Ampullae of Lorenzini

The ampullae of Lorenzini (sing. ampulla) are special sensing organs called electroreceptors, forming a network of jelly-filled pores. They are mostly discussed as being found in cartilaginous fish (sharks, rays, and chimaeras); however, they are also reported to be found in Chondrostei such as reedfish[1] and sturgeon.[2] Lungfish have also been reported to have them.[1] Teleosts have re-evolved a different type of electroreceptors.[2] They were first described by Stefano Lorenzini in 1678.

Lorenzini pores on snout of tiger shark
Pores with ampullae of Lorenzini in snout of Tiger shark

These sensory organs help fish to sense electric fields in the water. Each ampulla consists of a jelly-filled canal opening to the surface by a pore in the skin and ending blindly in a cluster of small pockets full of special jelly. The ampullae are mostly clustered into groups inside the body, each cluster having ampullae connecting with different parts of the skin, but preserving a left-right symmetry. The canal lengths vary from animal to animal, but the distribution of the pores is generally specific to each species. The ampullae pores are plainly visible as dark spots in the skin. They provide fish with an additional sense capable of detecting electric and magnetic fields as well as temperature gradients.

Electroreceptors in a sharks head
Electroreceptors (ampullae of Lorenzini) and lateral line canals in the head of a shark
Ampullae of Lorenzini inner side
Inner view of Ampullae of Lorenzini

Electric field sensing ability

The ampullae detect electric fields in the water, or more precisely the potential difference between the voltage at the skin pore and the voltage at the base of the electroreceptor cells. A positive pore stimulus would decrease the rate of nerve activity coming from the electroreceptor cells, and a negative pore stimulus would increase the rate of nerve activity coming from the electroreceptor cells. Each ampulla contains a single layer of cells that contains electrically excitable receptor cells separated by supporting cells. The cells are connected by apical tight junctions so that no current leaks between the cells. The apical faces of the receptor cells have a small surface area with a high concentration of voltage dependent calcium channels and calcium activated potassium channels.[3] Because the canal wall has a very high resistance, all of the voltage difference between the pore of the canal and the ampulla is dropped across the receptor epithelium which is about 50 microns thick. Because the basal membranes of the receptor cells have a lower resistance, most of the voltage is dropped across the apical faces which are excitable and are poised at threshold. Inward calcium current across the receptor cells depolarizes the basal faces causing presynaptic calcium release and release of excitatory transmitter onto the afferent nerve fibers. One of the first descriptions of calcium activated potassium channels was based on studies of the ampulla of Lorenzini in the skate. Large conductance calcium activated potassium channels (BK channels) have recently been demonstrated in the ampulla by cloning.

Sharks may be more sensitive to electric fields than any other animal, with a threshold of sensitivity as low as 5 nV/cm. That is 5/1,000,000,000 of a volt measured in a centimeter-long ampulla. All living creatures produce an electrical field by muscle contractions, and a shark may pick up weak electrical stimuli from the muscle contractions of animals, particularly prey.[4] On the other hand, the electrochemical fields generated by paralyzed prey were sufficient to elicit a feeding attack from sharks and rays in experimental tanks; therefore muscle contractions are not necessary to attract the animals. Sharks and rays can locate prey buried in the sand, or DC electric dipoles that simulate the main feature of the electric field of a prey buried in the sand.

Any moving conductor, such as sea water, induces an electric field when a magnetic field such as the Earth's is present. The electric fields induced in oceanic currents by the Earth's magnetic field are of the same order of magnitude as the electric fields that sharks and rays are capable of sensing. This could mean that sharks and rays can orient to the electric fields of oceanic currents, and use other sources of electric fields in the ocean for local orientation. Additionally, the electric field they induce in their bodies when swimming in the magnetic field of the Earth may enable them to sense their magnetic heading.

