How Animals Move: A Guide to Locomotion Types

Animal locomotion is the study of how animals move through their environments, encompassing walking, running, jumping, swimming, flying, and crawling. This guide explains the main categories of animal movement, the biomechanical principles behind each mode, and the neural circuits that coordinate these behaviors. Understanding locomotion matters for farmers, veterinarians, animal scientists, and researchers because movement quality directly affects animal welfare, productivity, and health assessment. This article covers the major locomotion types, their underlying mechanisms, and practical applications for observing and evaluating animal movement.

At a Glance: Locomotion Types and Key Adaptations

The table below summarizes the primary locomotion categories, representative examples, and the critical adaptations that enable each form of movement.

The Neural Basis of Locomotion

Central Pattern Generators

Locomotion originates from rhythmic limb movement patterns that are generated by circuits outside the brain. Central pattern generators are neural circuits that produce rhythmic and patterned output without requiring rhythmic external inputs. This property makes them essential for generating many kinds of rhythmic motor behaviors in insects, including flying, walking, swimming, and crawling. Insects represent the most diverse group of animals and utilize at least one of these locomotion types during some stage of their development. Researchers have extensively studied insects to understand the neural basis of rhythmic motor behaviors and the structure and operation of central pattern generators involved in locomotion. The contribution of central pattern generation to flying, walking, and crawling in insects reveals how the topology and structure of these circuits influence the generation of respective motor patterns. Sensory information is critical for establishing functional motor output, and behavior-specific adaptations indicate how coordination between different body parts is achieved. See the NCBI Literature Resources and the PubMed record on central pattern generating networks in insect locomotion for further background.

Brainstem Control of Movement

The brainstem serves as an important interface between upper motor centers involved in action planning and circuits in the spinal cord that execute body movements. Recent work using genetic and viral entry points has revealed functionally dedicated and frequently spatially intermingled brainstem populations essential for action diversification, a general principle conserved throughout evolution. Brainstem circuits with distinct organization and function control skilled forelimb behavior, orofacial movements, and locomotion. These circuits convey regulatory parameters to motor output structures and collaborate in the construction of complex natural motor behaviors. Functionally tuned brainstem neurons for different actions serve as important integrators of synaptic inputs from upstream centers, including the basal ganglia and cortex, to regulate and modulate behavioral function in different contexts. See the PubMed record on brainstem circuits controlling action diversification for details.

Forebrain Contributions to Locomotor Behavior

Activation of different areas in the forebrain evokes different types of goal-directed adaptive behaviors. An important component of these behavioral patterns is the locomotion that brings the animal to or away from a particular location. Projections from forebrain structures to the mesopontine tegmentum of the brainstem contain neural mechanisms for initiation of locomotion and regulation of postural muscle tone that are activated during locomotor behavior. The mesopontine tegmentum receives inhibitory efferents from the basal ganglia and excitatory efferents from the limbic-hypothalamic system and the neocortex. Functional gating mechanisms in the mesopontine tegmentum determine whether the subject will initiate and select volitionally guided or emotionally triggered locomotor behaviors, depending on the behavioral context. See the PubMed record on forebrain control of locomotor behaviors for further reading.

Molecular Signaling During Locomotion

The canonical model of striatal function predicts that animal locomotion is associated with opposing regulation of protein kinase A in direct and indirect pathway striatal spiny projection neurons by dopamine. However, precise dynamics of protein kinase A in dorsolateral spiny projection neurons during locomotion remained unclear, as did the involvement of other neuromodulators. Research using two-photon fluorescence lifetime imaging of a protein kinase A sensor through gradient index lenses measured protein kinase A activity within individual spiny projection neurons of the mouse dorsolateral striatum during locomotion. Consistent with the canonical view, dopamine activated protein kinase A activity in direct pathway spiny projection neurons during locomotion through the dopamine D1 receptor. Indirect pathway spiny projection neurons exhibited a greater increase in protein kinase A activity, which was largely abolished through the blockade of adenosine A2A receptors. Fiber photometry measurements of an adenosine sensor revealed an acute increase in extracellular adenosine during locomotion. Antagonism of dopamine or adenosine receptors resulted in distinct changes in spiny projection neuron protein kinase A activity, neuronal activity, and locomotion. These results suggest that acute adenosine accumulation interplays with dopamine release to orchestrate protein kinase A activity in spiny projection neurons and proper striatal function during animal locomotion. See the PubMed record on locomotion activating PKA through dopamine and adenosine for the full study.

