Animal locomotion
Encyclopedia
Animal locomotion, which is the act of self-propulsion by an animal, has many manifestations, including running
, swimming
, jumping
and flying
. Animals move for a variety of reasons, such as to find food, a mate, or a suitable microhabitat, and to escape predators. For many animals the ability to move is essential to survival and, as a result, selective pressures have shaped the locomotion methods and mechanisms employed by moving organisms. For example, migratory animals that travel vast distances (such as the Arctic Tern
) typically have a locomotion mechanism that costs very little energy per unit distance, whereas non-migratory animals that must frequently move quickly to escape predators (such as frog
s) are likely to have costly but very fast locomotion. The study of animal locomotion is typically considered to be a sub-field of biomechanics
.
Locomotion requires energy
to overcome friction
, drag
, inertia
, and gravity, though in many circumstances some of these factors are negligible. In terrestrial
environments gravity must be overcome, though the drag of air is much less of an issue. In aqueous environments however, friction (or drag) becomes the major challenge, with gravity being less of a concern. Although animals with natural buoyancy
need not expend much energy maintaining vertical position, some will naturally sink and must expend energy to remain afloat. Drag may also present a problem in flight
, and the aerodynamically efficient body shapes of bird
s highlight this point. Flight presents a different problem from movement in water however, as there is no way for a living organism to have lower density
than air. Limbless organisms moving on land must often contend with surface friction, but do not usually need to expend significant energy to counteract gravity.
Newton's third law of motion
is widely used in the study of animal locomotion: if at rest, to move forwards an animal must push something backwards. Terrestrial animals must push the solid ground, swimming and flying animals must push against a fluid
or gas
(either water
or air). The effect of forces during locomotion on the design of the skeletal system is also important, as is the interaction between locomotion and muscle physiology, in determining how the structures and effectors of locomotion enable or limit animal movement.
s and otter
s, move through more than one type of fluid. In some cases locomotion is facilitated by the substrate on which they move. Forms of locomotion include:
encountered in water is much higher than that of air, which is almost negligible at low speeds. Body shape is therefore important for efficient movement, which is essential for basic functions like catching prey
. A fusiform, torpedo
-like body form is seen in many marine animals, though the mechanisms they employ for movement are diverse. Movement of the body may be from side to side, as in shark
s and many fish
es, or up and down, as in marine mammal
s. Other animals, such as those from the class Cephalopoda, use jet-propulsion, taking in water then squirting it back out in an explosive burst. Others may rely predominantly on their limbs, much as humans do when swimming. Though life on land originated from the seas, terrestrial animals have returned to an aquatic lifestyle on several occasions, such as the fully aquatic cetaceans, now far removed from their terrestrial ancestors.
to ascend and remain airborne. Wing shape is crucial in achieving this, generating a pressure gradient that results in an upward force on the animal' body. The same principle applies to airplanes, the wings of which are also airfoil
s. Unlike aircraft however, flying animals must be very light to achieve flight, the largest living flying animals being birds of around 20 kilograms. Other structural modifications of flying animals include reduced and redistributed body weight, fusiform shape and powerful flight muscles.
Rather than fly, some animals simply reduce their rate of falling by gliding
. Flight has independently
evolved at least four times, in the insect
s, pterosaur
s, bird
s, and bat
s. Gliding has evolved on many more occasions.
, and crawling or slithering. Here friction and buoyancy are no longer an issue, but a strong skeletal and muscular framework are required in most terrestrial animals for structural support. Each step also requires much energy to overcome inertia
, and animals can store elastic potential energy in their tendon
s to help overcome this. Balance
is also required for movement on land. Human infant
s learn to crawl first before they are able to stand on two feet, which requires good coordination as well as physical development. Humans are bipedal animals, standing on two feet and keeping one on the ground at all times while walking
. When running
, only one foot is on the ground at any one time at most, and both leave the ground briefly. At higher speeds momentum
helps keep the body upright, so more energy can be used in movement. The number of legs an animal has varies greatly, resulting in differences in locomotion. Many familiar mammals have four legs; insects have six, while arachnid
s have eight. Centipedes and millipedes have many sets of legs that move in metachronal rhythm
. Some have none at all, relying on other modes of locomotion.