Behavioral studies have also provided evidence that sharks can detect changes in the geomagnetic field. In one experiment, sandbar sharks and scalloped hammerhead sharks were conditioned to associate a food reward with an artificial magnetic field. When the food reward was removed, the sharks continued to show a marked difference in behavior when the magnetic field was turned on as compared to when it was off.[5]

Temperature sensing ability

Early in the 20th century, the purpose of the ampullae was not clearly understood, and electrophysiological experiments suggested a sensibility to temperature, mechanical pressure and possibly salinity. It was not until 1960 that the ampullae were clearly identified as specialized receptor organs for sensing electric fields.[6][7] The ampullae may also allow the shark to detect changes in water temperature. Each ampulla is a bundle of sensory cells containing multiple nerve fibres. These fibres are enclosed in a gel-filled tubule which has a direct opening to the surface through a pore. The gel is a glycoprotein based substance with the same resistivity as seawater, and it has electrical properties similar to a semiconductor.[8] This has been suggested as a mechanism by which temperature changes are transduced into an electrical signal that the shark may use to detect temperature gradients, although it is a subject of debate in scientific literature.[9][10]

Material properties

The hydrogel, which contains keratan sulfate in 97% water, has a conductivity of about 1.8 mS/cm, the highest known amongst biological materials.[11]

See also

References

  1. ^ a b Roth A, Tscharntke H (October 1976). "Ultrastructure of the ampullary electroreceptors in lungfish and Brachiopterygii". Cell Tissue Res. 173 (1): 95–108. doi:10.1007/bf00219268. PMID 991235.
  2. ^ a b Gibbs MA, Northcutt RG (2004). "Development of the lateral line system in the shovelnose sturgeon". Brain Behav. Evol. 64 (2): 70–84. doi:10.1159/000079117. PMID 15205543.
  3. ^ Clusin, WT; Bennett, MV (February 1977). "Calcium-activated conductance in skate electroreceptors: current clamp experiments". The Journal of General Physiology. 69 (2): 121–43. doi:10.1085/jgp.69.2.121. PMC 2215012. PMID 190338.
  4. ^ Fields, R. Douglas (August 2007). "The Shark's Electric Sense" (PDF). Scientific American. Retrieved 2 December 2013.
  5. ^ Meyer, Carl G.; Holland, Kim N.; Papastamatiou, Yannis P. (2005). "Sharks can detect changes in the geomagnetic field". Journal of the Royal Society Interface. 2 (2): 129–130. doi:10.1098/rsif.2004.0021. PMC 1578252. PMID 16849172.
  6. ^ Murray RW (1960). "The Response of the Ampullae of Lorenzini of Elasmobranchs to Mechanical Stimulation". J Exp Biol. 37: 417–424.
  7. ^ Murray RW (1960). "Electrical sensitivity of the ampullae of Lorenzini". Nature. 187 (4741): 957. doi:10.1038/187957a0.
  8. ^ Brown BR (2003). "Sensing temperature without ion channels". Nature. 421 (6922): 495. doi:10.1038/421495a. PMID 12556879.
  9. ^ Fields, RD, Fields, KD, Fields, MC (2007). "Semiconductor gel in shark sense organs?". Neurosci. Lett. 426 (3): 166–170. doi:10.1016/j.neulet.2007.08.064. PMC 2211453. PMID 17904741.
  10. ^ Brown BR (2010). "Temperature response in electrosensors and thermal voltages in electrolytes". J Biol Phys. 36 (2): 121–134. doi:10.1007/s10867-009-9174-8. PMC 2825305.
  11. ^ Erik E. Josberger; Pegah Hassanzadeh; Yingxin Deng; Joel Sohn; Michael J. Rego; Chris T. Amemiya; Marco Rolandi (13 May 2016). "Proton conductivity in ampullae of Lorenzini jelly". Science Advances. 2 (5): e1600112. doi:10.1126/sciadv.1600112.

External links

Dwarf sawfish

The dwarf sawfish or Queensland sawfish, Pristis clavata, is a sawfish of the family Pristidae, found in tropical Australia. This endangered species is by far the smallest species in its family.

Electroreception

Electroreception or electroception is the biological ability to perceive natural electrical stimuli. It has been observed almost exclusively in aquatic or amphibious animals, because water is a much better conductor than air. The known exceptions are the monotremes

(echidnas and platypuses), cockroaches and bees. Electroreception is used in electrolocation (detecting objects) and for electrocommunication.

