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Deep-sea fish are fish that live in the
darkness Darkness is the condition resulting from a lack of illumination, or an absence of visible light. Human vision is unable to distinguish colors in conditions of very low luminance because the hue-sensitive photoreceptor cells on the retina a ...
below the sunlit surface waters, that is below the
epipelagic The photic zone (or euphotic zone, epipelagic zone, or sunlight zone) is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological ...
or
photic zone The photic zone (or euphotic zone, epipelagic zone, or sunlight zone) is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological ...
of the
sea A sea is a large body of salt water. There are particular seas and the sea. The sea commonly refers to the ocean, the interconnected body of seawaters that spans most of Earth. Particular seas are either marginal seas, second-order section ...
. The
lanternfish Lanternfish (or myctophids, from the Greek language, Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family (biology), family Myctophidae. One of two families in the order Myctophiformes, ...
is, by far, the most common deep-sea fish. Other
deep-sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...
fishes include the flashlight fish,
cookiecutter shark The cookiecutter shark (''Isistius brasiliensis''), also called the cigar shark, is a species of small squaliform shark in the family Dalatiidae. This shark lives in warm, oceanic waters worldwide, particularly near islands, and has been reco ...
, bristlemouths,
anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...
,
viperfish A viperfish is any species of marine fish in the genus ''Chauliodus''. Viperfish are mostly found in the mesopelagic zone and are characterized by long, needle-like teeth and hinged lower jaws. A typical viperfish grows to lengths of . Viperfis ...
, and some species of eelpout. Only about 2% of known marine species inhabit the
pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...
environment. This means that they live in the
water column The (oceanic) water column is a concept used in oceanography to describe the physical (temperature, salinity, light penetration) and chemical ( pH, dissolved oxygen, nutrient salts) characteristics of seawater at different depths for a defined ...
as opposed to the
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
organisms that live in or on the sea floor. Deep-sea organisms generally inhabit
bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
( deep) and
abyssopelagic The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. The word ''abyss'' comes from the Ancient Greek, Greek word (), meaning "bottomless". At depths of , this zone remains in perpetual darkness. It covers 83% of t ...
( deep) zones. However, characteristics of deep-sea organisms, such as
bioluminescence Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorgani ...
can be seen in the
mesopelagic The mesopelagic zone (Greek language, Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light ...
( deep) zone as well. The mesopelagic zone is the disphotic zone, meaning light there is minimal but still measurable. The oxygen minimum layer exists somewhere between a depth of depending on the place in the ocean. This area is also where nutrients are most abundant. The
bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
and
abyssopelagic The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. The word ''abyss'' comes from the Ancient Greek, Greek word (), meaning "bottomless". At depths of , this zone remains in perpetual darkness. It covers 83% of t ...
zones are aphotic, meaning that no light penetrates this area of the ocean. These zones make up about 75% of the inhabitable ocean space. The epipelagic zone (0–200 metres or 0–650 ft deep) is the area where light penetrates the water and photosynthesis occurs. This is also known as the photic zone. Because this typically extends only a few hundred meters below the water, the deep sea, about 90% of the ocean volume, is in darkness. The deep sea is also an extremely hostile environment, with temperatures that rarely exceed and fall as low as (with the exception of
hydrothermal vent Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hot ...
ecosystems that can exceed 350 °C, or 662 °F), low oxygen levels, and pressures between 20 and 1000  atm (2-100 
MPa MPA or mPa may refer to: Academia Academic degrees * Master of Performing Arts * Master of Professional Accountancy * Master of Public Administration * Master of Public Affairs Schools * Mesa Preparatory Academy * Morgan Park Academy * M ...
, 300–14,500 psi).