Other animals move in terrestrial habitats without the aid of legs. Earthworm
s crawl by a peristalsis
, the same rhythmic contractions that propel food through the digestive tract. Snakes move using several different modes of locomotion, depending upon substrate type and desired speed. Some animals even roll, though typically not as a primary means of locomotion.
Some animals are specialized for moving on non-horizontal surfaces. One common habitat for such climbing animals
is in trees, for example the gibbon
is specialized for arboreal movement , traveling rapidly by brachiation
. Another case is animals like the snow leopard
living on steep rock faces such as are found in mountain
s. Some light animals are able to climb up smooth sheer surfaces or hang upside down by adhesion
. Many insects can do this, though much larger animals such as gecko
s can also perform similar feats.
of water. Animals that move in such a way include the water strider
. Water striders have legs that are hydrophobic, preventing them from interfering with the structure of water. Another form of locomotion (in which the surface layer is broken) is used by the Basilisk lizard.
is a hole or tunnel dug into the ground by an animal to create a space suitable for habitation, temporary refuge, or as a byproduct of locomotion. In loose solids such a sand some animals, such as the golden mole
, marsupial mole
, and the pink fairy armadillo
, are able to move more rapidly, 'swimming' through the loose substrate. Burrowing animals include moles
, ground squirrels, naked mole rat
s, tilefish
, mole cricket
s, and earthworms.
The most common metric of energy use during locomotion is net cost of transport, defined as the calories needed above baseline metabolism to move a given distance, per unit body mass. For aerobic locomotion, most animals have a nearly constant cost of transport - moving a given distance requires the same caloric expenditure, regardless of speed. This constancy is usually accomplished by changes in gait
. The net cost of transport of swimming is lowest, followed by flight, with terrestrial limbed locomotion being the most expensive per unit distance. However, because of the speeds involved, flight requires the most energy per unit time. This does not mean that an animal that normally moves by running would be a more efficient swimmer, however; these comparisons assume an animal is specialized for that form of motion. Another consideration here is body mass—heavier animals, though using more total energy, require less energy per unit mass to move. Physiologists
generally measure energy use by the amount of oxygen
consumed, or the amount of carbon dioxide
produced, in an animal's respiration
.
All of the methods can be combined. For example, studies frequently combine EMG and kinematics to determine "motor pattern", the series of electrical and kinematic events which produce a given movement.
Running
Running is a means of terrestrial locomotion allowing humans and other animals to move rapidly on foot. It is simply defined in athletics terms as a gait in which at regular points during the running cycle both feet are off the ground...
, swimming
Aquatic locomotion
Swimming is biologically propelled motion through a liquid medium. Swimming has evolved a number of times in a range of organisms ranging from arthropods to fish to molluscs.-Evolution of swimming:...
, jumping
Jumping
Jumping or leaping is a form of locomotion or movement in which an organism or non-living mechanical system propels itself through the air along a ballistic trajectory...
and flying
Flying and gliding animals
A number of animals have evolved aerial locomotion, either by powered flight or by gliding. Flying and gliding animals have evolved separately many times, without any single ancestor. Flight has evolved at least four times, in the insects, pterosaurs, birds, and bats. Gliding has evolved on many...
. Animals move for a variety of reasons, such as to find food, a mate, or a suitable microhabitat, and to escape predators. For many animals the ability to move is essential to survival and, as a result, selective pressures have shaped the locomotion methods and mechanisms employed by moving organisms. For example, migratory animals that travel vast distances (such as the Arctic Tern
Arctic Tern
The Arctic Tern is a seabird of the tern family Sternidae. This bird has a circumpolar breeding distribution covering the Arctic and sub-Arctic regions of Europe, Asia, and North America...
