A rete mirabile (

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

for "wonderful net"; : retia mirabilia) is a complex of

arteries An artery () is a blood vessel in humans and most other animals that takes oxygenated blood away from the heart in the systemic circulation to one or more parts of the body. Exceptions that carry deoxygenated blood are the pulmonary arteries in ...

and

vein Veins () are blood vessels in the circulatory system of 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 those of the pulmonary and feta ...

s lying very close to each other, found in some

vertebrate Vertebrates () are animals with a vertebral column (backbone or spine), and a cranium, or skull. The vertebral column surrounds and protects the spinal cord, while the cranium protects the brain. The vertebrates make up the subphylum Vertebra ...

s, mainly

warm-blooded Warm-blooded is a term referring to animal species whose bodies maintain a temperature higher than that of their environment. In particular, homeothermic species (including birds and mammals) maintain a stable body temperature by regulating ...

ones. The rete mirabile utilizes countercurrent blood flow within the net (blood flowing in opposite directions) to act as a

countercurrent exchange Countercurrent exchange is a mechanism between two flowing bodies flowing in opposite directions to each other, in which there is a transfer of some property, usually heat or some chemical. The flowing bodies can be liquids, gases, or even solid ...

r. It exchanges

heat In thermodynamics, heat is energy in transfer between a thermodynamic system and its surroundings by such mechanisms as thermal conduction, electromagnetic radiation, and friction, which are microscopic in nature, involving sub-atomic, ato ...

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 convent ...

s, or

gas Gas is a state of matter that has neither a fixed volume nor a fixed shape and is a compressible fluid. A ''pure gas'' is made up of individual atoms (e.g. a noble gas like neon) or molecules of either a single type of atom ( elements such as ...

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 quantitatively expresses the attribute of hotness or coldness. Temperature is measurement, measured with a thermometer. It reflects the average kinetic energy of the vibrating and colliding atoms making ...

, or concentration of gases or

solutes In chemistry, a solution is defined by IUPAC as "A liquid or solid phase containing more than one substance, when for convenience one (or more) substance, which is called the solvent, is treated differently from the other substances, which are ...

. This term was coined by

Galen Aelius Galenus or Claudius Galenus (; September 129 – AD), often Anglicization, anglicized as Galen () or Galen of Pergamon, was a Ancient Rome, Roman and Greeks, Greek physician, surgeon, and Philosophy, philosopher. Considered to be one o ...

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 of the genus ''Thunnus''. {{Animal common name Commercial fish Thunnus Fish common names ...

, 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

bird Birds are a group of warm-blooded vertebrates constituting the class (biology), class Aves (), characterised by feathers, toothless beaked jaws, the Oviparity, laying of Eggshell, hard-shelled eggs, a high Metabolism, metabolic rate, a fou ...

s with webbed feet, 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 are a group of aquatic flightless birds from the family Spheniscidae () of the order Sphenisciformes (). They live almost exclusively in the Southern Hemisphere. Only one species, the Galápagos penguin, is equatorial, with a sm ...

s also have them in the flippers and nasal passages. Seabirds distill seawater using

in a so-called

salt gland The salt gland is an organ (anatomy), organ for excreting excess salt (chemistry), 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 r ...

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 before ...

petrel Petrels are tube-nosed seabirds in the phylogenetic order Procellariiformes. Description Petrels are a monophyletic group of marine seabirds, sharing a characteristic of a nostril arrangement that results in the name "tubenoses". Petrels enco ...

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 Paci ...

es,

gull Gulls, or colloquially seagulls, are seabirds of the subfamily Larinae. They are most closely related to terns and skimmers, distantly related to auks, and even more distantly related to waders. Until the 21st century, most gulls were placed ...

s and

tern Terns are seabirds in the family Laridae, subfamily Sterninae, that have a worldwide distribution and are normally found near the sea, rivers, or wetlands. Terns are treated in eleven genera in a subgroup of the family Laridae, which also ...

