Aquatic locomotion or swimming is biologically propelled motion through a liquid medium.

The simplest
propulsive systems are composed of cilia and flagella. Swimming has evolved a number of times in a
range of organisms including arthropods, fish, molluscs, amphibians, reptiles, birds, and mammals.

Contents

1 Evolution of swimming

2 Micro-organisms

2.1 Bacterial

2.2 Ciliates

2.3 Flagellates

2.4 Pseudopodia

3 Invertebrates

3.1 Jet propulsion

4 Fish

4.1 Body-caudal fin (BCF) propulsion

4.2 Median paired fin (MPF) propulsion

4.3 Hydrofoils

4.4 Drag powered swimming

5 Amphibians

6 Reptiles

7 Fin and flipper locomotion

8 Escape reactions

9 Efficiency

9.1 Minimizing drag

9.2 Buoyancy
9.3 Temperature

9.4 Submergence

10 Secondary evolution

11 Human swimming

12 See also

13 References

Evolution of swimming

Jellyfish in motion

Swimming evolved a number of times in unrelated lineages. Supposed jellyfish fossils occur in the
Ediacaran, but the first free-swimming animals appear in the Early to Middle Cambrian. These are mostly
related to the arthropods, and include the Anomalocaridids, which swam by means of lateral lobes in a
fashion reminiscent of today's cuttlefish. Cephalopods joined the ranks of the nekton in the late
Cambrian,[1] and chordates were probably swimming from the Early Cambrian.[2] Many terrestrial
animals retain some capacity to swim, however some have returned to the water and developed the
capacities for aquatic locomotion. Most apes (including humans), however, lost the swimming instinct.
[3]

In 2013 Pedro Renato Bender, a research fellow at the University of the Witwatersrand's Institute for
Human Evolution, proposed a theory to explain the loss of that instinct. Termed the Saci last common
ancestor hypothesis (after Saci, a Brazilian folklore character who cannot cross water barriers), it holds
that the loss of instinctive swimming ability in apes is best explained as a consequence of constraints
related to the adaptation to an arboreal life in the last common ancestor of apes.[4] Bender
hypothesized that the ancestral ape increasingly avoided deep-water bodies when the risks of being
exposed to water were clearly higher than the advantages of crossing them.[4] A decreasing contact
with water bodies then could have led to the disappearance of the doggy paddle instinct.[4]

Micro-organisms

Further information: Microswimmer and Protist locomotion

Microbial swimmers, sometimes called microswimmers, are microscopic entities that have the ability to
move in fluid or aquatic environment.[5] Natural microswimmers are found everywhere in the natural
world as biological microorganisms, such as bacteria, archaea, protists, sperm and microanimals.
Bacterial

Main article: Bacterial motility

Ciliates

Ciliates use small flagella called cilia to move through the water. One ciliate will generally have hundreds
to thousands of cilia that are densely packed together in arrays. During movement, an individual cilium
deforms using a high-friction power stroke followed by a low-friction recovery stroke. Since there are
multiple cilia packed together on an individual organism, they display collective behavior in a
metachronal rhythm. This means the deformation of one cilium is in phase with the deformation of its
neighbor, causing deformation waves that propagate along the surface of the organism. These
propagating waves of cilia are what allow the organism to use the cilia in a coordinated manner to
move. A typical example of a ciliated microorganism is the Paramecium, a one-celled, ciliated protozoan
covered by thousands of cilia. The cilia beating together allow the Paramecium to propel through the
water at speeds of 500 micrometers per second.[6]

The flagellum of a Gram-negative bacteria is rotated by a molecular motor at its base

Salmon spermatozoa for artificial propagation

Flagellates

Certain organisms such as bacteria and animal sperm have flagellum which have developed a way to
move in liquid environments. A rotary motor model shows that bacteria uses the protons of an
electrochemical gradient in order to move their flagella. Torque in the flagella of bacteria is created by
particles that conduct protons around the base of the flagellum. The direction of rotation of the flagella
in bacteria comes from the occupancy of the proton channels along the perimeter of the flagellar motor.
[7]