Evolution

It has been speculated that deep-sea ecosystems may have been inhospitable to vertebrate life prior to an increased influx of nutrients into the ocean during the
Late Jurassic The Late Jurassic is the third Epoch (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...
and Early Cretaceous following the rise of
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit. T ...
on land, which led to an increase in abyssal invertebrate life, allowing fish to in turn colonize these ecosystems. However, some modern deep-sea fish, such as
holocephali Holocephali (Sometimes spelled Holocephala; Romanization of Greek, Greek for "complete head" in reference to the fusion of Palatoquadrate, upper jaw with the rest of the skull) is a Subclass (biology), subclass of Chondrichthyes, cartilaginous fi ...
ans, are descendants of much older lineages, indicating that much earlier colonizations of the deep-sea by vertebrates may have occurred, although no fossil evidence of this is known. The earliest known records of deep-sea fish are
trace fossil A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contrast with body fossils, which are the fossilized remains of part ...
s of feeding and swimming behavior attributed to unidentified neoteleosts (referable to the ichnogenera ''
Piscichnus ''Piscichnus'' is an ichnogenus of representing trace fossils A trace fossil, also called an ichnofossil (; ), is a fossil record of biological activity by lifeforms, but not the preserved remains of the organism itself. Trace fossils contr ...
'' and '' Undichna''), from the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
(130 million-year-old) Palombini Shale of
Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...
, which is thought to have been deposited in the
abyssal plain An abyssal plain is an underwater plain on the deep ocean floor, usually found at depths between . Lying generally between the foot of a continental rise and a mid-ocean ridge, abyssal plains cover more than 50% of the Earth's surface. They ...
of the former Piemont-Liguria Ocean. Prior to the discovery of these fossils, there was no evidence for deep-sea bony fish older than 50 million years in the
Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...
. The Cretaceous origin for most modern deep-sea fish has been further affirmed with phylogenetic studies such as those of aulopiform fish, which indicate that many deep-sea lineages of these groups originated around this time. Although the records from the Palombini Shale represent the earliest records of deep-sea bony fish, formations that preserve deepwater
shark 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 ...
fossils are also known from later in the Cretaceous. These include the Northumberland Formation of Canada and similarly aged deposits in Angola, both of which preserve fossils of taxa such as hexanchids, chlamydoselachids, and catsharks, which are known from deepwater habitats today but rare in other formations of the time. Paleogene formations with fossil deep-sea shark teeth are known from New Zealand for the middle
Paleocene The Paleocene ( ), or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), ...
, and formations in Denmark, France, Austria, and Morocco during the Eocene. The Paratethys Sea still supported deepwater sharks and rays into the Miocene, which are preserved in formations in Hungary. During the Paleogene, some prominent formations that preserve well-articulated specimens of deep-sea bony fish are known. These include the Monte Solane lagerstatte of early
Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...
Italy, which preserves a
bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
habitat likely deposited under the sea, as well as the late Eocene Pabdeh Formation of
Iran Iran, officially the Islamic Republic of Iran (IRI) and also known as Persia, is a country in West Asia. It borders Iraq to the west, Turkey, Azerbaijan, and Armenia to the northwest, the Caspian Sea to the north, Turkmenistan to the nort ...
. The deep-sea environments preserved by both formations are apparent through their abundance of fossil stomiiform fish. Notable Neogene formations that preserve fossils of deep-sea bony fish are known from the Miocene of Italy, Japan, and California.


Environment

In the deep ocean, the waters extend far below the epipelagic zone, and support very different types of pelagic fishes adapted to living in these deeper zones. In deep water,
marine snow In the deep ocean, marine snow (also known as "ocean dandruff") is a continuous shower of mostly organic detritus falling from the upper layers of the water column. It is a significant means of exporting energy from the light-rich photic zone to ...
is a continuous shower of mostly organic
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
falling from the upper layers of the water column. Its origin lies in activities within the productive
photic zone The photic zone (or euphotic zone, epipelagic zone, or sunlight zone) is the uppermost layer of a body of water that receives sunlight, allowing phytoplankton to perform photosynthesis. It undergoes a series of physical, chemical, and biological ...
