Otolith
Encyclopedia
An otolith, also called statoconium or otoconium is a structure in the saccule
Saccule
The saccule is a bed of sensory cells situated in the inner ear. The saccule translates head movements into neural impulses which the brain can interpret. The saccule is sensitive to linear translations of the head, specifically movements up and down...

 or utricle of the inner ear
Inner ear
The inner ear is the innermost part of the vertebrate ear. In mammals, it consists of the bony labyrinth, a hollow cavity in the temporal bone of the skull with a system of passages comprising two main functional parts:...

, specifically in the vestibular labyrinth of vertebrates. The saccule and utricle, in turn, together make the otolith organs. They are sensitive to gravity and linear acceleration
Acceleration
In physics, acceleration is the rate of change of velocity with time. In one dimension, acceleration is the rate at which something speeds up or slows down. However, since velocity is a vector, acceleration describes the rate of change of both the magnitude and the direction of velocity. ...

. Because of their orientation in the head, the utricle is sensitive to a change in horizontal movement, and the saccule
Saccule
The saccule is a bed of sensory cells situated in the inner ear. The saccule translates head movements into neural impulses which the brain can interpret. The saccule is sensitive to linear translations of the head, specifically movements up and down...

 gives information about vertical acceleration (such as when in an elevator
Elevator
An elevator is a type of vertical transport equipment that efficiently moves people or goods between floors of a building, vessel or other structures...

).

Description

Endolymphatic infillings such as an otolith, (οτο-, oto-, ear + λιθος, lithos, a stone) or statoconia are structure in the saccule
Saccule
The saccule is a bed of sensory cells situated in the inner ear. The saccule translates head movements into neural impulses which the brain can interpret. The saccule is sensitive to linear translations of the head, specifically movements up and down...

 or utricle of the inner ear
Inner ear
The inner ear is the innermost part of the vertebrate ear. In mammals, it consists of the bony labyrinth, a hollow cavity in the temporal bone of the skull with a system of passages comprising two main functional parts:...

, specifically in the vestibular labyrinth of all vertebrates (fish, amphibians, reptiles, mammals and birds). In vertebrate the saccule and utricle together make the otolith organs. Both statoconia and otoliths are used as gravity, balance, movement, and directional indicators in all vertebrates and have a secondary function in sound detection in higher aquatic and terrestrial vertebrates. They are sensitive to gravity and linear acceleration
Acceleration
In physics, acceleration is the rate of change of velocity with time. In one dimension, acceleration is the rate at which something speeds up or slows down. However, since velocity is a vector, acceleration describes the rate of change of both the magnitude and the direction of velocity. ...

. Because of their orientation in the head, the utricle is sensitive to a change in horizontal movement, and the saccule
Saccule
The saccule is a bed of sensory cells situated in the inner ear. The saccule translates head movements into neural impulses which the brain can interpret. The saccule is sensitive to linear translations of the head, specifically movements up and down...

 gives information about vertical acceleration (such as when in an elevator
Elevator
An elevator is a type of vertical transport equipment that efficiently moves people or goods between floors of a building, vessel or other structures...

).

Statoliths can be found in many invertebrate groups but are not contained in the structure of an inner ear Mollusk statoliths are of a similar morphology to the displacement-sensitive organs of vertebrates; however, the function of the mollusk statocyst is restricted to gravity detection and possibly some detection of angular momentum. These are analogous structures, with similar form and function but not descended from a common structure.

Statoconia (also called otoconia) are numerous grains, often spherical in shape, between 1–50 microns; collectively, statoconia are also sometimes termed a statocyst. Otoliths (also called statoliths) are agglutinated crystals or crystals precipitated around a nucleus, with well defined morphology and together all may be termed endolymphatic infillings.

Mechanism

The semicircular canals and sacs in all vertebrates are attached to endolymphatic ducts, which in some groups (such as sharks) end in small openings, called endolymphatic pores, on the dorsal surface of the head. Extrinsic grains may enter through these openings, typically less than a millimeter in diameter. The size of material that enters is limited to sand-sized particles and in the case of sharks is bound together with endogenous
organic matrix that the animal secretes.

