Electroreception and electrogenesis

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The elephantnose fish is a weakly electric mormyrid fish which generates an electric field with its electric organ and then uses its electroreceptive knollenorgans and mormyromasts to locate nearby objects by the distortions they cause in the electric field. Electroreception system in Elephantfish.svg
The elephantnose fish is a weakly electric mormyrid fish which generates an electric field with its electric organ and then uses its electroreceptive knollenorgans and mormyromasts to locate nearby objects by the distortions they cause in the electric field.

Electroreception and electrogenesis are the closely related biological abilities to perceive electrical stimuli and to generate electric fields. Both are used to locate prey; stronger electric discharges are used in a few groups of fishes (most famously the electric eel, which is not actually an eel but a knifefish) to stun prey. The capabilities are found almost exclusively in aquatic or amphibious animals, since water is a much better conductor of electricity than air. In passive electrolocation, objects such as prey are detected by sensing the electric fields they create. In active electrolocation, fish generate a weak electric field and sense the different distortions of that field created by objects that conduct or resist electricity. Active electrolocation is practised by two groups of weakly electric fish, the Gymnotiformes (knifefishes) and the Mormyridae (elephantfishes), and by Gymnarchus niloticus, the African knifefish. An electric fish generates an electric field using an electric organ, modified from muscles in its tail. The field is called weak if it is only enough to detect prey, and strong if it is powerful enough to stun or kill. The field may be in brief pulses, as in the elephantfishes, or a continuous wave, as in the knifefishes. Some strongly electric fish, such as the electric eel, locate prey by generating a weak electric field, and then discharge their electric organs strongly to stun the prey; other strongly electric fish, such as the electric ray, electrolocate passively. The stargazers are unique in being strongly electric but not using electrolocation.

Contents

The electroreceptive ampullae of Lorenzini evolved early in the history of the vertebrates; they are found in both cartilaginous fishes such as sharks, and in bony fishes such as coelacanths and sturgeons, and must therefore be ancient. Most bony fishes have secondarily lost their ampullae of Lorenzini, but other non-homologous electroreceptors have repeatedly evolved, including in two groups of mammals, the monotremes (platypus and echidnas) and the cetaceans (Guiana dolphin).

History

Hans Lissmann discovered electroreception in 1950 through his observations of Gymnarchus niloticus. Gymnarchus niloticus005 (cropped).JPG
Hans Lissmann discovered electroreception in 1950 through his observations of Gymnarchus niloticus .

In 1678, while doing dissections of sharks, the Italian physician Stefano Lorenzini discovered organs on their heads now called ampullae of Lorenzini. He published his findings in Osservazioni intorno alle torpedini. [3] The electroreceptive function of these organs was established by R. W. Murray in 1960. [4] [5]

In 1921, the German anatomist Viktor Franz described the knollenorgans (tuberous organs) in the skin of the elephantfishes, again without knowledge of their function as electroreceptors. [6]

In 1949, the Ukrainian-British zoologist Hans Lissmann noticed that the African knife fish (Gymnarchus niloticus) was able to swim backwards at the same speed and with the same dexterity around obstacles as when it swam forwards, avoiding collisions. He demonstrated in 1950 that the fish was producing a variable electric field, and that the fish reacted to any change in the electric field around it. [2] [7]

Electrolocation

Electroreceptors in a sharks head.svg
The electroreceptive ampullae of Lorenzini (red dots) evolved from the mechanosensory lateral line organs (gray lines) of early vertebrates. [8] They are seen here in the head of a shark.
Ampullae of Lorenzini.svg
Ampullae of Lorenzini, found in several basal groups of fishes, are jelly-filled canals connecting pores in the skin to sensory bulbs. They detect small differences in electrical potential between their two ends.