Sensory Integration and Locomotion

Vision is an active process, and what we perceive strongly depends on our actions, intentions, and expectations. During visual processing, internal signals need to be integrated with visual information from the retina. Advances in recording and manipulating neuronal activity in specific cell types and axonal projections, together with tools for circuit tracing, are beginning to reveal the neuronal circuit mechanisms of how internal contextual signals shape sensory representations. Research primarily in mice has advanced understanding of these processes, focusing on contextual signals related to locomotion, behavioral relevance, and predictions. See the PubMed record on contextual signals in visual cortex for details.

Cortical computation arises from the interaction of multiple neuronal types, including pyramidal cells and interneurons expressing somatostatin, vasoactive intestinal peptide, or parvalbumin. Recordings in darkness were consistent with a disinhibitory model in which locomotion activates vasoactive intestinal peptide cells, thus inhibiting somatostatin cells and disinhibiting pyramidal cells. However, the disinhibitory model failed when visual stimuli were present. Locomotion increased somatostatin cell responses to large stimuli and vasoactive intestinal peptide cell responses to small stimuli. A recurrent network model successfully predicted each cell type's activity from the measured activity of other types. Capturing the effects of locomotion required allowing it to increase feedforward synaptic weights and modulate recurrent weights. This network model summarizes interneuron interactions and suggests that locomotion may alter cortical computation by changing effective synaptic connectivity. See the PubMed record on vision and locomotion shaping neuron type interactions for the full study.

Cell-Type Specific Activity in Locomotion Centers

The activity of neurotransmitter-based cell types in the cuneiform and pedunculopontine nuclei during locomotion, non-locomotor behaviors, and following sensory stimulation is not fully understood. Using fiber photometry in mice, researchers found cell-type specific responses to sensory stimuli. Glutamatergic and GABAergic cells responded to sound, visual looming, and air puffs, except for pedunculopontine GABAergic cells, which did not respond to visual looming. Cholinergic cells responded to air puffs. 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. During locomotion evoked by visual looming or sound, activity in pedunculopontine glutamatergic neurons was higher than when no locomotion occurred. In the open-field arena, mice exhibited spontaneous low-speed locomotion during which activity increased in pedunculopontine glutamatergic cells. Activity also increased in a cell-type specific manner during grooming or rearing. This study shows cell-type specific activity in the cuneiform or pedunculopontine nuclei during locomotion, non-locomotor behaviors, and following sensory stimulation. Sensory responsiveness likely has relevance in Parkinson's disease, where sensory circuits are increasingly targeted to improve walking. See the PubMed record on cell-type specific sensory and motor activity in locomotion nuclei for the full study.

Terrestrial Locomotion

Walking and Gait Patterns

Walking is the most common terrestrial locomotion mode and involves coordinated limb movements that maintain stability while supporting body weight. The neural control of walking requires descending commands from the brain that modulate rhythmic limb movement patterns originating in circuits outside the brain. Research on walking Drosophila has identified descending neurons whose activity predicts steering maneuvers. Two descending cell types downstream of distinct brain networks evoke specific 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, and networks are positioned to implement this phase-specific gating. Purposeful locomotion emerges from specific, coordinated modulations of low-level patterns. See the PubMed record on fine-grained descending control of steering in walking Drosophila for the full study.