In mammals, otoliths are small particles, composed of a combination of a gelatinous matrix and calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...

 in the viscous fluid of the saccule and utricle. The inertia
Inertia
Inertia is the resistance of any physical object to a change in its state of motion or rest, or the tendency of an object to resist any change in its motion. It is proportional to an object's mass. The principle of inertia is one of the fundamental principles of classical physics which are used to...

 of these small particles causes them to stimulate hair cells when the head moves. The hair cells are made up of 40 to 70 stereocilia and one hair cell that is connected to the nerve called kinocilium. When the body changes position or begins a movement the weight of the membrane bends the stereocilia and stimulates the hair cells. Hair cells send signals down sensory nerve fibers, which are interpreted by the brain as motion. The brain interprets the orientation of the head by comparing the input from the utricules and saccules from both ears to the input from the eyes, allowing the brain to discriminate a tilted head from movement of the entire body. When the head is in a normal upright position, the otolith presses on the sensory hair cell receptors. This pushes the hair cell processes down and prevents them from moving side to side. However, when the head is tilted, the pull of gravity on statoconia shift the hair cell processes to the side, distorting them and sending a message to the central nervous system
Central nervous system
The central nervous system is the part of the nervous system that integrates the information that it receives from, and coordinates the activity of, all parts of the bodies of bilaterian animals—that is, all multicellular animals except sponges and radially symmetric animals such as jellyfish...

 that the head is no longer level but now tilted. (see: BPPV) This theory may have to reevaluated because of an experiment in which a blindfolded owl in zero gravity was able to keep its head level while a handler was rocking its body back and forth.

In 1991, Martin Lenhardt of the University of Michigan
University of Michigan
The University of Michigan is a public research university located in Ann Arbor, Michigan in the United States. It is the state's oldest university and the flagship campus of the University of Michigan...

 discovered that people can hear ultrasonic
Ultrasound
Ultrasound is cyclic sound pressure with a frequency greater than the upper limit of human hearing. Ultrasound is thus not separated from "normal" sound based on differences in physical properties, only the fact that humans cannot hear it. Although this limit varies from person to person, it is...

 speech, perhaps using the saccule as a hearing organ.

Paleontology and evolution

After the death and decomposition of a fish, otoliths and statoconia may preserved within the body of an organism or be dispersed before burial and fossil
Fossil
Fossils are the preserved remains or traces of animals , plants, and other organisms from the remote past...

ization. Dispersed otoliths are one of the many microfossils which can be found though a micropalaeontological analysis of a fine sediment. Their stratigraphic significance is minimal, but can still be used to characterize a level or interval. Fossil otoliths are rarely found in situ (on the remains of the animal), likely because they are not recognized separately from the surrounding rock matrix. In some cases, due to differences in colour, grain size, or a distinctive shape, they can be identified. These rare cases are of special significance, since the presence, composition, and morphology of the material can clarify the relationship of species and groups, as in the case of lineages of primitive fish which shows that endolymphatic infillings were similar in elemental composition to the rock matrix but were restricted to coarse grained material, which presumably is better for the detection of gravity, displacement, and sound. The presence of these extrinsic grains, in osteostracans, chondrichthyans, and acanthodians indicates a common inner ear physiology and presence of open endolymphatic ducts.

Composition

The composition of fish otoliths is also proving useful to fisheries scientists. The calcium carbonate that the otolith is composed of is primarily derived from the water. As the otolith grows, new calcium carbonate, usually aragonite
Aragonite
Aragonite is a carbonate mineral, one of the two common, naturally occurring, crystal forms of calcium carbonate, CaCO3...

 but sometimes vaterite
Vaterite
Vaterite is a mineral, a polymorph of calcium carbonate. It was named after the German mineralogist Heinrich Vater. It is also known as mu-calcium carbonate and has a JCPDS number of 13-192. Vaterite, like aragonite, is a metastable phase of calcium carbonate at ambient conditions at the surface...