Electroreceptive animals use the sense to locate objects around them. This is important in ecological niches where the animal cannot depend on vision: for example in caves, in murky water, and at night. Electrolocation can be passive, sensing electric fields such as those generated by the muscle movements of buried prey, or active, the electrogenic predator generating a weak electric field to allow it to distinguish between conducting and non-conducting objects in its vicinity. [9]

Passive electrolocation

In passive electrolocation, the animal senses the weak bioelectric fields generated by other animals and uses it to locate them. These electric fields are generated by all animals due to the activity of their nerves and muscles. A second source of electric fields in fish is the ion pump associated with osmoregulation at the gill membrane. This field is modulated by the opening and closing of the mouth and gill slits. [10] [11] Passive electroreception usually relies upon ampullary receptors such as ampullae of Lorenzini which are sensitive to low frequency stimuli, below 50 Hz. These receptors have a jelly-filled canal leading from the sensory receptors to the skin surface. [8] [9]

Active electrolocation

Knollenorgan by Viktor Franz 1921.gif
A knollenorgan, a tuberous electroreceptor of weakly electric fish. RC=receptor cell; b.m.=basal membrane; n=nerve.
Mormyromast diagram.svg
A Mormyromast, a type of electroreceptor found only in Mormyrid fishes

In active electrolocation, [12] the animal senses its surrounding environment by generating weak electric fields (electrogenesis) and detecting distortions in these fields using electroreceptor organs. This electric field is generated by means of a specialised electric organ consisting of modified muscle or nerves. [13] Animals that use active electroreception include the weakly electric fish, which either generate small electrical pulses (termed "pulse-type"), as in the Mormyridae, or produce a quasi-sinusoidal discharge from the electric organ (termed "wave-type"), as in the Gymnotidae. [14]

Many of these fish, such as Gymnarchus and Apteronotus , keep their body rather rigid, swimming forwards or backwards with equal facility by undulating fins that extend most of the length of their bodies. Swimming backwards may help them to search for and assess prey using electrosensory cues. Experiments by Lannoo and Lannoo in 1993 support Lissmann's proposal that this style of swimming with a straight back works effectively given the constraints of active electrolocation. Apteronotus can select and catch larger Daphnia water fleas among smaller ones, and they do not discriminate against artificially-darkened water fleas, in both cases with or without light. [7] [15]

These fish create a potential usually smaller than one volt (1 V). Weakly electric fish can discriminate between objects with different resistance and capacitance values, which may help in identifying objects. Active electroreception typically has a range of about one body length, though objects with an electrical impedance similar to that of the surrounding water are nearly undetectable. [12] [13] [14]

Active electrolocation relies upon tuberous electroreceptors which are sensitive to high frequency (20-20,000  Hz) stimuli. These receptors have a loose plug of epithelial cells which capacitively couples the sensory receptor cells to the external environment. Elephantfish (Mormyridae) from Africa have tuberous electroreceptors known as Knollenorgans and Mormyromasts in their skin. [16] [17] [18]

Elephantfish emit short pulses to locate their prey. Capacitative and resistive objects affect the electric field differently, enabling the fish to locate objects of different types within a distance of about a body length. Resistive objects increase the amplitude of the pulse; capacitative objects introduce distortions. [1]

The Gymnotiformes, including the glass knifefish (Sternopygidae) and the electric eel (Gymnotidae), differ from the Mormyridae in emitting a continuous wave, approximating a sine wave, from their electric organ. As in the Mormyridae, the generated electric field enables them to discriminate accurately between capacitative and resistive objects. [1]

Electroreception of Capacitative and Resistive Objects in Glass Knifefish.svg

Electrolocation of capacitative and resistive objects in glass knifefish.
Many gymnotid fish generate a continuous electrical wave, which is
distorted differently by objects according to their conductivity.

 Electric eel's electric organs.svg

The electric eel's electric organs occupy much of its body.
They can discharge both weakly for electrolocation
and strongly to stun prey.