) typically have a locomotion mechanism that costs very little energy per unit distance, whereas non-migratory animals that must frequently move quickly to escape predators (such as frog
Frog
Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...
s) are likely to have costly but very fast locomotion. The study of animal locomotion is typically considered to be a sub-field of biomechanics
Biomechanics
Biomechanics is the application of mechanical principles to biological systems, such as humans, animals, plants, organs, and cells. Perhaps one of the best definitions was provided by Herbert Hatze in 1974: "Biomechanics is the study of the structure and function of biological systems by means of...
.
Locomotion requires energy
Energy
In physics, energy is an indirectly observed quantity. It is often understood as the ability a physical system has to do work on other physical systems...
to overcome friction
Friction
Friction is the force resisting the relative motion of solid surfaces, fluid layers, and/or material elements sliding against each other. There are several types of friction:...
, drag
Drag (physics)
In fluid dynamics, drag refers to forces which act on a solid object in the direction of the relative fluid flow velocity...
, inertia
Inertia
Inertia is the resistance of any physical object to a change in its state of motion or rest, or the tendency of an object to resist any change in its motion. It is proportional to an object's mass. The principle of inertia is one of the fundamental principles of classical physics which are used to...
, and gravity, though in many circumstances some of these factors are negligible. In terrestrial
Terrestrial planet
A terrestrial planet, telluric planet or rocky planet is a planet that is composed primarily of silicate rocks or metals. Within the Solar System, the terrestrial planets are the inner planets closest to the Sun...
environments gravity must be overcome, though the drag of air is much less of an issue. In aqueous environments however, friction (or drag) becomes the major challenge, with gravity being less of a concern. Although animals with natural buoyancy
Buoyancy
In physics, buoyancy is a force exerted by a fluid that opposes an object's weight. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus a column of fluid, or an object submerged in the fluid, experiences greater pressure at the bottom of the...
need not expend much energy maintaining vertical position, some will naturally sink and must expend energy to remain afloat. Drag may also present a problem in flight
Flight
Flight is the process by which an object moves either through an atmosphere or beyond it by generating lift or propulsive thrust, or aerostatically using buoyancy, or by simple ballistic movement....
, and the aerodynamically efficient body shapes of bird
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...
s highlight this point. Flight presents a different problem from movement in water however, as there is no way for a living organism to have lower density
Density
The mass density or density of a material is defined as its mass per unit volume. The symbol most often used for density is ρ . In some cases , density is also defined as its weight per unit volume; although, this quantity is more properly called specific weight...
than air. Limbless organisms moving on land must often contend with surface friction, but do not usually need to expend significant energy to counteract gravity.
Newton's third law of motion
Newton's laws of motion
Newton's laws of motion are three physical laws that form the basis for classical mechanics. They describe the relationship between the forces acting on a body and its motion due to those forces...
is widely used in the study of animal locomotion: if at rest, to move forwards an animal must push something backwards. Terrestrial animals must push the solid ground, swimming and flying animals must push against a fluid
Fluid
In physics, a fluid is a substance that continually deforms under an applied shear stress. Fluids are a subset of the phases of matter and include liquids, gases, plasmas and, to some extent, plastic solids....
or gas
Gas
Gas is one of the three classical states of matter . Near absolute zero, a substance exists as a solid. As heat is added to this substance it melts into a liquid at its melting point , boils into a gas at its boiling point, and if heated high enough would enter a plasma state in which the electrons...
(either water
Water
Water is a chemical substance with the chemical formula H2O. A water molecule contains one oxygen and two hydrogen atoms connected by covalent bonds. Water is a liquid at ambient conditions, but it often co-exists on Earth with its solid state, ice, and gaseous state . Water also exists in a...
or air). The effect of forces during locomotion on the design of the skeletal system is also important, as is the interaction between locomotion and muscle physiology, in determining how the structures and effectors of locomotion enable or limit animal movement.