Euryhaline

Euryhaline organisms are able to adapt to a wide range of salinities. An example of a euryhaline fish is the molly (Poecilia sphenops) which can live in fresh water, brackish water, or salt water. The green crab (Carcinus maenas) is an example of a euryhaline invertebrate that can live in salt and brackish water. Euryhaline organisms are commonly found in habitats such as estuaries and tide pools where the salinity changes regularly. However, some organisms are euryhaline because their life cycle involves migration between freshwater and marine environments, as is the case with salmon and eels.

The opposite of euryhaline organisms are stenohaline ones, which can only survive within a narrow range of salinities. Most freshwater organisms are stenohaline, and will die in seawater, and similarly most marine organisms are stenohaline, and cannot live in fresh water.

Fear of fish

Fear of fish or ichthyophobia ranges from cultural phenomena such as fear of eating fish, fear of touching raw fish, or fear of dead fish, up to irrational fear (specific phobia). Galeophobia is the fear specifically of sharks.

Freshwater fish

Freshwater fish are those that spend some or all of their lives in fresh water, such as rivers and lakes, with a salinity of less than 0.05%. These environments differ from marine conditions in many ways, the most obvious being the difference in levels of salinity. To survive fresh water, the fish need a range of physiological adaptations.

41.24% of all known species of fish are found in fresh water. This is primarily due to the rapid speciation that the scattered habitats make possible. When dealing with ponds and lakes, one might use the same basic models of speciation as when studying island biogeography.

Goblin shark

The goblin shark (Mitsukurina owstoni) is a rare species of deep-sea shark. Sometimes called a "living fossil", it is the only extant representative of the family Mitsukurinidae, a lineage some 125 million years old. This pink-skinned animal has a distinctive profile with an elongated, flattened snout, and highly protrusible jaws containing prominent nail-like teeth. It is usually between 3 and 4 m (10 and 13 ft) long when mature, though it can grow considerably larger. Goblin sharks inhabit upper continental slopes, submarine canyons, and seamounts throughout the world at depths greater than 100 m (330 ft), with adults found deeper than juveniles.

Various anatomical features of the goblin shark, such as its flabby body and small fins, suggest that it is sluggish in nature. This species hunts for teleost fishes, cephalopods, and crustaceans both near the sea floor and in the middle of the water column. Its long snout is covered with ampullae of Lorenzini that enable it to sense minute electric fields produced by nearby prey, which it can snatch up by rapidly extending its jaws. Small numbers of goblin sharks are unintentionally caught by deepwater fisheries. The International Union for Conservation of Nature (IUCN) has assessed it as Least Concern, despite its rarity, citing its wide distribution and low incidence of capture.

Lorenzini

Lorenzini is an Italian surname. Notable people with the surname include:

Carlo Lorenzini (1826 – 1890), better known by the pen name C. Collodi, a Florentine's children's writer known for the fairy tale The Adventures of Pinocchio

Carola Lorenzini (1889-1941), Argentine aviator

Davide Lorenzini (born 1969), Italian diver

Orlando Lorenzini (1890-1941), Italian general during World War II

Roberto Lorenzini (born 1966), Italian professional football coach and a former player

Magnetic shark repellent

Magnetic shark repellents utilize permanent magnets, which exploit the sensitivity of the Ampullae of Lorenzini in sharks and rays (electrosense). This organ is not found on bony fishes (teleosts), therefore, this type of shark repellent is selective to sharks and rays. Permanent magnets do not require power input, making them ideal for use in fisheries and as bycatch reduction devices.

Magnetoreception

Magnetoreception (also magnetoception) is a sense which allows an organism to detect a magnetic field to perceive direction, altitude or location. This sensory modality is used by a range of animals for orientation and navigation, and as a method for animals to develop regional maps. For the purpose of navigation, magnetoreception deals with the detection of the Earth's magnetic field.