Movement of sperm is called sperm motility. The middle of the mammalian spermatozoon contains
mitochondria that power the movement of the flagellum of the sperm. The motor around the base
produces torque, just like in bacteria for movement through the aqueous environment.[8]

Pseudopodia
Movement using a pseudopod is accomplished through increases in pressure at one point on the cell
membrane. This pressure increase is the result of actin polymerization between the cortex and the
membrane. As the pressure increases the cell membrane is pushed outward creating the pseudopod.
When the pseudopod moves outward, the rest of the body is pulled forward by cortical tension. The
result is cell movement through the fluid medium. Furthermore, the direction of movement is
determined by chemotaxis. When chemoattraction occurs in a particular area of the cell membrane,
actin polymerization can begin and move the cell in that direction.[9] An excellent example of an
organism that utilizes pseudopods is Naegleria fowleri.[10]

A Simple Animation

Invertebrates

Shrimp paddle with special swimming legs (pleopods)

Daphnia swims by beating its antennae

Among the radiata, jellyfish and their kin, the main form of swimming is to flex their cup shaped bodies.
All jellyfish are free-swimming, although many of these spend most of their time swimming passively.
Passive swimming is akin to gliding; the organism floats, using currents where it can, and does not exert
any energy into controlling its position or motion. Active swimming, in contrast, involves the
expenditure of energy to travel to a desired location.

In bilateria, there are many methods of swimming. The arrow worms (chaetognatha) undulate their
finned bodies, not unlike fish. Nematodes swim by undulating their fin-less bodies. Some Arthropod
groups can swim - including many crustaceans. Most crustaceans, such as shrimp, will usually swim by
paddling with special swimming legs (pleopods). Swimming crabs swim with modified walking legs
(pereiopods). Daphnia, a crustacean, swims by beating its antennae instead.

There are also a number of forms of swimming molluscs. Many free-swimming sea slugs, such as sea
angels, flap fin-like structures. Some shelled molluscs, such as scallops can briefly swim by clapping their
two shells open and closed. The molluscs most evolved for swimming are the cephalopods. Violet sea-
snails exploit a buoyant foam raft stabilized by amphiphilic mucins to float at the sea surface.[11][12]
Among the Deuterostomia, there are a number of swimmers as well. Feather stars can swim by
undulating their many arms. Salps move by pumping waters through their gelatinous bodies. The
deuterostomes most evolved for swimming are found among the vertebrates, notably the fish.

Jet propulsion

Octopuses swim headfirst, with arms trailing behind

Jellyfish pulsate their bell for a type of jet locomotion

Scallops swim by clapping their two shells open and closed

Main article: Jet propulsion

Jet propulsion is a method of aquatic locomotion where animals fill a muscular cavity and squirt out
water to propel them in the opposite direction of the squirting water. Most organisms are equipped
with one of two designs for jet propulsion; they can draw water from the rear and expel it from the rear,
such as jellyfish, or draw water from front and expel it from the rear, such as salps. Filling up the cavity
causes an increase in both the mass and drag of the animal. Because of the expanse of the contracting
cavity, the animal's velocity fluctuates as it moves through the water, accelerating while expelling water
and decelerating while vacuuming water. Even though these fluctuations in drag and mass can be
ignored if the frequency of the jet-propulsion cycles is high enough, jet-propulsion is a relatively
inefficient method of aquatic locomotion.