Introduction
Animals move through a variety of fluids, such as water, air and mud. Some, for example sealPinniped
Pinnipeds or fin-footed mammals are a widely distributed and diverse group of semiaquatic marine mammals comprising the families Odobenidae , Otariidae , and Phocidae .-Overview: Pinnipeds are typically sleek-bodied and barrel-shaped...
s and otter
Otter
The Otters are twelve species of semi-aquatic mammals which feed on fish and shellfish, and also other invertebrates, amphibians, birds and small mammals....
s, move through more than one type of fluid. In some cases locomotion is facilitated by the substrate on which they move. Forms of locomotion include:
Swimming
In the water staying afloat is possible through buoyancy. Provided an aquatic animal's body is no denser than its aqueous environment, it should be able to stay afloat well enough. Though this means little energy need be expended maintaining vertical position, it makes movement in the horizontal plane much more difficult. The dragDrag (physics)
In fluid dynamics, drag refers to forces which act on a solid object in the direction of the relative fluid flow velocity...
encountered in water is much higher than that of air, which is almost negligible at low speeds. Body shape is therefore important for efficient movement, which is essential for basic functions like catching prey
Predation
In ecology, predation describes a biological interaction where a predator feeds on its prey . Predators may or may not kill their prey prior to feeding on them, but the act of predation always results in the death of its prey and the eventual absorption of the prey's tissue through consumption...
. A fusiform, torpedo
Torpedo
The modern torpedo is a self-propelled missile weapon with an explosive warhead, launched above or below the water surface, propelled underwater towards a target, and designed to detonate either on contact with it or in proximity to it.The term torpedo was originally employed for...
-like body form is seen in many marine animals, though the mechanisms they employ for movement are diverse. Movement of the body may be from side to side, as in shark
Shark
Sharks are a type of fish with a full cartilaginous skeleton and a highly streamlined body. The earliest known sharks date from more than 420 million years ago....
s and many fish
Fish
Fish are a paraphyletic group of organisms that consist of all gill-bearing aquatic vertebrate animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish, as well as various extinct related groups...
es, or up and down, as in marine mammal
Marine mammal
Marine mammals, which include seals, whales, dolphins, and walruses, form a diverse group of 128 species that rely on the ocean for their existence. They do not represent a distinct biological grouping, but rather are unified by their reliance on the marine environment for feeding. The level of...
s. Other animals, such as those from the class Cephalopoda, use jet-propulsion, taking in water then squirting it back out in an explosive burst. Others may rely predominantly on their limbs, much as humans do when swimming. Though life on land originated from the seas, terrestrial animals have returned to an aquatic lifestyle on several occasions, such as the fully aquatic cetaceans, now far removed from their terrestrial ancestors.
Flight
Gravity is a major problem for flight through the air. Because it is impossible for any organism to approach the density of air, flying animals must generate enough liftLift (force)
A fluid flowing past the surface of a body exerts a surface force on it. Lift is the component of this force that is perpendicular to the oncoming flow direction. It contrasts with the drag force, which is the component of the surface force parallel to the flow direction...
to ascend and remain airborne. Wing shape is crucial in achieving this, generating a pressure gradient that results in an upward force on the animal' body. The same principle applies to airplanes, the wings of which are also airfoil
Airfoil
An airfoil or aerofoil is the shape of a wing or blade or sail as seen in cross-section....
s. Unlike aircraft however, flying animals must be very light to achieve flight, the largest living flying animals being birds of around 20 kilograms. Other structural modifications of flying animals include reduced and redistributed body weight, fusiform shape and powerful flight muscles.
Rather than fly, some animals simply reduce their rate of falling by gliding
Gliding (flight)
Gliding flight is heavier-than-air flight without the use of thrust. It is employed by gliding animals and by aircraft such as gliders. The most common human application of gliding flight is in sport and recreation using aircraft designed for this purpose...
. Flight has independently
Convergent evolution
Convergent evolution describes the acquisition of the same biological trait in unrelated lineages.The wing is a classic example of convergent evolution in action. Although their last common ancestor did not have wings, both birds and bats do, and are capable of powered flight. The wings are...
evolved at least four times, in the insect
Insect flight
Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how...
s, pterosaur
Pterosaur
Pterosaurs were flying reptiles of the clade or order Pterosauria. They existed from the late Triassic to the end of the Cretaceous Period . Pterosaurs are the earliest vertebrates known to have evolved powered flight...
s, bird
Bird flight
Flight is the main mode of locomotion used by most of the world's bird species. Flight assists birds while feeding, breeding and avoiding predators....
s, and bat
Bat
Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...
s. Gliding has evolved on many more occasions.