, crystals form. As with any crystal structure, lattice vacancies will exist during crystal formation allowing trace elements from the water to bind with the otolith. Studying the trace elemental composition or isotopic signatures of trace elements within a fish otolith gives insight to the water bodies fish have previously occupied. The most studied trace and isotopic signatures are strontium due to the same charge and similar ionic radius to calcium; however, scientists can study multiple trace elements within an otolith to discriminate more specific signatures. A common tool used to measure trace elements in an otolith is a laser ablation inductively coupled plasma mass spectrometer. This tool can measure a variety of trace elements simultaneously. A secondary ion mass spectrometer can also be used. This instrument can allow for greater chemical resolution but can only measure one trace element at a time. The hope of this research is to provide scientists with valuable information on where fish have traveled. Combined with otolith annuli, scientists can add how old fish were when they traveled through different water bodies. All this information can be used to determine fish life cycles so that fisheries scientists can make better informed decisions about fish stocks.

Growth rate and age

Finfish (class Osteichthyes
Osteichthyes
Osteichthyes , also called bony fish, are a taxonomic group of fish that have bony, as opposed to cartilaginous, skeletons. The vast majority of fish are osteichthyes, which is an extremely diverse and abundant group consisting of over 29,000 species...

) have three pairs of otoliths – the sagittae (singular sagitta), lapilli (singular lapillus), and asterisci (singular asteriscus). The sagittae are largest, found just behind the eyes and approximately level with them vertically. The lapilli and asterisci (smallest of the three) are located within the semicircular canals.

The shapes and proportional sizes of the otoliths vary with fish species. In general, fish from highly structured habitats such as reefs or rocky bottoms (e.g. snappers
Lutjanidae
Snappers are a family of perciform fish, mainly marine but with some members inhabiting estuaries, feeding in freshwater. Some are important food fish. One of the best known is the red snapper....

, grouper
Grouper
Groupers are fish of any of a number of genera in the subfamily Epinephelinae of the family Serranidae, in the order Perciformes.Not all serranids are called groupers; the family also includes the sea basses. The common name grouper is usually given to fish in one of two large genera: Epinephelus...

s, many drums and croakers
Sciaenidae
Sciaenidae is a family of fish commonly called drums, croakers, or hardheads for the repetitive throbbing or drumming sounds they make...

) will have larger otoliths than fish that spend most of their time swimming at high speed in straight lines in the open ocean (e.g. tuna
Tuna
Tuna is a salt water fish from the family Scombridae, mostly in the genus Thunnus. Tuna are fast swimmers, and some species are capable of speeds of . Unlike most fish, which have white flesh, the muscle tissue of tuna ranges from pink to dark red. The red coloration derives from myoglobin, an...

, mackerel
Mackerel
Mackerel is a common name applied to a number of different species of fish, mostly, but not exclusively, from the family Scombridae. They may be found in all tropical and temperate seas. Most live offshore in the oceanic environment but a few, like the Spanish mackerel , enter bays and can be...

, dolphinfish
Coryphaenidae
The Coryphaenidae are a family of marine ray-finned fish belonging to the Order Perciformes. The family contains only one genus, Coryphaena, which contains two species, both of which have compressed heads and single dorsal fins that run the entire length of the fish's bodies...

). Flying fish have unusually large otoliths, possibly due to their need for balance when launching themselves out of the water to "fly" in the air. Often, the fish species can be identified from distinct morphological characteristics of an isolated otolith.

Fish otoliths accrete layers of calcium carbonate
Calcium carbonate
Calcium carbonate is a chemical compound with the formula CaCO3. It is a common substance found in rocks in all parts of the world, and is the main component of shells of marine organisms, snails, coal balls, pearls, and eggshells. Calcium carbonate is the active ingredient in agricultural lime,...

 and gelatinous matrix throughout their lives. The accretion rate varies with growth of the fish – often less growth in winter and more in summer – which results in the appearance of rings that resemble tree rings. By counting the rings, it is possible to determine the age of the fish in years. Typically the sagitta is used, as it is largest, but sometimes lapilli are used if they have a more convenient shape. The asteriscus, which is smallest of the three, is rarely used in age and growth studies.

In addition, in most species the accretion of calcium carbonate and gelatinous matrix alternates on a daily cycle. It is therefore also possible to determine fish age in days. This latter information is often obtained under a microscope, and provides significant data to early life history studies.

By measuring the thickness of individual rings, it has been assumed (at least in some species) to estimate fish growth because fish growth is directly proportional to otolith growth. However, some studies disprove a direct link between body growth and otolith growth. At times of lower or zero body growth the otolith continues to accrete leading some researchers to believe the direct link is to metabolism, not growth per se. Otoliths, unlike scales, do not reabsorb during times of decreased energy making it even more useful tool to age a fish. Fish never stop growing entirely, though growth rate in mature fish is reduced. Rings corresponding to later parts of the life cycle tend to be closer together as a result.