Electrocommunication

Electric eels create electric fields powerful enough to stun prey using modified muscles. Some weakly electric knifefishes appear to mimic the electric eel's discharge patterns; this may be Batesian mimicry, to deceive predators that they are too dangerous to attack. Sidderaal (4039238527).jpg
Electric eels create electric fields powerful enough to stun prey using modified muscles. Some weakly electric knifefishes appear to mimic the electric eel's discharge patterns; this may be Batesian mimicry, to deceive predators that they are too dangerous to attack.

Weakly electric fish can communicate by modulating the electrical waveform they generate. They may use this to attract mates and in territorial displays. [21] Electric catfish frequently use their electric discharges to ward off other species from their shelter sites, whereas with their own species they have ritualized fights with open-mouth displays and sometimes bites, but rarely use electric organ discharges. [22]

When two glass knifefishes (Sternopygidae) come close together, both individuals shift their discharge frequencies in a jamming avoidance response. [13]

In bluntnose knifefishes, Brachyhypopomus , the electric discharge pattern is similar to the low voltage electrolocative discharge of the electric eel, Electrophorus. This is hypothesized to be Batesian mimicry of the powerfully-protected electric eel. [20] Brachyhypopomus males produce a continuous electric "hum" to attract females; this consumes 11–22% of their total energy budget, whereas female electrocommunication consumes only 3%. Large males produced signals of larger amplitude, and these are preferred by the females. The cost to males is reduced by a circadian rhythm, with more activity coinciding with night-time courtship and spawning, and less at other times. [23]

Fish that prey on electrolocating fish may "eavesdrop" [24] on the discharges of their prey to detect them. The electroreceptive African sharptooth catfish (Clarias gariepinus) may hunt the weakly electric mormyrid, Marcusenius macrolepidotus in this way. [25] This has driven the prey, in an evolutionary arms race, to develop more complex or higher frequency signals that are harder to detect. [26]

Some shark embryos and pups "freeze" when they detect the characteristic electric signal of their predators. [10]

Evolution and taxonomic distribution

In vertebrates, passive electroreception is an ancestral trait, meaning that it was present in their last common ancestor. [27] The ancestral mechanism is called ampullary electroreception, from the name of the receptive organs involved, ampullae of Lorenzini. These evolved from the mechanical sensors of the lateral line, and exist in cartilaginous fishes (sharks, rays, and chimaeras), lungfishes, bichirs, coelacanths, sturgeons, paddlefishes, aquatic salamanders, and caecilians. Ampullae of Lorenzini appear to have been lost early in the evolution of bony fishes and tetrapods, though the evidence for absence in many groups is incomplete and unsatisfactory. [27] Where electroreception does occur in these groups, it has secondarily been acquired in evolution, using organs other than and not homologous with ampullae of Lorenzini. [8] [27]

Electric organs have evolved at least eight separate times, each one forming a clade: twice during the evolution of cartilaginous fishes, creating the electric skates and rays, and six times during the evolution of the bony fishes. [28] Passively-electrolocating groups, including those that move their heads to direct their electroreceptors, are shown without symbols. Non-electrolocating species are not shown. [27] Actively electrolocating fish are marked with a small yellow lightning flash Farm-Fresh lightning.png and their characteristic discharge waveforms. [29] Fish able to deliver electric shocks are marked with a red lightning flash Lightning Symbol.svg . [27]

Vertebrates
Lampreys

Petromyzon marinus.jpg

End‑bud recep.
Jawed fishes
Cartilaginous fishes

Selachimorpha (sharks) Tiburon portada.jpg

Batoidea

Torpediniformes (electric rays) Farm-Fresh lightning.png Lightning Symbol.svg Electric ray waveform.svg Fish4345 - Flickr - NOAA Photo Library (white background).jpg

other rays Mobula mobular.jpg

Rajidae (skates) Farm-Fresh lightning.png Skate waveform.svg Raja montagui2.jpg

430  mya
Bony fishes
Lobe-finned fishes

Coelacanths Coelacanth flipped.png

Lungfishes Barramunda coloured.jpg

Amphibians

(aquatic salamanders, caecilians; others: lost) Aquatic life (1916-1917) (19559021800) (cropped).jpg

Mammals
Monotremes

(platypus, echidna) Feeding Platypus (6811147158) (white background).jpg Zaglossus bartoni - MUSE.JPG

glands in snout
Cetaceans

(Guiana dolphin) Sotalia guianensis (white background).jpg

vibrissal crypts 
(lost)
Ray-finned fishes
425  mya
Amp. of Lorenz.