All cephalopods can move by jet propulsion, but this is a very energy-consuming way to travel compared
to the tail propulsion used by fish.[13] The relative efficiency of jet propulsion decreases further as
animal size increases. Since the Paleozoic, as competition with fish produced an environment where
efficient motion was crucial to survival, jet propulsion has taken a back role, with fins and tentacles used
to maintain a steady velocity.[14] The stop-start motion provided by the jets, however, continues to be
useful for providing bursts of high speed - not least when capturing prey or avoiding predators.[14]
Indeed, it makes cephalopods the fastest marine invertebrates,[15]: Preface  and they can out accelerate
most fish.[16] Oxygenated water is taken into the mantle cavity to the gills and through muscular
contraction of this cavity, the spent water is expelled through the hyponome, created by a fold in the
mantle. Motion of the cephalopods is usually backward as water is forced out anteriorly through the
hyponome, but direction can be controlled somewhat by pointing it in different directions.[17] Most
cephalopods float (i.e. are neutrally buoyant), so do not need to swim to remain afloat.[13] Squid swim
more slowly than fish, but use more power to generate their speed. The loss in efficiency is due to the
amount of water the squid can accelerate out of its mantle cavity.[18]

Jellyfish use a one-way water cavity design which generates a phase of continuous cycles of jet-
propulsion followed by a rest phase. The Froude efficiency is about 0.09, which indicates a very costly
method of locomotion. The metabolic cost of transport for jellyfish is high when compared to a fish of
equal mass.

Other jet-propelled animals have similar problems in efficiency. Scallops, which use a similar design to
jellyfish, swim by quickly opening and closing their shells, which draws in water and expels it from all
sides. This locomotion is used as a means to escape predators such as starfish. Afterwards, the shell acts
as a hydrofoil to counteract the scallop's tendency to sink. The Froude efficiency is low for this type of
movement, about 0.3, which is why it's used as an emergency escape mechanism from predators.
However, the amount of work the scallop has to do is mitigated by the elastic hinge that connects the
two shells of the bivalve. Squids swim by drawing water into their mantle cavity and expelling it through
their siphon. The Froude efficiency of their jet-propulsion system is around 0.29, which is much lower
than a fish of the same mass.

Much of the work done by scallop muscles to close its shell is stored as elastic energy in abductin tissue,
which acts as a spring to open the shell. The elasticity causes the work done against the water to be low
because of the large openings the water has to enter and the small openings the water has to leave. The
inertial work of scallop jet-propulsion is also low. Because of the low inertial work, the energy savings
created by the elastic tissue is so small that it's negligible. Medusae can also use their elastic mesoglea
to enlarge their bell. Their mantle contains a layer of muscle sandwiched between elastic fibers. The
muscle fibers run around the bell circumferentially while the elastic fibers run through the muscle and
along the sides of the bell to prevent lengthening. After making a single contraction, the bell vibrates
passively at the resonant frequency to refill the bell. However, in contrast with scallops, the inertial work
is similar to the hydrodynamic work due to how medusas expel water - through a large opening at low
velocity. Because of this, the negative pressure created by the vibrating cavity is lower than the positive
pressure of the jet, meaning that inertial work of the mantle is small. Thus, jet-propulsion is shown as an
inefficient swimming technique.[18]

Fish
Open water fish, like this Atlantic bluefin tuna, are usually streamlined for straightline speed, with a
deeply forked tail and a smooth body shaped like a spindle tapered at both ends.

Many reef fish, like this queen angelfish, have a body flattened like a pancake, with pectoral and pelvic
fins that act with the flattened body to maximize manoeuvrability.

See also: Fish locomotion

Many fish swim through water by creating undulations with their bodies or oscillating their fins. The
undulations create components of forward thrust complemented by a rearward force, side forces which
are wasted portions of energy, and a normal force that is between the forward thrust and side force.
Different fish swim by undulating different parts of their bodies. Eel-shaped fish undulate their entire
body in rhythmic sequences. Streamlined fish, such as salmon, undulate the caudal portions of their
bodies. Some fish, such as sharks, use stiff, strong fins to create dynamic lift and propel themselves. It is
common for fish to use more than one form of propulsion, although they will display one dominant
mode of swimming [19] Gait changes have even been observed in juvenile reef fish of various sizes.
Depending on their needs, fish can rapidly alternate between synchronized fin beats and alternating fin
beats.[20]

According to Guinness World Records 2009, Hippocampus zosterae (the dwarf seahorse) is the slowest
moving fish, with a top speed of about 5 feet (150 cm) per hour.[21] They swim very poorly, rapidly
fluttering a dorsal fin and using pectoral fins (located behind their eyes) to steer. Seahorses have no
caudal fin.

