Animal locomotion on the surface layer of water
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Animal locomotion on the surface layer of water is the study of
animal locomotion Animal locomotion, in ethology, is any of a variety of methods that animals use to move from one place to another. Some modes of locomotion are (initially) self-propelled, e.g., running, swimming, jumping, flying, hopping, soaring and gliding. Th ...
in the case of small
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s that live on the
surface A surface, as the term is most generally used, is the outermost or uppermost layer of a physical object or space. It is the portion or region of the object that can first be perceived by an observer using the senses of sight and touch, and is ...
layer of
water Water (chemical formula ) is an Inorganic compound, inorganic, transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance, which is the main constituent of Earth's hydrosphere and the fluids of all known living ...
, relying on surface tension to stay afloat. There are two types of animal locomotion on water, determined by the ratio of the animal's
weight In science and engineering, the weight of an object is the force acting on the object due to gravity. Some standard textbooks define weight as a vector quantity, the gravitational force acting on the object. Others define weight as a scalar qua ...
to the water's surface tension: those whose weight is supported by the surface tension at rest, and can therefore easily remain on the water's surface without much exertion, and those whose weight is not supported by the water's surface tension at rest, and must therefore exert additional motion in a direction parallel to the water's surface in order to remain above it. A creature such as the basilisk lizard, often dubbed the 'Jesus lizard', has a weight which is larger than the surface tension can support, and is widely known for running across the surface of water. Another example, the
western grebe The western grebe (''Aechmophorus occidentalis'') is a species in the grebe family of water birds. Folk names include "dabchick", "swan grebe" and "swan-necked grebe". Western grebe fossils from the Late Pleistocene of southwest North America ...
, performs a mating ritual that includes running across the surface of water. Surface living animals such as the
water strider The Gerridae are a family of insects in the order Hemiptera, commonly known as water striders, water skeeters, water scooters, water bugs, pond skaters, water skippers, or water skimmers. Consistent with the classification of the Gerridae as tr ...
typically have
hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, t ...
feet covered in small hairs that prevent the feet from breaking the surface and becoming wet. Another insect known to walk on the water surface is the ant species '' Polyrhachis sokolova''. The pygmy gecko ('' Coleodactylus amazonicus''), due to its small size and hydrophobic skin, is also able to walk on the water surface. According to
biophysicist Biophysics is an interdisciplinary science that applies approaches and methods traditionally used in physics to study biological phenomena. Biophysics covers all scales of biological organization, from molecular to organismic and populations. ...
David L. Hu, there are at least 342 species of water striders. As striders increase in size, their legs become proportionately longer, with '' Gigantometra gigas'' having a length of over 20 cm requiring a surface tension force of about 40 millinewtons. Water striders generate thrust by shedding
vortices In fluid dynamics, a vortex ( : vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in th ...
in the water: a series of "U"-shaped vortex filaments is created during the power stroke. The two free ends of the "U" are attached to the water surface. These vortices transfer enough (backward) momentum to the water to propel the animal forwards (note that some momentum is transferred by
capillary wave A capillary wave is a wave traveling along the phase boundary of a fluid, whose dynamics and phase velocity are dominated by the effects of surface tension. Capillary waves are common in nature, and are often referred to as ripples. The wav ...
s; see Denny's paradox for a more detailed discussion.)


Meniscus climbing

To pass from the water surface to land, a water-walking insect must contend with the slope of the
meniscus Meniscus may refer to: *Meniscus (anatomy), crescent-shaped fibrocartilaginous structure that partly divides a joint cavity *Meniscus (liquid) The meniscus (plural: ''menisci'', from the Greek for "crescent") is the curve in the upper surface ...
at the water's edge. Many such insects are unable to climb this meniscus using their usual propulsion mechanism. David Hu and coworker John W. M. Bush have shown that such insects climb meniscuses by assuming a fixed body posture. This deforms the water surface and generates capillary forces that propels the insect up the slope without moving its appendages. Hu and Bush conclude that meniscus climbing is an unusual means of propulsion in that the insect propels itself in a quasi-static configuration, without moving its appendages. Biolocomotion is generally characterized by the transfer of muscular strain energy to the kinetic and
gravitational potential energy Gravitational energy or gravitational potential energy is the potential energy a massive object has in relation to another massive object due to gravity. It is the potential energy associated with the gravitational field, which is released (conver ...
of the creature, and the kinetic energy of the suspending fluid. In contrast, meniscus climbing has a different energy pathway: by deforming the free surface, the insect converts muscular strain to the surface energy that powers its ascent.


Marangoni propulsion

Many insects, including some terrestrial insects, can release a surfactant and propel themselves using the
Marangoni effect The Marangoni effect (also called the Gibbs–Marangoni effect) is the mass transfer along an interface between two phases due to a gradient of the surface tension. In the case of temperature dependence, this phenomenon may be called thermo-capill ...
. Hu and Bush report that ''
Microvelia ''Microvelia'' is a genus of aquatic bugs in the family Veliidae Veliidae is a family of gregarious predatory insects in the suborder Heteroptera. They are commonly known as riffle bugs, small water striders, or broad-shouldered water stri ...
'' can attain a peak speed of 17 cm/s, which is twice its peak walking speed, using Marangoni propulsion. Marangoni propulsion by a
wetting Wetting is the ability of a liquid to maintain contact with a solid surface, resulting from intermolecular interactions when the two are brought together. This happens in presence of a gaseous phase or another liquid phase not miscible with ...
arthropod is precisely analogous to a soap boat but the situation for insects such as water striders is more complex. Hu and Bush state that "for nonwetting arthropods, the transfer of chemical to kinetic energy is more subtle, as the Marangoni stress must be communicated across the creature’s complex surface layer".


Sailing

''
Velella ''Velella'' is a monospecific genus of hydrozoa in the Porpitidae family. Its only known species is ''Velella velella'', a cosmopolitan free-floating hydrozoan that lives on the surface of the open ocean. It is commonly known by the names sea raf ...
'', the by-the-wind sailor, is a
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that ...
n with no means of propulsion other than
sailing Sailing employs the wind—acting on sails, wingsails or kites—to propel a craft on the surface of the ''water'' (sailing ship, sailboat, raft, windsurfer, or kitesurfer), on ''ice'' (iceboat) or on ''land'' (land yacht) over a chosen cou ...
. A small rigid sail projects into the air and catches the wind. ''Velella'' sails always align along the direction of the wind where the sail may act as an
aerofoil An airfoil (American English) or aerofoil (British English) is the cross-sectional shape of an object whose motion through a gas is capable of generating significant lift, such as a wing, a sail, or the blades of propeller, rotor, or turbine. ...
, so that the animals tend to sail downwind at a small angle to the wind.


See also

* Robostrider *
Stone skipping Stone skipping and stone skimming are considered related but distinct activities: both refer to the art of throwing a flat stone across the water in such a way (usually sidearm) that it bounces off the surface. The objective of "skipping" is ...


References

* John W. M. Bush and David L. Hu
''WALKING ON WATER: Biolocomotion at the Interface''
Annual Review of Fluid Mechanics, vol. 38, p. 339-369, January 2006.


External links



(photos) {{locomotion Animal locomotion Water transport