Magnetoreception is present in bacteria, arthropods, molluscs and members of all major taxonomic groups of vertebrates. Humans are not thought to have a magnetic sense, but there is a protein (a cryptochrome) in the eye which could serve this function.

Metridium Fields

Metridium Fields is the major label debut by San Francisco-based doom metal outfit Giant Squid. It is a re-recorded version of their album Metridium Field, which was released independently by the band two years earlier.

The band's original debut album, Metridium Field was self-released in 2004, and after signing with The End Records in 2005, the band decided to remaster the album for its world-wide release. Upon finding the original master tracks had become unusable, Giant Squid re-recorded the entire album and re-released it as their major label debut and first "official" album. Fields was released on August 22, 2006, and brought the band critical acclaim and a devoted fanbase.

The title of the album and its closing track is named for Metridium, a genus of sea anemone.

Neopterygii

Neopterygii are a group of fish. Neopterygii means "new fins" (from Greek νέος neos, new, and πτέρυξ pteryx, fin). Only a few changes occurred during their evolution from the earlier actinopterygians. They appeared sometime in the Late Permian, before the time of the dinosaurs. The Neopterygii were a very successful group of fish, because they could move more rapidly than their ancestors. Their scales and skeletons began to lighten during their evolution, and their jaws became more powerful and efficient. While electroreception and the ampullae of Lorenzini are present in all other groups of fish, with the exception of hagfish (although hagfish are not Actinopterygii, they are Agnathans), Neopterygii have lost this sense, even if it has later been re-evolved within Gymnotiformes and catfishes, which possess nonhomologous teleost ampullae.

Niger stingray

The Niger stingray or smooth freshwater stingray, Dasyatis garouaensis, is a species of stingray in the family Dasyatidae, native to rivers in Nigeria and Cameroon. Attaining a width of 40 cm (16 in), this species can be distinguished by its thin, almost circular pectoral fin disk, slightly projecting snout tip, and mostly smooth skin with small or absent dermal denticles. The Niger stingray feeds on aquatic insect larvae and is ovoviviparous. The long stinging spine on the tail of this ray can inflict a painful wound. It has been assessed as Vulnerable by the International Union for Conservation of Nature (IUCN), as its numbers are declining in some areas and it faces heavy fishing pressure and habitat degradation.

Outline of sharks

The following outline is provided as an overview of and topical guide to sharks:

Sharks (superorder Selachimorpha) are a type of fish with a full cartilaginous skeleton and a highly streamlined body. The earliest known sharks date from more than 440 million years ago, before the time of the dinosaurs.

Passive electrolocation in fish

Passive electrolocation is a process where certain species of fish or aquatic amphibians can detect electric fields using specialized electroreceptors to detect and to locate the source of an external electric field in its environment creating the electric field. These external electric fields can be produced by any bioelectrical process in an organism, especially by actions of the nerves or muscles of fish, or indeed by the specially developed electric organs of fish. Other fields are induced by movement of a conducting organism through the earth's magnetic field, or from atmospheric electricity.Electrolocating fish use this ability to detect prey, locate other fish, avoid predators, and navigate by the Earth's magnetic field. Electroreceptors probably evolved once or twice early in vertebrate evolution, but the sense was apparently lost in amniotes, and in a large number of the Actinopterygii (ray finned fishes) only to reappear independently in two teleost clades. In fish, the ampullary receptor is a specialized receptor that it uses to sense these electric fields and allows the fish to follow electric field lines to their source. Sharks primarily use specialized receptors, called Ampullae of Lorenzini, to detect their prey's low frequency DC fields and may also use their receptors in navigation by the Earth's magnetic field. Weakly electric fish use their ampullary receptors and tuberous receptors to detect the weakly electric fields produced by other fish, as well as for possible predator avoidance. Passive electrolocation contrasts with active electrolocation, in which the animal emits its own weak self generated electric field and detects nearby objects by detecting the distortion of its produced electric field. In active electrolocation the animal senses its own electromotor discharge or reafference instead of some externally generated electric field or discharge

Proton conductor

A proton conductor is an electrolyte, typically a solid electrolyte, in which H+ are the primary charge carriers.