. Marine snow includes dead or dying
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
,
protist A protist ( ) or protoctist is any eukaryotic organism that is not an animal, land plant, or fungus. Protists do not form a natural group, or clade, but are a paraphyletic grouping of all descendants of the last eukaryotic common ancest ...
s (
diatom A diatom (Neo-Latin ''diatoma'') is any member of a large group comprising several Genus, genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's B ...
s), fecal matter, sand, soot and other inorganic dust. The "snowflakes" grow over time and may reach several centimetres in diameter, travelling for weeks before reaching the ocean floor. However, most organic components of marine snow are consumed by
microbe A microorganism, or microbe, is an organism of microscopic size, which may exist in its single-celled form or as a colony of cells. The possible existence of unseen microbial life was suspected from antiquity, with an early attestation in ...
s,
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
and other filter-feeding animals within the first of their journey, that is, within the epipelagic zone. In this way, marine snow may be considered the foundation of deep-sea
mesopelagic The mesopelagic zone (Greek language, Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light ...
and
benthic The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...
s: as sunlight cannot reach them, deep-sea organisms rely heavily on marine snow as an energy source. Since there is no light in the deep sea (aphotic), there is a lack of primary producers. Therefore, most organisms in the bathypelagic rely on the marine snow from regions higher in the vertical column. Some deep-sea pelagic groups, such as the
lanternfish Lanternfish (or myctophids, from the Greek language, Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family (biology), family Myctophidae. One of two families in the order Myctophiformes, ...
,
ridgehead Ridgeheads, also known as bigscales, are a family (Melamphaidae, from the Greek ''melanos'' lackand ''amphi'' y both sides of small, deep-sea beryciform fish. The family contains approximately 37 species in five genera; their distribution is ...
,
marine hatchetfish Marine hatchetfishes or deep-sea hatchetfishes are small deep-sea mesopelagic ray-finned fish of the stomiiform subfamily Sternoptychinae. They should not be confused with the freshwater hatchetfishes, which are not particularly closely relate ...
, and lightfish families, are sometimes termed ''pseudoceanic'' because, rather than having an even distribution in open water, they occur in significantly higher abundances around structural oases, notably
seamount A seamount is a large submarine landform that rises from the ocean floor without reaching the water surface (sea level), and thus is not an island, islet, or cliff-rock. Seamounts are typically formed from extinct volcanoes that rise abruptly a ...
s and over
continental slope A continental margin is the outer edge of continental crust abutting oceanic crust under coastal waters. It is one of the three major zones of the ocean floor, the other two being deep-ocean basins and mid-ocean ridges. The continental margi ...
s. The phenomenon is explained by the likewise abundance of prey species which are also attracted to the structures. Hydrostatic pressure increases by 1 atm (0.1 MPa) for every in depth. Deep-sea organisms have the same pressure within their bodies as is exerted on them from the outside, so they are not crushed by the extreme pressure. Their high internal pressure, however, results in the reduced fluidity of their membranes because molecules are squeezed together. Fluidity in cell membranes increases efficiency of biological functions, most importantly the production of proteins, so organisms have adapted to this circumstance by increasing the proportion of unsaturated fatty acids in the lipids of the cell membranes. In addition to differences in internal pressure, these organisms have developed a different balance between their metabolic reactions from those organisms that live in the epipelagic zone. David Wharton, author of ''Life at the Limits: Organisms in Extreme Environments'', notes "Biochemical reactions are accompanied by changes in volume. If a reaction results in an increase in volume, it will be inhibited by pressure, whereas, if it is associated with a decrease in volume, it will be enhanced". This means that their metabolic processes must ultimately decrease the volume of the organism to some degree. Most fish that have evolved in this harsh environment are not capable of surviving in laboratory conditions, and attempts to keep them in captivity have led to their deaths. Deep-sea organisms contain gas-filled spaces (vacuoles). Gas is compressed under high pressure and expands under low pressure. Because of this, these organisms have been known to blow up if they come to the surface.