Terrestrial
Forms of locomotion on land include walking, running, hopping or jumpingJumping
Jumping or leaping is a form of locomotion or movement in which an organism or non-living mechanical system propels itself through the air along a ballistic trajectory...
, and crawling or slithering. Here friction and buoyancy are no longer an issue, but a strong skeletal and muscular framework are required in most terrestrial animals for structural support. Each step also requires much energy to overcome inertia
Inertia
Inertia is the resistance of any physical object to a change in its state of motion or rest, or the tendency of an object to resist any change in its motion. It is proportional to an object's mass. The principle of inertia is one of the fundamental principles of classical physics which are used to...
, and animals can store elastic potential energy in their tendon
Tendon
A tendon is a tough band of fibrous connective tissue that usually connects muscle to bone and is capable of withstanding tension. Tendons are similar to ligaments and fasciae as they are all made of collagen except that ligaments join one bone to another bone, and fasciae connect muscles to other...
s to help overcome this. Balance
Equilibrioception
Equilibrioception or sense of balance is one of the physiological senses. It helps prevent humans and animals from falling over when walking or standing still. Balance is the result of a number of body systems working together: the eyes , ears and the body's sense of where it is in space ideally...
is also required for movement on land. Human infant
Infant
A newborn or baby is the very young offspring of a human or other mammal. A newborn is an infant who is within hours, days, or up to a few weeks from birth. In medical contexts, newborn or neonate refers to an infant in the first 28 days after birth...
s learn to crawl first before they are able to stand on two feet, which requires good coordination as well as physical development. Humans are bipedal animals, standing on two feet and keeping one on the ground at all times while walking
Walking
Walking is one of the main gaits of locomotion among legged animals, and is typically slower than running and other gaits. Walking is defined by an 'inverted pendulum' gait in which the body vaults over the stiff limb or limbs with each step...
. When running
Running
Running is a means of terrestrial locomotion allowing humans and other animals to move rapidly on foot. It is simply defined in athletics terms as a gait in which at regular points during the running cycle both feet are off the ground...
, only one foot is on the ground at any one time at most, and both leave the ground briefly. At higher speeds momentum
Momentum
In classical mechanics, linear momentum or translational momentum is the product of the mass and velocity of an object...
helps keep the body upright, so more energy can be used in movement. The number of legs an animal has varies greatly, resulting in differences in locomotion. Many familiar mammals have four legs; insects have six, while arachnid
Arachnid
Arachnids are a class of joint-legged invertebrate animals in the subphylum Chelicerata. All arachnids have eight legs, although in some species the front pair may convert to a sensory function. The term is derived from the Greek words , meaning "spider".Almost all extant arachnids are terrestrial...
s have eight. Centipedes and millipedes have many sets of legs that move in metachronal rhythm
Metachronal rhythm
A metachronal rhythm or metachronal wave refers to wavy movements produced by the sequential action of structures such as cilia, segments of worms or legs. These movements produce the appearance of a travelling wave. A Mexican wave is a large scale example of a metachronal wave...
. Some have none at all, relying on other modes of locomotion.
Other animals move in terrestrial habitats without the aid of legs. Earthworm
Earthworm
Earthworm is the common name for the largest members of Oligochaeta in the phylum Annelida. In classical systems they were placed in the order Opisthopora, on the basis of the male pores opening posterior to the female pores, even though the internal male segments are anterior to the female...
s crawl by a peristalsis
Peristalsis
Peristalsis is a radially symmetrical contraction and relaxation of muscles which propagates in a wave down the muscular tube, in an anterograde fashion. In humans, peristalsis is found in the contraction of smooth muscles to propel contents through the digestive tract. Earthworms use a similar...
, the same rhythmic contractions that propel food through the digestive tract. Snakes move using several different modes of locomotion, depending upon substrate type and desired speed. Some animals even roll, though typically not as a primary means of locomotion.
