a recessive HYAL1 nonsense variant affecting longevity in Holstein cattle).

Genetic parameters for locomotion, body condition score, and linear type traits have been estimated in Czech Holstein cattle (Genetic parameters for female fertility, locomotion, body condition score, and linear type traits in Czech Holstein cattle) and in Holstein cattle in southern China (Genetic parameters estimates for locomotion score, body condition score and final type score of Holstein cattle in southern China). These studies provide the genetic basis for including locomotion traits in breeding programs.

Practical Locomotion Assessment Protocol

Farmers and veterinarians can implement a structured locomotion assessment protocol using the following steps:

1. Observe cows walking on a flat, non-slip surface with adequate space for several strides
2. Score each cow using a standardized locomotion scoring system that evaluates gait symmetry, stride length, back arch, and head bob
3. Record scores in a herd health management system with dates and animal identification
4. Investigate cows with locomotion scores indicating moderate or severe lameness
5. Examine feet and legs of affected cows for injuries, infections, or conformational abnormalities
6. Consult a veterinarian for diagnosis and treatment planning when lameness persists or affects multiple animals
7. Monitor response to treatment and track locomotion scores over time to evaluate intervention effectiveness

Records and Measurements

Maintaining accurate locomotion records supports herd health management and genetic improvement. Recommended records include:

At a Glance: Major Locomotion Modes

Practical Applications of Locomotion Research

Robotics and Bioinspired Design

Locomotion research informs the design of mobile robotic systems. Engineers study biological locomotion principles to develop robots capable of navigating diverse terrains. Research on locomotion principles for mobile robotic systems examines how different movement mechanisms suit different operational requirements (Locomotion Principles for Mobile Robotic Systems).

Quadruped robots with locomotion rhythm generators based on pulse-type hardware neural networks demonstrate how CPG principles can be implemented in artificial systems (Development of quadruped robot with locomotion rhythm generator using pulse-type hardware neural networks). These robots achieve naturalistic locomotion through rhythmic pattern generation analogous to biological CPGs.

Movement Analysis Technology

Advances in movement analysis technology enable detailed study of animal locomotion in natural environments. A method for reconstructing three-dimensional animal trajectories from monocular video combines deep learning tools and the pinhole camera model. This approach was tested in aerial-based and ground-based scenarios and applied to a bat-predation biomechanics study. The estimated three-dimensional coordinates had an average bias of 0.09 meters for aerial motion and 0.044 meters for ground motion, with robust distance estimation when faced with foreground occlusion (A simple method for rapid reconstruction of 3D animal trajectory from monocular video).

This low-cost, single-camera approach allows analysis of animal locomotion in natural environments without requiring multiple devices or precise calibration. Researchers can extract kinematic parameters and gait frequencies from video recordings, facilitating intelligent monitoring of wild animals and enhancing understanding of their locomotion data.

Biomechanical Modeling

Computational models are critical for understanding how neural, biomechanical, and physical systems interact to orchestrate animal behaviors. The first three-dimensional, data-driven musculoskeletal model of Drosophila legs, implemented in both OpenSim and MuJoCo simulation environments, incorporates Hill-type muscle representations based on high-resolution X-ray scans. This model enables investigation of motor control in an experimentally tractable model organism and can be used to control embodied artificial agents to generate naturalistic and compliant locomotion in simulated environments (Musculoskeletal simulation of limb movement biomechanics in Drosophila melanogaster).

Locomotion and Aging

Locomotion capacity changes with age, and these changes can serve as indicators of overall health and functional decline. Intrinsic capacity, a crucial determinant of healthy aging, is operationalized through five essential functional domains: locomotion, cognition, vitality, psychological, and sensory capacities. Longitudinal studies in mouse and fish models found that locomotor capacity, measured via voluntary activity and rotarod, showed initial impairment in middle age followed by progressive decline, especially in female mice. In contrast, impairments in locomotor activity in killifish emerged predominantly in the last third of life (Intrinsic capacity evolution during aging in mouse and fish: Longitudinal perspectives from a narrative review).

These findings have implications for animal agriculture, where locomotion changes can signal health problems that affect productivity and welfare. Monitoring locomotion over time can identify animals at risk of mobility-related conditions and support timely intervention.

Common Failure Patterns in Locomotion

Neurological Causes

Locomotion failure can result from dysfunction at any level of the motor system. Damage to CPGs, descending pathways, or sensory feedback circuits disrupts coordinated movement. Neurological conditions may present as ataxia, paresis, or abnormal gait patterns.

Musculoskeletal Causes

Injuries to bones, joints, muscles, tendons, or ligaments impair locomotion. Lameness in livestock commonly results from hoof lesions, joint infections, or traumatic injuries. Conformational abnormalities can predispose animals to locomotion problems.

