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A rete mirabile (
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
for "wonderful net"; plural retia mirabilia) is a complex of
arteries An artery (plural arteries) () is a blood vessel in humans and most animals that takes blood away from the heart to one or more parts of the body (tissues, lungs, brain etc.). Most arteries carry oxygenated blood; the two exceptions are the p ...
and
vein Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated ...
s lying very close to each other, found in some
vertebrate Vertebrates () comprise all animal taxon, taxa within the subphylum Vertebrata () (chordates with vertebral column, backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the ...
s, mainly
warm-blooded Warm-blooded is an informal term referring to animal species which can maintain a body temperature higher than their environment. In particular, homeothermic species maintain a stable body temperature by regulating metabolic processes. The onl ...
ones. The rete mirabile utilizes countercurrent blood flow within the net (blood flowing in opposite directions) to act as a countercurrent exchanger. It exchanges
heat In thermodynamics, heat is defined as the form of energy crossing the boundary of a thermodynamic system by virtue of a temperature difference across the boundary. A thermodynamic system does not ''contain'' heat. Nevertheless, the term is al ...
,
ion An ion () is an atom or molecule with a net electrical charge. The charge of an electron is considered to be negative by convention and this charge is equal and opposite to the charge of a proton, which is considered to be positive by conv ...
s, or
gas Gas is one of the four fundamental states of matter (the others being solid, liquid, and plasma). A pure gas may be made up of individual atoms (e.g. a noble gas like neon), elemental molecules made from one type of atom (e.g. oxygen), or ...
es between vessel walls so that the two bloodstreams within the rete maintain a gradient with respect to
temperature Temperature is a physical quantity that expresses quantitatively the perceptions of hotness and coldness. Temperature is measurement, measured with a thermometer. Thermometers are calibrated in various Conversion of units of temperature, temp ...
, or concentration of gases or
solutes In chemistry, a solution is a special type of homogeneous mixture composed of two or more substances. In such a mixture, a solute is a substance dissolved in another substance, known as a solvent. If the attractive forces between the solvent ...
. This term was coined by
Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be one of ...
.


Effectiveness

The effectiveness of retia is primarily determined by how readily the heat, ions, or gases can be exchanged. For a given length, they are most effective with respect to gases or heat, then small ions, and decreasingly so with respect to other substances. The retia can provide for extremely efficient exchanges. In
bluefin tuna Bluefin tuna is a common name used to refer to several species of tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae ( mackerel) family. The Thunnini comprise 15 species across five genera, ...
, for example, nearly all of the metabolic heat in the venous blood is transferred to the arterial blood, thus conserving muscle temperature; that heat exchange approaches 99% efficiency.


Birds

In birds with
webbed feet The webbed foot is a specialized limb with interdigital membranes (webbings) that aids in aquatic locomotion, present in a variety of tetrapod vertebrates. This adaptation is primarily found in semiaquatic species, and has convergently evolved ...
, retia mirabilia in the legs and feet transfer heat from the outgoing (hot) blood in the arteries to the incoming (cold) blood in the veins. The effect of this biological
heat exchanger A heat exchanger is a system used to transfer heat between a source and a working fluid. Heat exchangers are used in both cooling and heating processes. The fluids may be separated by a solid wall to prevent mixing or they may be in direct contac ...
is that the internal temperature of the feet is much closer to the ambient temperature, thus reducing heat loss.
Penguin Penguins ( order Sphenisciformes , family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain t ...
s also have them in the flippers and nasal passages. Seabirds distill seawater using countercurrent exchange in a so-called
salt gland The salt gland is an organ for excreting excess salts. It is found in the cartilaginous fishes subclass elasmobranchii (sharks, rays, and skates), seabirds, and some reptiles. Salt glands can be found in the rectum of sharks. Birds and reptile ...
with a rete mirabile. The gland secretes highly concentrated brine stored near the nostrils above the beak. The bird then "sneezes" the brine out. As freshwater is not usually available in their environments, some seabirds, such as
pelican Pelicans (genus ''Pelecanus'') are a genus of large water birds that make up the family Pelecanidae. They are characterized by a long beak and a large throat pouch used for catching prey and draining water from the scooped-up contents befor ...
s,
petrel Petrels are tube-nosed seabirds in the bird order Procellariiformes. Description The common name does not indicate relationship beyond that point, as "petrels" occur in three of the four families within that group (all except the albatross ...
s,
albatross Albatrosses, of the biological family Diomedeidae, are large seabirds related to the procellariids, storm petrels, and diving petrels in the order Procellariiformes (the tubenoses). They range widely in the Southern Ocean and the North Pac ...
es,
gull Gulls, or colloquially seagulls, are seabirds of the family Laridae in the suborder Lari. They are most closely related to the terns and skimmers and only distantly related to auks, and even more distantly to waders. Until the 21st century, ...
s and
tern Terns are seabirds in the family Laridae that have a worldwide distribution and are normally found near the sea, rivers, or wetlands. Terns are treated as a subgroup of the family Laridae which includes gulls and skimmers and consists of ...
s, possess this gland, which allows them to drink the salty water from their environments while they are hundreds of miles away from land.