Cartilaginous fish

Sharks and rays ( Elasmobranchii ) rely on electrolocation using their ampullae of Lorenzini in the final stages of their attacks, as can be demonstrated by the robust feeding response elicited by electric fields similar to those of their prey. Sharks are the most electrically sensitive animals known, responding to direct current fields as low as 5 nV/cm. [30] [31] [32] [33]

Bony fish

Two groups of teleost fishes are weakly electric and actively electroreceptive: the Neotropical knifefishes (Gymnotiformes) and the African elephantfishes (Notopteroidei), enabling them to navigate and find food in turbid water. [34] The Gymnotiformes include the electric eel, which besides the group's use of low-voltage electrolocation, is able to generate high voltage electric shocks to stun its prey. Such powerful electrogenesis makes use of large electric organs modified from muscles. These consist of a stack of electrocytes, each capable of generating a small voltage; the voltages are effectively added together (in series) to provide a powerful electric organ discharge. [35] [36]

Monotremes

The platypus is a monotreme mammal that has secondarily acquired electroreception. Its receptors are arranged in stripes on its bill, giving it high sensitivity to the sides and below; it makes quick turns of its head as it swims to detect prey. Platypus electrolocation.svg
The platypus is a monotreme mammal that has secondarily acquired electroreception. Its receptors are arranged in stripes on its bill, giving it high sensitivity to the sides and below; it makes quick turns of its head as it swims to detect prey.

The monotremes, including the semi-aquatic platypus and the terrestrial echidnas, are one of the only groups of mammals that have evolved electroreception. While the electroreceptors in fish and amphibians evolved from mechanosensory lateral line organs, those of monotremes are based on cutaneous glands innervated by trigeminal nerves. The electroreceptors of monotremes consist of free nerve endings located in the mucous glands of the snout. Among the monotremes, the platypus (Ornithorhynchus anatinus) has the most acute electric sense. [37] [38] The platypus localises its prey using almost 40,000 electroreceptors arranged in front-to-back stripes along the bill. [34] The arrangement is highly directional, being most sensitive off to the sides and below. By making short quick head movements called saccades, platypuses accurately locate their prey. The platypus appears to use electroreception along with pressure sensors to determine the distance to prey from the delay between the arrival of electrical signals and pressure changes in water. [38]

The electroreceptive capabilities of the four species of echidna are much simpler. Long-beaked echidnas (genus Zaglossus) have some 2,000 receptors, while short-beaked echidnas (Tachyglossus aculeatus) have around 400, near the end of the snout. [34] This difference can be attributed to their habitat and feeding methods. Western long-beaked echidnas feed on earthworms in leaf litter in tropical forests, wet enough to conduct electrical signals well. Short-beaked echidnas feeds mainly on termites and ants, which live in nests in dry areas; the nest interiors are presumably humid enough for electroreception to work. [39] Experiments have shown that echidnas can be trained to respond to weak electric fields in water and moist soil. The electric sense of the echidna is hypothesised to be an evolutionary remnant from a platypus-like ancestor. [38]

Dolphins

Dolphins have evolved electroreception in structures different from those of fish, amphibians and monotremes. The hairless vibrissal crypts on the rostrum of the Guiana dolphin (Sotalia guianensis), originally associated with mammalian whiskers, are capable of electroreception as low as 4.8 μV/cm, sufficient to detect small fish. This is comparable to the sensitivity of electroreceptors in the platypus. [40]

Bees

Until recently, electroreception was known only in vertebrates. Recent research has shown that bees can detect the presence and pattern of a static charge on flowers. [41]

See also

Related Research Articles

<span class="mw-page-title-main">Platypus</span> Species of mammal

The platypus, sometimes referred to as the duck-billed platypus, is a semiaquatic, egg-laying mammal endemic to eastern Australia, including Tasmania. The platypus is the sole living representative or monotypic taxon of its family Ornithorhynchidae and genus Ornithorhynchus, though a number of related species appear in the fossil record.