Metabolic and Nutritional Causes

Metabolic disorders can affect locomotion through effects on muscle function, nerve function, or bone health. Nutritional deficiencies or imbalances may contribute to musculoskeletal problems.

Environmental Causes

Slippery floors, poorly designed handling facilities, and inadequate space can increase the risk of locomotion injuries. Environmental factors that cause stress or fear may also affect movement patterns.

Limitations of Locomotion Research

Species Differences

Findings from one species do not always transfer to others. Neural circuits, biomechanical properties, and locomotion strategies vary across species, requiring species-specific research for accurate understanding.

Laboratory Versus Natural Conditions

Laboratory studies may not capture the full complexity of locomotion in natural environments. Terrain variability, predator pressure, and social factors influence movement patterns in ways that are difficult to replicate in controlled settings.

Measurement Challenges

Accurate measurement of locomotion requires appropriate technology and methodology. Some measurement approaches require multiple cameras or specialized equipment, limiting their application in field settings. The development of monocular video reconstruction methods addresses some of these limitations (A simple method for rapid reconstruction of 3D animal trajectory from monocular video).

Individual Variation

Locomotion parameters vary among individuals due to genetics, age, health status, and experience. Understanding this variation is important for interpreting locomotion measurements and making management decisions.

Welfare and Safety Considerations

Locomotion as a Welfare Indicator

Locomotion quality is a widely used indicator of animal welfare. Impaired locomotion is associated with pain, discomfort, and reduced ability to perform natural behaviors. Regular locomotion assessment supports early detection of health problems and timely intervention.

Safe Handling of Animals with Locomotion Impairment

Animals with locomotion impairment require special handling considerations. Providing non-slip flooring, reducing required walking distances, and ensuring access to food, water, and resting areas supports recovery and prevents further injury.

Human Safety During Locomotion Assessment

Locomotion assessment requires working in close proximity to animals. Assessors should follow established safety protocols, including maintaining escape routes, using appropriate handling equipment, and being alert to animal behavior that indicates agitation or fear.

Professional Escalation Criteria

Farmers and animal caretakers should seek professional veterinary assistance when:

1. Locomotion scores indicate moderate or severe lameness that does not improve with basic foot care
2. Multiple animals develop locomotion problems within a short period, suggesting a possible infectious or environmental cause
3. Locomotion impairment is accompanied by systemic signs such as fever, reduced appetite, or weight loss
4. Animals are unable to bear weight on a limb or show signs of severe pain
5. Locomotion problems persist despite treatment, suggesting the need for diagnostic imaging or specialized care
6. Genetic testing reveals risk variants for locomotion-related conditions, requiring breeding plan adjustments

Frequently Asked Questions

What is animal locomotion?

Animal locomotion is the self-propelled movement of an animal from one location to another. It involves coordinated activity of the nervous system, which generates and patterns motor commands, and the musculoskeletal system, which executes those commands through muscle contraction and joint movement. Locomotion serves essential biological functions including finding food, escaping predators, locating mates, and exploring territory.

What are the main types of animal locomotion?

The main types of animal locomotion are walking, running, jumping, swimming, crawling, and flying. Each mode involves distinct biomechanical principles and neural control mechanisms. Walking uses inverted pendulum mechanics with continuous ground contact, running uses spring-mass mechanics with aerial phases, swimming involves undulatory or fin-based propulsion in water, and flying generates lift and thrust through wing movements.

How do central pattern generators control locomotion?

Central pattern generators are neural circuits that produce rhythmic, patterned output without requiring rhythmic external input. They generate the motor patterns underlying walking, flying, swimming, and crawling. Sensory information continuously adjusts CPG output to accommodate changes in terrain, load, and speed, and descending commands from the brain modulate CPG activity to produce purposeful movement (Central pattern generating networks in insect locomotion).

How is locomotion assessed in dairy cattle?

Locomotion in dairy cattle is assessed using standardized scoring systems that evaluate gait symmetry, stride length, back arch, and head bob. Objective kinematic measurements using inertial measurement units can detect subtle changes associated with lameness, including decreased maximum protraction and retraction angles of the limbs (Kinematic changes in dairy cows with induced hindlimb lameness: transferring methodology from the field of equine biomechanics). Genetic evaluations for locomotion traits support selective breeding for improved mobility.

What causes lameness in livestock?