Fish

Fish have evolved retia mirabilia multiple times to raise the temperature (
endotherm An endotherm (from Greek ἔνδον ''endon'' "within" and θέρμη ''thermē'' "heat") is an organism that maintains its body at a metabolically favorable temperature, largely by the use of heat released by its internal bodily functions inste ...
y) or the oxygen concentration of a body part above the ambient level. In many
fish Fish are aquatic, craniate, gill-bearing 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. Approximately 95% of li ...
, a rete mirabile helps fill the
swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their current water depth wi ...
with
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as well ...
, increasing the fish's
buoyancy Buoyancy (), or upthrust, is an upward force exerted by a fluid that opposes the weight of a partially or fully immersed object. In a column of fluid, pressure increases with depth as a result of the weight of the overlying fluid. Thus the pr ...
. The rete mirable is an essential part of the system that pumps dissolved oxygen from a low
partial pressure In a mixture of gases, each constituent gas has a partial pressure which is the notional pressure of that constituent gas as if it alone occupied the entire volume of the original mixture at the same temperature. The total pressure of an ideal gas ...
() of 0.2 atmospheres into a gas filled bladder that is at a pressure of hundreds of atmospheres. A rete mirabile called the choroid rete mirabile is found in most living
teleost Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Teleo ...
s and raises the of the retina. The higher supply of oxygen allows the teleost retina to be thick and have few blood vessels thereby increasing its sensitivity to light. In addition to raising the , the choroid rete has evolved to raise the temperature of the eye in some teleosts and
sharks Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimor ...
. A countercurrent exchange system is utilized between the venous and arterial capillaries. Lowering the pH levels in the venous capillaries causes oxygen to unbind from blood
hemoglobin Hemoglobin (haemoglobin BrE) (from the Greek word αἷμα, ''haîma'' 'blood' + Latin ''globus'' 'ball, sphere' + ''-in'') (), abbreviated Hb or Hgb, is the iron-containing oxygen-transport metalloprotein present in red blood cells (erythrocyt ...
because of the root effect. This causes an increase in venous blood oxygen partial pressure, allowing the oxygen to diffuse through the capillary membrane and into the arterial capillaries, where oxygen is still sequestered to hemoglobin. The cycle of diffusion continues until the partial pressure of oxygen in the arterial capillaries exceeds that in the swim bladder. At this point, the dissolved oxygen in the arterial capillaries diffuses into the swim bladder via the gas gland. The rete mirabile allows for an increase in muscle temperature in regions where this network of vein and arteries is found. The fish is able to thermoregulate certain areas of its body. Additionally, this increase in temperature leads to an increase in basal metabolic temperature. The fish is now able to split ATP at a higher rate and ultimately can swim faster. The
opah Opahs, also commonly known as moonfish, sunfish (not to be confused with Molidae), kingfish, redfin ocean pan are large, colorful, deep-bodied pelagic lampriform fishes comprising the small family Lampridae (also spelled Lamprididae). The fami ...
utilizes retia mirabilia to conserve heat, making it the newest addition to the list of regionally endothermic fish. Blood traveling through capillaries in the gills must carry cold blood due to their exposure to cold water, but retia mirabilia in the opah's
gills A gill () is a respiratory organ that many aquatic organisms use to extract dissolved oxygen from water and to excrete carbon dioxide. The gills of some species, such as hermit crabs, have adapted to allow respiration on land provided they are ...
are able to transfer heat from warm blood in arterioles coming from the heart that heats this colder blood in arterioles leaving the gills. The huge
pectoral muscles Pectoral muscles (colloquially referred to as "pecs") are the muscles that connect the front of the human chest with the bones of the upper arm and shoulder. This region contains four muscles that provide movements to the upper limbs or ribs. Pe ...
of the opah, which generate most of the body heat, are thus able to control the temperature of the rest of the body.