Some animals are specialized for moving on non-horizontal surfaces. One common habitat for such climbing animals
Arboreal locomotion
Arboreal locomotion is the locomotion of animals in trees. In every habitat in which trees are present, animals have evolved to move in them. Some animals may only scale trees occasionally, while others are exclusively arboreal. These habitats pose numerous mechanical challenges to animals...
is in trees, for example the gibbon
Gibbon
Gibbons are apes in the family Hylobatidae . The family is divided into four genera based on their diploid chromosome number: Hylobates , Hoolock , Nomascus , and Symphalangus . The extinct Bunopithecus sericus is a gibbon or gibbon-like ape which, until recently, was thought to be closely related...
is specialized for arboreal movement , traveling rapidly by brachiation
Brachiation
Brachiation is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms.- Brachiators :...
. Another case is animals like the snow leopard
Snow Leopard
The snow leopard is a moderately large cat native to the mountain ranges of South Asia and Central Asia...
living on steep rock faces such as are found in mountain
Mountain
Image:Himalaya_annotated.jpg|thumb|right|The Himalayan mountain range with Mount Everestrect 58 14 160 49 Chomo Lonzorect 200 28 335 52 Makalurect 378 24 566 45 Mount Everestrect 188 581 920 656 Tibetan Plateaurect 250 406 340 427 Rong River...
s. Some light animals are able to climb up smooth sheer surfaces or hang upside down by adhesion
Adhesion
Adhesion is any attraction process between dissimilar molecular species that can potentially bring them in close contact. By contrast, cohesion takes place between similar molecules....
. Many insects can do this, though much larger animals such as gecko
Gecko
Geckos are lizards belonging to the infraorder Gekkota, found in warm climates throughout the world. They range from 1.6 cm to 60 cm....
s can also perform similar feats.
On water
While animals like ducks can swim in water by floating, some small animals move across it without breaking through the surface. This surface locomotion takes advantage of the surface tensionSurface tension
Surface tension is a property of the surface of a liquid that allows it to resist an external force. It is revealed, for example, in floating of some objects on the surface of water, even though they are denser than water, and in the ability of some insects to run on the water surface...
of water. Animals that move in such a way include the water strider
Water strider
Gerridae is a family of true bugs in the order Hemiptera, commonly known as water striders, water bugs, magic bugs, pond skaters, skaters, skimmers, water scooters, water skaters, water skeeters, water skimmers, water skippers, water spiders, or Jesus bugs...
. Water striders have legs that are hydrophobic, preventing them from interfering with the structure of water. Another form of locomotion (in which the surface layer is broken) is used by the Basilisk lizard.
Through a solid medium
Some animals move through solids such as soil by burrowing using claws, teeth, or other methods. A burrowBurrow
A burrow is a hole or tunnel dug into the ground by an animal to create a space suitable for habitation, temporary refuge, or as a byproduct of locomotion. Burrows provide a form of shelter against predation and exposure to the elements, so the burrowing way of life is quite popular among the...
is a hole or tunnel dug into the ground by an animal to create a space suitable for habitation, temporary refuge, or as a byproduct of locomotion. In loose solids such a sand some animals, such as the golden mole
Golden mole
Golden moles are small, insectivorous burrowing mammals native to southern Africa. They form the family Chrysochloridae, and are taxonomically distinct from the true moles which they resemble due to convergence...
, marsupial mole
Marsupial mole
Marsupial moles is a family of marsupials of the order Notoryctemorphia, consisting of only two extant species:* Notoryctes typhlops * Notoryctes caurinus ...
, and the pink fairy armadillo
Pink Fairy Armadillo
The pink fairy armadillo or pichiciego is the smallest species of armadillo...
, are able to move more rapidly, 'swimming' through the loose substrate. Burrowing animals include moles
Mole (animal)
Moles are small cylindrical mammals adapted to a subterranean lifestyle. They have velvety fur; tiny or invisible ears and eyes; and short, powerful limbs with large paws oriented for digging. The term is especially and most properly used for the true moles, those of the Talpidae family in the...