<span class="mw-page-title-main">Echidna</span> Family of mammals

Echidnas, sometimes known as spiny anteaters, are quill-covered monotremes belonging to the family Tachyglossidae, living in Australia and New Guinea. The four extant species of echidnas and the platypus are the only living mammals that lay eggs and the only surviving members of the order Monotremata. The diet of some species consists of ants and termites, but they are not closely related to the American true anteaters or to hedgehogs. Their young are called puggles.

<span class="mw-page-title-main">Gymnotiformes</span> Order of bony fishes

The Gymnotiformes are an order of teleost bony fishes commonly known as Neotropical knifefish or South American knifefish. They have long bodies and swim using undulations of their elongated anal fin. Found almost exclusively in fresh water, these mostly nocturnal fish are capable of producing electric fields to detect prey, for navigation, communication, and, in the case of the electric eel, attack and defense. A few species are familiar to the aquarium trade, such as the black ghost knifefish, the glass knifefish, and the banded knifefish.

<span class="mw-page-title-main">Lateral line</span> Sensory system in fish

The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial cells, known as hair cells, which respond to displacement caused by motion and transduce these signals into electrical impulses via excitatory synapses. Lateral lines play an important role in schooling behavior, predation, and orientation.

<span class="mw-page-title-main">Electric fish</span> Fish that can generate electric fields

An electric fish is any fish that can generate electric fields. Most electric fish are also electroreceptive, meaning that they can sense electric fields. The only exception is the stargazer family (Uranoscopidae). Electric fish, although a small minority of all fishes, include both oceanic and freshwater species, and both cartilaginous and bony fishes.

<span class="mw-page-title-main">Black ghost knifefish</span> Species of fish

The black ghost knifefish is a tropical fish belonging to the ghost knifefish family (Apteronotidae). They originate in freshwater habitats in South America where they range from Venezuela to the Paraguay–Paraná River, including the Amazon Basin. They are popular in aquaria. The fish is all black except for two white rings on its tail, and a white blaze on its nose, which can occasionally extend into a stripe down its back. It moves mainly by undulating a long fin on its underside. It will grow to a length of 18"-20". Only a fish for those with large aquariums, minimum 100 gallons.

<span class="mw-page-title-main">Magnetoreception</span> Biological ability to perceive magnetic fields

Magnetoreception is a sense which allows an organism to detect the Earth's magnetic field. Animals with this sense include some arthropods, molluscs, and vertebrates. The sense is mainly used for orientation and navigation, but it may help some animals to form regional maps. Experiments on migratory birds provide evidence that they make use of a cryptochrome protein in the eye, relying on the quantum radical pair mechanism to perceive magnetic fields. This effect is extremely sensitive to weak magnetic fields, and readily disturbed by radio-frequency interference, unlike a conventional iron compass.

<span class="mw-page-title-main">Ampullae of Lorenzini</span> Sensory organs in some fish that detect electrical fields

Ampullae of Lorenzini are electroreceptors, sense organs able to detect electric fields. They form a network of mucus-filled pores in the skin of cartilaginous fish and of basal bony fishes such as reedfish, sturgeon, and lungfish. They are associated with and evolved from the mechanosensory lateral line organs of early vertebrates. Most bony fishes and terrestrial vertebrates have lost their ampullae of Lorenzini.