Lameness in livestock can result from hoof lesions, joint infections, traumatic injuries, conformational abnormalities, metabolic disorders, and nutritional deficiencies. Environmental factors such as slippery floors and poorly designed facilities increase the risk of locomotion injuries. Genetic factors also contribute to locomotion problems, with heritability estimates for locomotion-related traits ranging from 0.10 to 0.28 in dairy cattle (Genomic dissection of hoof and leg conformation in Swiss dairy cattle populations reveals polygenic architecture and a recessive HYAL1 nonsense variant affecting longevity in Holstein cattle).

How does locomotion research inform robotics?

Locomotion research informs robotics through the application of biological principles to artificial systems. Engineers have developed quadruped robots with locomotion rhythm generators based on pulse-type hardware neural networks that mimic biological CPGs (Development of quadruped robot with locomotion rhythm generator using pulse-type hardware neural networks). Musculoskeletal models of animal limbs enable the design of robots that generate naturalistic and compliant locomotion in simulated environments (Musculoskeletal simulation of limb movement biomechanics in Drosophila melanogaster).

How does locomotion change with age?

Locomotion capacity typically declines with age, although the pattern of decline varies across species. Longitudinal studies in mice found that locomotor capacity showed initial impairment in middle age followed by progressive decline, especially in females. In killifish, impairments in locomotor activity emerged predominantly in the last third of life (Intrinsic capacity evolution during aging in mouse and fish: Longitudinal perspectives from a narrative review). Monitoring locomotion over time can identify age-related changes that affect health and welfare.

What technology is available for measuring animal locomotion?

Technology for measuring animal locomotion includes inertial measurement units attached to the body, motion capture systems using multiple cameras, and monocular video reconstruction methods. A recent method combining deep learning tools and the pinhole camera model reconstructs three-dimensional animal trajectories from single-camera video with average bias of 0.09 meters for aerial motion and 0.044 meters for ground motion (A simple method for rapid reconstruction of 3D animal trajectory from monocular video). This low-cost approach enables locomotion analysis in natural environments.

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References and Further Reading

• NCBI Literature Resources. National Center for Biotechnology Information.
• PubMed. National Library of Medicine.
• Locomotion activates PKA through dopamine and adenosine in striatal neurons.. Nature, 2022.
• Brainstem Circuits Controlling Action Diversification.. Annual review of neuroscience, 2019.
• Contextual signals in visual cortex.. Current opinion in neurobiology, 2018.
• Fine-grained descending control of steering in walking Drosophila.. Cell, 2024.
• Central pattern generating networks in insect locomotion.. Developmental neurobiology, 2020.
• Forebrain control of locomotor behaviors.. Brain research reviews, 2008.
• Cell-type specific sensory and motor activity in the cuneiform nucleus and pedunculopontine nucleus in mice.. Scientific reports, 2025.
• Vision and Locomotion Shape the Interactions between Neuron Types in Mouse Visual Cortex.. Neuron, 2018.
• PIEZO provides an ancient molecular framework for animal heading control. 2026.
• Intrinsic capacity evolution during aging in mouse and fish: Longitudinal perspectives from a narrative review.. 2026.
• Genomic dissection of hoof and leg conformation in Swiss dairy cattle populations reveals polygenic architecture and a recessive HYAL1 nonsense variant affecting longevity in Holstein cattle.. 2026.
• Musculoskeletal simulation of limb movement biomechanics in Drosophila melanogaster. arXiv.org, 2025.
• Functional morphology and biomechanics of arthropods. Journal of Comparative Physiology, 2023.
• Investigating the Biomechanics to Mimic Blinking using an Electromagnetic Actuator. Annual International Conference of the IEEE Engineering in Medicine and Biology Society, 2025.
• CHARMS - re-imagining the body in motion: the embodied armoured flesh and the biomechanics research lab. RUUKKU Studies in Artistic Research, 2025.
• Kinematic changes in dairy cows with induced hindlimb lameness: transferring methodology from the field of equine biomechanics.. Animal, 2024.
• A simple method for rapid reconstruction of 3D animal trajectory from monocular video. Movement Ecology, 2025.
• Development of quadruped robot with locomotion rhythm generator using pulse-type hardware neural networks. Artificial Life and Robotics, 2015.
• Animal and Robotic Locomotion on Wet Granular Media. Lecture Notes in Computer Science Including Subseries Lecture Notes in Artificial Intelligence and Lecture Notes in Bioinformatics, 2017.
• Genetic parameters for female fertility, locomotion, body condition score, and linear type traits in Czech Holstein cattle. Journal of Dairy Science, 2011.
• Genetic parameters estimates for locomotion score, body condition score and final type score of Holstein cattle in southern China. Journal of Applied Animal Research, 2013.
• Locomotion Principles for Mobile Robotic Systems. Procedia Computer Science, 2017.

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