, ground squirrels, naked mole rat
Naked Mole Rat
The naked mole rat , also known as the sand puppy or desert mole rat, is a burrowing rodent native to parts of East Africa and the only species currently classified in the genus Heterocephalus...
s, tilefish
Tilefish
Tilefishes, also known as blanquillo, are mostly small perciform marine fish comprising the family Malacanthidae.They are usually found in sandy areas, especially near coral reefs....
, mole cricket
Mole cricket
The mole crickets compose family Gryllotalpidae, of thick-bodied insects about long, with large beady eyes and shovel-like forelimbs highly developed for burrowing and swimming. They can also fly: the adult mole cricket may fly as far as during mating season, is active most of the year, and...
s, and earthworms.
Energetics
The energetics of locomotion involves the energy expenditure by animals in moving. Energy consumed in locomotion is not available for other efforts, so animals typically have evolved to use the minimum energy possible during movement. However, in the case of certain behaviors, such as locomotion to escape a predator, performance (such as speed or maneuverability) is more crucial, and such movements may be energetically expensive. Furthermore, animals may use energetically expensive methods of locomotion when environmental conditions (such as being within a tunnel) preclude other modes.The most common metric of energy use during locomotion is net cost of transport, defined as the calories needed above baseline metabolism to move a given distance, per unit body mass. For aerobic locomotion, most animals have a nearly constant cost of transport - moving a given distance requires the same caloric expenditure, regardless of speed. This constancy is usually accomplished by changes in gait
Gait
Gait is the pattern of movement of the limbs of animals, including humans, during locomotion over a solid substrate. Most animals use a variety of gaits, selecting gait based on speed, terrain, the need to maneuver, and energetic efficiency...
. The net cost of transport of swimming is lowest, followed by flight, with terrestrial limbed locomotion being the most expensive per unit distance. However, because of the speeds involved, flight requires the most energy per unit time. This does not mean that an animal that normally moves by running would be a more efficient swimmer, however; these comparisons assume an animal is specialized for that form of motion. Another consideration here is body mass—heavier animals, though using more total energy, require less energy per unit mass to move. Physiologists
Physiology
Physiology is the science of the function of living systems. This includes how organisms, organ systems, organs, cells, and bio-molecules carry out the chemical or physical functions that exist in a living system. The highest honor awarded in physiology is the Nobel Prize in Physiology or...
generally measure energy use by the amount of oxygen
Oxygen
Oxygen is the element with atomic number 8 and represented by the symbol O. Its name derives from the Greek roots ὀξύς and -γενής , because at the time of naming, it was mistakenly thought that all acids required oxygen in their composition...
consumed, or the amount of carbon dioxide
Carbon dioxide
Carbon dioxide is a naturally occurring chemical compound composed of two oxygen atoms covalently bonded to a single carbon atom...
produced, in an animal's respiration
Respiration (physiology)
'In physiology, respiration is defined as the transport of oxygen from the outside air to the cells within tissues, and the transport of carbon dioxide in the opposite direction...
.
Methods of study
A variety of methods and equipment are used to study animal locomotion:- KinematicsKinematicsKinematics is the branch of classical mechanics that describes the motion of bodies and systems without consideration of the forces that cause the motion....
is the study of the motion of an entire animal or parts of its body. It is typically accomplished by placing visual markers at particular anatomical locations on the animal and then recording videoVideoVideo is the technology of electronically capturing, recording, processing, storing, transmitting, and reconstructing a sequence of still images representing scenes in motion.- History :...
of its movement. The video is often captured from multiple angles, with frame rates exceeding 2000 frames per second when capturing high speed movement. The location of each marker is determined for each video frame, and data from multiple views is integrated to give positions of each point through time. Computers are sometimes used to track the markers, although this task must often be performed manually. The kinematic data can be used to determine fundamental motion attributes such as velocity, acceleration, joint angles, and the sequencing and timing of kinematic events. These fundamental attributes can be used to quantify various higher level attributes, such as the physical abilities of the animal (e.g., its maximum running speed, how steep a slope it can climb), neural control of locomotion, gait, and responses to environmental variation. These, in turn, can aid in formulation of hypotheses about the animal or locomotion in general.