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 inst ...

y) or the oxygen concentration of a body part above the ambient level. In many

fish A fish (: fish or fishes) is an aquatic animal, aquatic, Anamniotes, anamniotic, gill-bearing vertebrate animal with swimming fish fin, fins and craniate, a hard skull, but lacking limb (anatomy), limbs with digit (anatomy), digits. Fish can ...

, a rete mirabile helps fill the

swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ (anatomy), organ in bony fish that functions to modulate buoyancy, and thus allowing the fish to stay at desired water depth without having to maintain lift ...

with

oxygen Oxygen is a chemical element; it has chemical symbol, symbol O and atomic number 8. It is a member of the chalcogen group (periodic table), group in the periodic table, a highly reactivity (chemistry), reactive nonmetal (chemistry), non ...

, increasing the fish's

buoyancy Buoyancy (), or upthrust, is the force exerted by a fluid opposing the weight of a partially or fully immersed object (which may be also be a parcel of fluid). In a column of fluid, pressure increases with depth as a result of the weight of t ...

. The rete mirabile 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 g ...

(

) 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 (; Ancient Greek, Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts (), is, by far, the largest group of ray-finned fishes (class Actinopterygii), with 96% of all neontology, extant species of f ...

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 cartilaginous fish characterized by a ribless endoskeleton, dermal denticles, five to seven gill slits on each side, and pectoral fins that are not fused to the head. Modern sharks are classified within the ...

. A

countercurrent exchange system Countercurrent exchange is a mechanism between two flowing bodies flowing in opposite directions to each other, in which there is a transfer of some property, usually heat or some chemical. The flowing bodies can be liquids, gases, or even solid ...

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, Hb or Hgb) is a protein containing iron that facilitates the transportation of oxygen in red blood cells. Almost all vertebrates contain hemoglobin, with the sole exception of the fish family Channichthyidae. Hemoglobin ...

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 The opah, also commonly known as moonfish, sunfish (not to be confused with Molidae), cowfish, kingfish, and redfin ocean pan are large, colorful, deep-bodied Pelagic zone, pelagic lampriform Fish, fishes comprising the genus ''Lampris'', of ...

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. P ...

of the opah, which generate most of the body heat, are thus able to control the temperature of the rest of the body.

Mammals

mammal A mammal () is a vertebrate animal of the Class (biology), class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three ...

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 conver ...

s of juxtamedullary glomeruli is important in maintaining the hypertonicity of the

renal medulla The renal medulla (Latin: ''medulla renis'' 'marrow of the kidney') 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 ...

. It is the hypertonicity of this zone, resorbing water osmotically from the renal collecting ducts as they exit the

kidney In humans, the kidneys are two reddish-brown bean-shaped blood-filtering organ (anatomy), organs that are a multilobar, multipapillary form of mammalian kidneys, usually without signs of external lobulation. They are located on the left and rig ...

, that makes possible the excretion of a hypertonic

urine Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...

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 ( testes) is the gonad in all male bilaterians, including humans, and is homologous to the ovary in females. Its primary functions are the production of sperm and the secretion of androgens, primarily testosterone. The ...

increasing their productivity. In the neck of the

dog The dog (''Canis familiaris'' or ''Canis lupus familiaris'') is a domesticated descendant of the gray wolf. Also called the domestic dog, it was selectively bred from a population of wolves during the Late Pleistocene by hunter-gatherers. ...

, 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 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

mistakenly thought that

human Humans (''Homo sapiens'') or modern humans are the most common and widespread species of primate, and the last surviving species of the genus ''Homo''. They are Hominidae, great apes characterized by their Prehistory of nakedness and clothing ...

s also have a rete mirabile in the neck, apparently based on dissection of

sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal 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 d ...

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 carot ...

, and ascribed important properties to it; it fell to Berengario da Carpi first, and then to

Vesalius Andries van Wezel (31 December 1514 – 15 October 1564), Latinization of names, latinized as Andreas Vesalius (), was an anatomist and physician who wrote ''De humani corporis fabrica, De Humani Corporis Fabrica Libri Septem'' (''On the fabric ...

to demonstrate the error.

References

External links