Age and growth studies of fish are important for understanding such things as timing and magnitude of spawning, recruitment and habitat use, larval and juvenile duration, and population age structure
Age class structure
Age class structure in fisheries and wildlife management is a part of population assessment. Age can be determined by counting growth rings in fish scales, otoliths, cross-sections of fin spines for species with thick spines such as triggerfish, or teeth for a few species. Each method has its...

. Such knowledge is in turn important for designing appropriate fisheries
Fishery
Generally, a fishery is an entity engaged in raising or harvesting fish which is determined by some authority to be a fishery. According to the FAO, a fishery is typically defined in terms of the "people involved, species or type of fish, area of water or seabed, method of fishing, class of boats,...

 management policies.

Diet research

Since the compounds in fish otoliths are resistant to digestion
Digestion
Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones....

, they are found in the digestive tracts and scat
Scat
-Education:* School and College Ability Test* Somerset College of Arts and Technology, a community college in Somerset, England* Shrewsbury College of Arts & Technology, a community college in Shropshire, England-Games:* Thirty-one , a card game...

s of piscivorous marine mammals, such as dolphins, seals
Pinniped
Pinnipeds or fin-footed mammals are a widely distributed and diverse group of semiaquatic marine mammals comprising the families Odobenidae , Otariidae , and Phocidae .-Overview: Pinnipeds are typically sleek-bodied and barrel-shaped...

, sea lions and walruses. Many fish can be identified to genus
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

 and species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 by their sagittal otoliths. Otoliths can therefore, to some extent, be used to reconstruct the prey composition of marine mammal diets.


Sagittal otoliths (sagittae) are bilaterally symmetrical, with each fish having one right and one left. Separating recovered otoliths into right and left, therefore, allows one to infer a minimum number of prey individuals ingested for a given fish species. Otolith size is also proportional to the length and weight of a fish. They can therefore be used to back-calculate prey size and biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....

, useful when trying to estimate marine mammal
Marine mammal
Marine mammals, which include seals, whales, dolphins, and walruses, form a diverse group of 128 species that rely on the ocean for their existence. They do not represent a distinct biological grouping, but rather are unified by their reliance on the marine environment for feeding. The level of...

 prey consumption, and potential impacts on fish stocks.


Otoliths cannot be used alone to reliably estimate cetacean or pinniped
Pinniped
Pinnipeds or fin-footed mammals are a widely distributed and diverse group of semiaquatic marine mammals comprising the families Odobenidae , Otariidae , and Phocidae .-Overview: Pinnipeds are typically sleek-bodied and barrel-shaped...

 diets, however. They may suffer partial or complete erosion
Erosion
Erosion is when materials are removed from the surface and changed into something else. It only works by hydraulic actions and transport of solids in the natural environment, and leads to the deposition of these materials elsewhere...

 in the digestive tract, skewing measurements of prey number and biomass
Biomass
Biomass, as a renewable energy source, is biological material from living, or recently living organisms. As an energy source, biomass can either be used directly, or converted into other energy products such as biofuel....

. Species with fragile, easily digested otoliths may be underestimated in the diet. To address these biases, otolith correction factors have been developed through captive feeding experiments, in which seals are fed fish of known size, and the degree of otolith erosion is quantified for different prey taxa.


The inclusion of fish vertebrae, jaw bones, teeth, and other informative skeletal elements improves prey identification and quantification over otolith analysis alone. This is especially true for fish species with fragile otoliths, but other distinctive bones, such as Atlantic mackerel
Atlantic mackerel
The Atlantic mackerel , is a pelagic schooling species of mackerel found on both sides of the North Atlantic Ocean. The species is also called Boston mackerel, or just mackerel....

 (Scomber scombrus), and Atlantic herring
Atlantic herring
Atlantic herring is a fish in the family Clupeidae. It is one of the most abundant fish species on earth. Herring can be found on both sides of the Atlantic Ocean, congregating in large schools. They can grow up to in length and weigh more than...

(Clupea harengus).
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