For livestock producers, observing gait patterns is a routine management practice. Lameness detection relies on identifying deviations from normal walking patterns. Cattle with locomotion issues show altered stride length, reduced weight bearing on affected limbs, and changes in back posture during walking. Genetic parameters for locomotion in dairy cattle have been estimated in breeding programs, indicating that locomotion traits are heritable and can be improved through selection. See the Elsevier record on genetic parameters for locomotion in Czech Holstein cattle for the study details.

Running and Elastic Energy Storage

Running involves a phase where all limbs leave the ground simultaneously, requiring greater muscle power and elastic energy storage compared to walking. Tendons play a critical role in running by storing and releasing elastic energy during each stride. The nuchal ligament in horses demonstrates this principle. The horse's head and neck account for about 10% of body weight, and this combined with their cantilevered position in front of the trunk produces considerable leverage around the joints at the base of the neck. During locomotion, the neck is moved primarily by gravitational and inertial forces controlled by eccentric-concentric cycles of contraction in the topline muscles assisted by energy-saving stretch-recoil cycles in the nuchal ligament. The functional part of the nuchal ligament is the caudal funicular region and the lamellae to the second cervical vertebra that acts as the point of suspension for the entire neck. See the record on biomechanics of the head and neck for details.

Jumping and Leaping

Jumping requires rapid generation of force to overcome gravity and propel the body through the air. Animals that jump effectively typically have powerful hindlimb muscles, elastic energy storage mechanisms, and lightweight skeletons. Frogs and kangaroos exemplify specialized jumping adaptations. Grasshoppers use elastic energy stored in their exoskeleton to achieve rapid takeoff. The biomechanical principles of jumping involve force production, energy storage, and precise timing of muscle activation.

Aquatic Locomotion

Swimming Mechanics

Swimming involves propulsion through water using various body structures. Fish use undulatory body movements and fins to generate thrust. Whales and seals use flippers and tail movements for propulsion. Streamlined body shapes reduce drag and improve swimming efficiency. The density and viscosity of water create different mechanical challenges compared to terrestrial locomotion. Aquatic animals must manage buoyancy, drag, and thrust generation simultaneously.

Locomotion on Wet Granular Media

Some animals move on substrates that are neither solid ground nor open water. Wet granular media such as mud, sand, and wet soil present unique locomotion challenges. Research on animal and robotic locomotion on wet granular media examines how animals adjust their movement strategies on these substrates. See the Elsevier record on animal and robotic locomotion on wet granular media for the publication details.

Aerial Locomotion

Flight Mechanics

Flight requires specialized anatomical adaptations including wings, lightweight bones, powerful flight muscles, and aerodynamic body shapes. Birds, bats, and insects have evolved distinct flight mechanisms. The Pterosauria comprises Mesozoic flying reptiles whose unique anatomy imposes challenging constraints for biomechanical studies. These animals left no modern descendants, making understanding their motion dynamics challenging. Research relies heavily on technological devices including modeling, 3D simulations, and computer graphics software. The number of articles on pterosaur locomotion has increased continuously since the 1990s. Evolution and flight were the most frequent words cited together in scientific production. Flight was the trending topic before 2001 and played an important role as expected for volant reptiles. Further scientific studies are needed to better understand land locomotion, musculoskeletal systems, mechanostat, density and properties, neuromechanical aspects, and biomimetics in pterosaurs. An increasing integration between paleontology and fields such as engineering, physics, and computational sciences is anticipated. This interdisciplinary approach is expected to advance understanding of the movement dynamics of these flying reptiles while also enabling the application of their biological principles in the development of bioinspired engineering technologies. See the record on pterosaur biomechanics for the full review.

Insect Flight

Insects utilize central pattern generators for flying, walking, swimming, and crawling. The contribution of central pattern generation to flying in insects involves specific neural circuit topologies that generate the rhythmic wing movements required for flight. Sensory information is important for establishing functional motor output during flight, and behavior-specific adaptations indicate how coordination between different body parts is achieved. See the PubMed record on central pattern generating networks in insect locomotion for details.