<span class="mw-page-title-main">Mormyridae</span> Family of fishes

The Mormyridae, sometimes called "elephantfish", are a superfamily of weakly electric fish in the order Osteoglossiformes native to Africa. It is by far the largest family in the order, with around 200 species. Members of the family can be popular, if challenging, aquarium species. These fish have a large brain size and unusually high intelligence.

<span class="mw-page-title-main">Hans Lissmann (zoologist)</span> British zoologist

Hans Werner Lissmann FRS was a British zoologist of Ukrainian provenance, specialising in animal behaviour.

<span class="mw-page-title-main">Peters's elephantnose fish</span> Species of fish

Peters's elephant-nose fish is an African freshwater elephantfish in the genus Gnathonemus. Other names in English include elephantnose fish, long-nosed elephant fish, and Ubangi mormyrid, after the Ubangi River. The Latin name petersii is probably for the German naturalist Wilhelm Peters. The fish uses electrolocation to find prey, and has the largest brain-to-body oxygen use ratio of all known vertebrates.

<span class="mw-page-title-main">Electric organ (fish)</span> Organ in electric fish

In biology, the electric organ is an organ that an electric fish uses to create an electric field. Electric organs are derived from modified muscle or in some cases nerve tissue, called electrocytes, and have evolved at least six times among the elasmobranchs and teleosts. These fish use their electric discharges for navigation, communication, mating, defence, and in strongly electric fish also for the incapacitation of prey.

<i>Gymnarchus</i> Genus of ray-finned fishes

Gymnarchus niloticus – commonly known as the aba, aba aba, frankfish, freshwater rat-tail, poisson-cheval, or African knifefish – is an electric fish, and the only species in the genus Gymnarchus and the family Gymnarchidae within the order Osteoglossiformes. It is found in swamps, lakes and rivers in the Nile, Turkana, Chad, Niger, Volta, Senegal, and Gambia basins.

<span class="mw-page-title-main">Knollenorgan</span>

A Knollenorgan is an electroreceptor in the skin of weakly electric fish of the family Mormyridae (Elephantfish) from Africa. The structure was first described by Viktor Franz (1921), a German anatomist unaware of its function. They are named after "Knolle", German for "tuberous root" which describes their structure.

Sternarchogiton porcinum is a species of weakly electric knifefish in the family Apteronotidae. It is native to deep river channels in the Río Huallaga, Río Napo, and Río Amazonas in Peru, and in the Río Orinoco in Venezuela. Many specimens once identified as S. porcinum from the Brazilian Amazon Basin and the Venezuelan Orinoco are now known to be a different species, S. preto.

<span class="mw-page-title-main">Jamming avoidance response</span> Behavior performed by weakly electric fish to prevent jamming of their sense of electroreception

The jamming avoidance response is a behavior of some species of weakly electric fish. It occurs when two electric fish with wave discharges meet – if their discharge frequencies are very similar, each fish shifts its discharge frequency to increase the difference between the two. By doing this, both fish prevent jamming of their sense of electroreception.

<span class="mw-page-title-main">Hydrodynamic reception</span> Ability of an organism to sense water movements

In animal physiology, hydrodynamic reception refers to the ability of some animals to sense water movements generated by biotic or abiotic sources. This form of mechanoreception is useful for orientation, hunting, predator avoidance, and schooling. Frequent encounters with conditions of low visibility can prevent vision from being a reliable information source for navigation and sensing objects or organisms in the environment. Sensing water movements is one resolution to this problem.

Most fish possess highly developed sense organs. Nearly all daylight fish have colour vision that is at least as good as a human's. 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.

<span class="mw-page-title-main">Mormyroidea</span> Superfamily of fishes

The Mormyroidea are a superfamily of fresh water fishes endemic to Africa that, together with the families Hiodontidae, Osteoglossidae, Pantodontidae and Notopteridae, represents one of the main groups of living Osteoglossiformes. They stand out for their use of weak electric fields, which they use to orient themselves, reproduce, feed, and communicate.

The history of bioelectricity dates back to ancient Egypt, where the shocks delivered by the electric catfish were used medicinally.

References

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