Body-caudal fin (BCF) propulsion

Anguilliform: Anguilliform swimmers are typically slow swimmers. They undulate the majority of their
body and use their head as the fulcrum for the load they are moving. At any point during their
undulation, their body has an amplitude between 0.5-1.0 wavelengths. The amplitude that they move
their body through allows them to swim backwards. Anguilliform locomotion is usually seen in fish with
long, slender bodies like eels, lampreys, oarfish, and a number of catfish species.

Subcarangiform, Carangiform, Thunniform: These swimmers undulate the posterior half of their body
and are much faster than anguilliform swimmers. At any point while they are swimming, a wavelength
<1 can be seen in the undulation pattern of the body. Some Carangiform swimmers include nurse
sharks, bamboo sharks, and reef sharks. Thunniform swimmers are very fast and some common
Thunniform swimmers include tuna, white sharks, salmon, jacks, and mako sharks[citation needed].
Thunniform swimmers only undulate their high aspect ratio caudal fin, so they are usually very stiff to
push more water out of the way.
Ostraciiform: Ostraciiform swimmers oscillate their caudal region, making them relatively slow
swimmers. Boxfish, torpedo rays, and momyrs employ Ostraciiform locomotion. The cow fish uses
Osctraciiform locomotion to hover in the water column.[22]

Median paired fin (MPF) propulsion

Tetraodoniform, Balistiform, Diodontiform: These swimmers oscillate their median fins. They are
typically slow swimmers, and some notable examples include the oceanic sunfish (which has extremely
modified anal and dorsal fins), puffer fish, and triggerfish.

Rajiform, Amiiform, Gymnotiform: This locomotory mode is accomplished by undulation of the pectoral
and median fins. During their undulation pattern, a wavelength >1 can be seen in their fins. They are
typically slow to moderate swimmers, and some examples include rays, bowfin, and knife fishes. The
black ghost knife fish is a Gymnotiform swimmer that has a very long ventral ribbon fin. Thrust is
produced by passing waves down the ribbon fin while the body remains rigid. This also allows the ghost
knife fish to swim in reverse.

Labriform: Labriform swimmers are also slow swimmers. They oscillate their pectoral fins to create
thrust. Oscillating fins create thrust when a starting vortex is shed from the trailing edge of the fin. As
the foil departs from the starting vortex, the effect of that vortex diminishes, while the bound circulation
remains, producing lift. Labriform swimming can be viewed as continuously starting and stopping.
Wrasses and surf perch are common Labriform swimmers.[22]

Hydrofoils

The leopard shark angles its pectoral fins so they behave as hydrofoils to control the animal's pitch

Hydrofoils, or fins, are used to push against the water to create a normal force to provide thrust,
propelling the animal through water. Sea turtles and penguins beat their paired hydrofoils to create lift.
Some paired fins, such as pectoral fins on leopard sharks, can be angled at varying degrees to allow the
animal to rise, fall, or maintain its level in the water column. The reduction of fin surface area helps to
minimize drag, and therefore increase efficiency. Regardless of size of the animal, at any particular
speed, maximum possible lift is proportional to (wing area) x (speed)2. Dolphins and whales have large,
horizontal caudal hydrofoils, while many fish and sharks have vertical caudal hydrofoils. Porpoising (seen
in cetaceans, penguins, and pinnipeds) may save energy if they are moving fast. Since drag increases
with speed, the work required to swim unit distance is greater at higher speeds, but the work needed to
jump unit distance is independent of speed. Seals propel themselves through the water with their
caudal tail, while sea lions create thrust solely with their pectoral flippers.[19]