Mammals

In
mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...
s, an elegant rete mirabile in the
efferent arteriole The efferent arterioles are blood vessels that are part of the urinary tract of organisms. Efferent (from Latin ex + ferre) means "outgoing", in this case meaning carrying blood out away from the glomerulus. The efferent arterioles form a conve ...
s of juxtamedullary glomeruli is important in maintaining the hypertonicity of the
renal medulla The renal medulla is the innermost part of the kidney. The renal medulla is split up into a number of sections, known as the renal pyramids. Blood enters into the kidney via the renal artery, which then splits up to form the segmental arteries whi ...
. It is the hypertonicity of this zone, resorbing water osmotically from the renal collecting ducts as they exit the
kidney The kidneys are two reddish-brown bean-shaped organs found in vertebrates. They are located on the left and right in the retroperitoneal space, and in adult humans are about in length. They receive blood from the paired renal arteries; blood ...
, that makes possible the excretion of a hypertonic
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellular m ...
and maximum conservation of body water. Vascular retia mirabilia are also found in the limbs of a range of mammals. These reduce the temperature in the extremities. Some of these probably function to prevent heat loss in cold conditions by reducing the temperature gradient between the limb and the environment. Others reduce the temperature of the
testes A testicle or testis (plural testes) is the male reproductive gland or gonad in all bilaterians, including humans. It is homologous to the female ovary. The functions of the testes are to produce both sperm and androgens, primarily testost ...
increasing their productivity. In the neck of the
dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the wolf. Also called the domestic dog, it is derived from the extinct Pleistocene wolf, and the modern wolf is the dog's nearest living relative. ...
, a rete mirabile protects the brain when the body overheats during hunting; the venous blood is cooled down by panting before entering the net. Retia mirabilia also occur frequently in mammals that burrow, dive or have arboreal lifestyles that involve clinging with the limbs for lengthy periods. In these cases, the rete mirabile may lower limb temperature and therefore the metabolic requirement for oxygen and nutrients in the tissues of the limb. This would be desirable when the blood supply to the limb is limited as a result of a "diving reflex" diverting blood away from the limbs during diving or burrowing or because of postural restrictions on blood supply in the case of clinging, arboreal animals. In the last case, slow-moving arboreal mammals such as sloths, lorises and arboreal anteaters possess retia of the highly developed type known as vascular bundles. The structure and function of these mammalian retia mirabilia are reviewed by O'Dea (1990). The ancient physician
Galen Aelius Galenus or Claudius Galenus ( el, Κλαύδιος Γαληνός; September 129 – c. AD 216), often Anglicized as Galen () or Galen of Pergamon, was a Greek physician, surgeon and philosopher in the Roman Empire. Considered to be one of ...
mistakenly thought that
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, an ...
s also have a rete mirabile in the neck, apparently based on dissection of
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated sh ...
and misidentifying the results with the human
carotid sinus In human anatomy, the carotid sinus is a dilated area at the base of the internal carotid artery just superior to the bifurcation of the internal carotid and external carotid at the level of the superior border of thyroid cartilage. The carotid s ...
, and ascribed important properties to it; it fell to Berengario da Carpi first, and then to
Vesalius Andreas Vesalius (Latinized from Andries van Wezel) () was a 16th-century anatomist, physician, and author of one of the most influential books on human anatomy, ''De Humani Corporis Fabrica Libri Septem'' (''On the fabric of the human body'' '' ...
to demonstrate the error.


See also

*
Pampiniform plexus The pampiniform plexus (from Latin ''pampinus'', a tendril, + ''forma'', form) is a venous plexus – a network of many small veins found in the human male spermatic cord, and the suspensory ligament of the ovary. In the male, it is formed by the ...
, a countercurrent heat-exchanging structure in the spermatic cord


References


External links


Fish gas bladder with an adjacent rete mirabile
{{DEFAULTSORT:Rete Mirabile Circulatory system Vertebrate anatomy Latin biological phrases