- Force platesForce platformForce platforms or force plates are measuring instruments that measure the ground reaction forces generated by a body standing on or moving across them, to quantify balance, gait and other parameters of biomechanics. Most common areas of application are medicine and sports.The simplest force...
are platforms, usually part of a trackway, that can be used to measure the magnitude and direction of forces of an animal's step. When used with kinematics and a sufficiently detailed model of anatomy, inverse dynamics solutions can determine the forces not just at the contact with the ground, but at each joint in the limb.
- ElectromyographyElectromyographyElectromyography is a technique for evaluating and recording the electrical activity produced by skeletal muscles. EMG is performed using an instrument called an electromyograph, to produce a record called an electromyogram. An electromyograph detects the electrical potential generated by muscle...
(EMG) is a method of detecting the electrical activity that occurs when muscles are activated, thus determining which muscles are used when in a given movement. This can be accomplished either by surface electrodes (usually in large animals) or implanted electrodes (often wires thinner than a human hair). Furthermore, the intensity of electrical activity can correlate to the level of muscle activity, with greater activity implying (though not definitively showing) greater force.
- SonomicrometrySonomicrometrySonomicrometry is a technique of measuring the distance between piezoelectric crystals based on the speed of acoustic signals through the medium they are embedded in. Typically, the crystals will be coated with an epoxy 'lens' and placed into the material facing each other...
employs a pair of piezoelectric crystals implanted in a muscle or tendon to continuously measure the length of a muscle or tendon. This is useful because surface kinematics may be inaccurate due to skin movement. Similarly, if an elastic tendon is in series with the muscle, the muscle length may not be accurately reflected by the joint angle.
- Tendon force buckles measure the force produced by a single muscle by measuring the strain of a tendon. After the experiment, the tendon's elastic modulus is determined and used to compute the exact force produced by the muscle. However, this can only be used on muscles with long tendons.
- Particle image velocimetryParticle image velocimetryParticle image velocimetry is an optical method of flow visualization used in education and research. It is used to obtain instantaneous velocity measurements and related properties in fluids...
is used in aquatic and aerial systems to measure the flow of fluid around and past a moving aquatic organism, allowing fluid dynamics calculations to determine pressure gradients, speeds, etc.
- FluoroscopyFluoroscopyFluoroscopy is an imaging technique commonly used by physicians to obtain real-time moving images of the internal structures of a patient through the use of a fluoroscope. In its simplest form, a fluoroscope consists of an X-ray source and fluorescent screen between which a patient is placed...
allows real-time X-ray video, for precise kinematics of moving bones. Markers which are opaque to X-rays can allow simultaneous tracking of muscle length.
All of the methods can be combined. For example, studies frequently combine EMG and kinematics to determine "motor pattern", the series of electrical and kinematic events which produce a given movement.
See also
- FeatherFeatherFeathers are one of the epidermal growths that form the distinctive outer covering, or plumage, on birds and some non-avian theropod dinosaurs. They are considered the most complex integumentary structures found in vertebrates, and indeed a premier example of a complex evolutionary novelty. They...
- JointJointA joint is the location at which two or more bones make contact. They are constructed to allow movement and provide mechanical support, and are classified structurally and functionally.-Classification:...
- Kinesis (biology)
- Role of skin in locomotionRole of Skin in LocomotionRole of skin in locomotion describes how the integumentary system is involved in locomotion. Typically the integumentary system can be thought of as skin, however the integumentary system also includes the segmented exoskeleton in arthropods and feathers of birds. The primary role of the...
- TaxisTaxisA taxis is an innate behavioral response by an organism to a directional stimulus or gradient of stimulus intensity. A taxis differs from a tropism in that the organism has motility and demonstrates guided movement towards or away from the stimulus source ...
Further reading
- McNeill Alexander, Robert. (2003) Principles of Animal Locomotion. Princeton University Press, Princeton, N.J. ISBN 0691086788