Crawling and Limbless Locomotion

Muscular Hydrostats

Crawling involves movement using muscular hydrostats, which are structures composed primarily of muscle tissue that maintain volume while changing shape. Earthworms use segmental muscles and bristles to move through soil. Snakes use lateral undulation, rectilinear movement, and other strategies depending on substrate conditions. Caterpillars use a combination of segmental muscles and prolegs for gripping surfaces.

Subterranean Locomotion

Animals that move through soil face unique challenges including high friction, limited visibility, and the need to displace substrate material. Earthworms use peristaltic contractions to move through soil. Burrowing animals such as moles have specialized forelimbs for digging. The mechanical properties of soil affect locomotion efficiency and energy expenditure.

Practical Assessment of Locomotion in Farm Animals

Observing Locomotion in the Field

Farmers and veterinarians routinely assess locomotion in livestock. A structured observation protocol improves consistency and early detection of problems. The following steps provide a practical framework for locomotion assessment.

First, observe animals in a familiar environment with level, non-slip footing. Allow animals to move freely without pressure. Second, watch from the side to evaluate stride length, foot placement, and back posture. Third, watch from behind to assess pelvic symmetry and weight distribution. Fourth, observe animals turning to detect reluctance or stiffness. Fifth, compare the animal to herd mates to identify deviations from normal movement patterns.

Recording Locomotion Scores

Locomotion scoring systems provide a standardized method for documenting movement quality. A simple numerical scale allows consistent recording and trend monitoring. Record the animal identification, date, environment conditions, and locomotion score. Note any visible injuries, swelling, or heat in limbs. Track changes over time to identify worsening or improvement.

Common Failure Patterns in Locomotion Assessment

Several common errors reduce the reliability of locomotion assessment. Observing animals on slippery or uneven surfaces can mask or exaggerate gait abnormalities. Assessing animals immediately after transport or handling may reflect fatigue instead of chronic conditions. Failing to account for individual variation in normal gait can lead to false positives. Inconsistent scoring between observers reduces data quality. Infrequent assessment misses gradual deterioration.

Professional Escalation Criteria

Certain observations warrant professional veterinary evaluation. Acute lameness with non-weight bearing requires immediate attention. Swelling, heat, or wounds in limbs should be examined promptly. Locomotion changes accompanied by fever, reduced appetite, or other systemic signs require veterinary assessment. Worsening locomotion despite management changes indicates the need for professional diagnosis. Herd-level increases in locomotion scores suggest environmental or management factors requiring investigation.

Biomechanical Principles Across Locomotion Types

Force Generation and Transmission

All forms of locomotion require force generation by muscles and transmission through skeletal structures to the environment. The musculoskeletal system converts muscle contraction into movement. Tendons transmit force from muscles to bones and can store elastic energy. Joints allow controlled movement while maintaining stability.

Energy Efficiency

Different locomotion modes have different energy costs. Walking is generally more energy-efficient than running at low speeds, while running becomes more efficient at higher speeds. Elastic energy storage in tendons improves running efficiency. Swimming is energetically efficient due to buoyancy support but faces drag forces. Flight is energetically expensive but enables rapid travel over long distances.

Stability and Control

Maintaining stability during locomotion requires continuous sensory feedback and motor adjustment. The vestibular system provides information about head position and movement. Proprioceptors in muscles and joints provide information about limb position. Visual input helps animals navigate and avoid obstacles. The integration of these sensory signals allows animals to adapt their locomotion to changing conditions.

Robotics and Locomotion Principles

Bioinspired Locomotion

Understanding animal locomotion has informed the development of robotic systems. Quadruped robots with locomotion rhythm generators using pulse-type hardware neural networks demonstrate how biological principles can be applied to engineering. See the Elsevier record on quadruped robot locomotion rhythm generators for the publication details.

Locomotion Principles for Mobile Robotic Systems

Mobile robotic systems apply locomotion principles derived from biological systems. The study of locomotion principles for mobile robotic systems examines how different movement strategies can be implemented in engineered platforms. See the Elsevier record on locomotion principles for mobile robotic systems for the publication details.