Sensory systems in fish

Most fish possess highly developed sense organs. Nearly all daylight fish have color vision that is at least as good as a human's (see vision in fishes). Many fish also have chemoreceptors that are responsible for extraordinary senses of taste and smell. Although they have ears, many fish may not hear very well. Most fish have sensitive receptors that form the lateral line system, which detects gentle currents and vibrations, and senses the motion of nearby fish and prey. Sharks can sense frequencies in the range of 25 to 50 Hz through their lateral line.Fish orient themselves using landmarks and may use mental maps based on multiple landmarks or symbols. Fish behavior in mazes reveals that they possess spatial memory and visual discrimination.

Stefano Lorenzini

Stefano Lorenzini (born around 1652, Florence, Italy — date of death unknown) was an Italian physician and noted ichthyologist. He studied medicine in Pisa and surgery at the Hospital of St. Florence Maria Nuova, with teachers including Francesco Redi, Nicholas Steno, John Fynch among other prominent scholars. He fell into disgrace with the Grand Duke Cosimo III de' Medici, who imprisoned him along with his brother Lorenzo Lorenzini, a famous mathematician.

His observations on sharks, published in Florence in 1678, offers an example of the extraordinary depth study on animal anatomy and physiology, based on new mechanistic and corpuscular perspectives applied to the study of the living. He is most famous for the discovery of the so-called Ampullae of Lorenzini, special electromagnetic sense organs possessed by the Elasmobranchii (sharks and rays), which are located in front of the head and form a network of canals filled with gel.

Stingray

Stingrays are a group of sea rays, which are cartilaginous fish related to sharks. Many species are endangered. They are classified in the suborder Myliobatoidei of the order Myliobatiformes and consist of eight families: Hexatrygonidae (sixgill stingray), Plesiobatidae (deepwater stingray), Urolophidae (stingarees), Urotrygonidae (round rays), Dasyatidae (whiptail stingrays), Potamotrygonidae (river stingrays), Gymnuridae (butterfly rays), and Myliobatidae (eagle rays).Stingrays are common in coastal tropical and subtropical marine waters throughout the world. Some species, such as Dasyatis thetidis, are found in warmer temperate oceans, and others, such as Plesiobatis daviesi, are found in the deep ocean. The river stingrays, and a number of whiptail stingrays (such as the Niger stingray), are restricted to fresh water. Most myliobatoids are demersal (inhabiting the next-to-lowest zone in the water column), but some, such as the pelagic stingray and the eagle rays, are pelagic.There are about 220 known stingray species organized into 10 families and 29 genera. Stingray species are progressively becoming threatened or vulnerable to extinction, particularly as the consequence of unregulated fishing. As of 2013, 45 species have been listed as vulnerable or endangered by the IUCN. The status of some other species is poorly known, leading to their being listed as data deficient.

Winghead shark

The winghead shark (Eusphyra blochii) is a species of hammerhead shark, and part of the family Sphyrnidae. Reaching a length of 1.9 m (6.2 ft), this small brown to gray shark has a slender body with a tall, sickle-shaped first dorsal fin. Its name comes from its exceptionally large "hammer", or cephalofoil, which can be as wide as half of the shark's total length. The function of this structure is unclear, but may relate to the shark's senses. The wide spacing of its eyes grants superb binocular vision, while the extremely long nostrils on the leading margin of the cephalofoil may allow for better detection and tracking of odor trails in the water. The cephalofoil also provides a large surface area for its ampullae of Lorenzini and lateral line, with potential benefits for electroreception and mechanoreception, respectively.

Inhabiting the shallow coastal waters of the central and western Indo-Pacific, the winghead shark feeds on small bony fishes, crustaceans, and cephalopods. It gives birth to live young, with the developing embryos receiving nourishment through a placental connection. Females produce annual litters of 6 to 25 pups; depending on region, birthing may occur from February to June after a gestation period of 8–11 months. This harmless species is widely fished for meat, fins, liver oil, and fishmeal. The International Union for Conservation of Nature has assessed it as Endangered in 2016, as it is thought to have declined in some parts of its range due to overfishing.

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