Welfare and Safety Context

Locomotion as a Welfare Indicator

Locomotion quality serves as an important welfare indicator in farm animals. Impaired locomotion often indicates pain, injury, or disease. Regular locomotion assessment allows early detection and intervention. Poor locomotion can reduce feed intake, productivity, and reproductive performance. Addressing locomotion problems improves both animal welfare and farm profitability.

Safe Handling During Locomotion Assessment

When assessing locomotion, prioritize handler safety. Work with animals in well-designed handling facilities. Avoid entering pens with animals that are agitated or fearful. Use appropriate restraint when necessary. Maintain escape routes and follow established safety protocols. Document any incidents and review handling procedures regularly.

Limitations and Knowledge Gaps

Species-Specific Differences

Locomotion mechanisms vary substantially across species. Findings from one species may not directly apply to others. Research on mice, Drosophila, and other model organisms provides insights into general principles, but species-specific adaptations require direct study. Farmers and researchers should exercise caution when extrapolating findings across species.

Environmental Influences

Locomotion is influenced by environmental conditions including substrate, temperature, and terrain. Laboratory studies may not fully capture the complexity of natural locomotion. Field observations complement laboratory research by documenting locomotion under real-world conditions.

Ongoing Research

Research on animal locomotion continues to advance understanding of neural control, biomechanics, and evolutionary adaptations. New technologies including advanced imaging, genetic tools, and computational modeling are expanding research capabilities. The integration of multiple disciplines including neuroscience, biomechanics, and engineering is expected to yield further insights.

Frequently Asked Questions

What is animal locomotion?

Animal locomotion is the study of how animals move through their environments. It encompasses all forms of movement including walking, running, jumping, swimming, flying, and crawling. The field examines the biomechanical principles, neural control mechanisms, and evolutionary adaptations that enable animal movement.

What are the main types of animal locomotion?

The main types of animal locomotion are walking, running, jumping, swimming, flying, and crawling. Each type involves distinct biomechanical adaptations and is suited to particular environments. Some animals use multiple locomotion types depending on their life stage or environmental conditions.

How do central pattern generators control locomotion?

Central pattern generators are neural circuits that produce rhythmic and patterned output without requiring rhythmic external inputs. They generate the basic rhythmic patterns for walking, flying, swimming, and crawling. Sensory information modulates central pattern generator output to produce functional movement adapted to current conditions.

Why is locomotion assessment important in farm animals?

Locomotion assessment is important because movement quality reflects animal health and welfare. Impaired locomotion often indicates pain, injury, or disease. Regular assessment allows early detection and intervention, which improves animal welfare and reduces economic losses.

How does elastic energy storage improve running efficiency?

Elastic energy storage in tendons allows animals to store energy during one phase of the stride and release it during another phase. This reduces the muscular work required for running. The nuchal ligament in horses demonstrates this principle by storing and releasing energy during neck movement.

What role does the brainstem play in locomotion?

The brainstem serves as an interface between upper motor centers involved in action planning and spinal cord circuits that execute movement. Brainstem circuits control skilled forelimb behavior, orofacial movements, and locomotion. They integrate inputs from the basal ganglia and cortex to regulate behavioral function in different contexts.

How do sensory signals influence locomotion?

Sensory signals provide information about the environment and the animal's position within it. Visual, vestibular, and proprioceptive inputs are integrated to guide movement. Research shows that locomotion itself influences sensory processing, with locomotion altering cortical computation by changing effective synaptic connectivity.

What are the limitations of current locomotion research?

Current locomotion research has several limitations. Findings from model organisms may not directly apply to other species. Laboratory conditions may not fully capture natural locomotion complexity. Many studies focus on single locomotion types instead of the integration of multiple movement modes. Ongoing research aims to address these gaps.

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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.
• Biomechanics of the Head and Neck.. 2026.
• Gaining Ground On Pterosaur Biomechanics: A General Overview.. 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.
• 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.