Characteristics

The fish of the deep-sea have evolved various adaptations to survive in this region. Since many of these fish live in regions where there is no natural illumination, they cannot rely solely on their eyesight for locating prey and mates and avoiding predators; deep-sea fish have
evolved Evolution is the change in the heritable Phenotypic trait, characteristics of biological populations over successive generations. It occurs when evolutionary processes such as natural selection and genetic drift act on genetic variation, re ...
appropriately to the extreme sub-photic region in which they live. Many of these organisms are blind and rely on their other senses, such as sensitivities to changes in local pressure and smell, to catch their food and avoid being caught. Those that aren't blind have large and sensitive eyes that can use
bioluminescent Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some fungi, microorganisms inc ...
light. These eyes can be as much as 100 times more sensitive to light than human eyes.
Rhodopsin Rhodopsin, also known as visual purple, is a protein encoded by the ''RHO'' gene and a G-protein-coupled receptor (GPCR). It is a light-sensitive receptor protein that triggers visual phototransduction in rod cells. Rhodopsin mediates dim ...
(Rh1) is a protein found in the eye's rod cells that helps animals see in dim light. While most vertebrates usually have one Rh1 opsin gene, some deep-sea fish have several Rh1 genes, and one species, the
silver spinyfin The silver spinyfin (''Diretmus argenteus'') is a spinyfin of the genus ''Diretmus'', found around the world except the Mediterranean, at depths down to 2,000 m. It belongs to the monotypic genus ''Diretmus''. Their length is between 30 and 40 ...
(''Diretmus argenteus''), has 38. This proliferation of Rh1 genes may help deep-sea fish to see in the depths of the ocean. Also, to avoid predation, many species are dark to blend in with their environment. Many deep-sea fish are
bioluminescent Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some fungi, microorganisms inc ...
, with extremely large eyes adapted to the dark. Bioluminescent organisms are capable of producing light biologically through the agitation of molecules of luciferin, which then produce light. This process must be done in the presence of oxygen. These organisms are common in the mesopelagic region and below ( and below). More than 50% of deep-sea fish, as well as some species of shrimp and squid, are capable of bioluminescence. About 80% of these organisms have photophores – light producing glandular cells that contain luminous bacteria bordered by dark colourings. Some of these photophores contain lenses, much like those in the eyes of humans, which can intensify or lessen the emanation of light. The ability to produce light only requires 1% of the organism's energy and has many purposes: It is used to search for food and attract prey, like the anglerfish; claim territory through patrol; communicate and find a mate, and distract or temporarily blind predators to escape. Also, in the mesopelagic where some light still penetrates, some organisms camouflage themselves from predators below them by illuminating their bellies to match the colour and intensity of light from above so that no shadow is cast. This tactic is known as counter-illumination. The lifecycle of deep-sea fish can be exclusively deep-water, although some species are born in shallower water and sink upon maturation. Regardless of the depth where eggs and larvae reside, they are typically pelagic. This planktonic — drifting — lifestyle requires neutral buoyancy. In order to maintain this, the eggs and larvae often contain oil droplets in their plasma. When these organisms are in their fully matured state they need other adaptations to maintain their positions in the water column. In general, water's density causes upthrust — the aspect of buoyancy that makes organisms float. To counteract this, the density of an organism must be greater than that of the surrounding water. Most animal tissues are denser than water, so they must find an equilibrium to make them float. Many organisms develop
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 ...
s (gas cavities) to stay afloat, but because of the high pressure of their environment, deep-sea fishes usually do not have this organ. Instead they exhibit structures similar to hydrofoils in order to provide hydrodynamic lift. It has also been found that the deeper a fish lives, the more jelly-like its flesh and the more minimal its bone structure. They reduce their tissue density through high fat content, reduction of skeletal weight — accomplished through reductions of size, thickness and mineral content — and water accumulation makes them slower and less agile than surface fish. The body shapes of deep-sea fish are generally better adapted for periodic bursts of swimming rather than constant swimming. Due to the poor level of
photosynthetic Photosynthesis ( ) is a Biological system, system of biological processes by which Photoautotrophism, photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical ener ...
light reaching deep-sea environments, most fish need to rely on organic matter sinking from higher levels, or, in rare cases,
hydrothermal vents Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hots ...
for nutrients. This makes the deep-sea much poorer in
productivity Productivity is the efficiency of production of goods or services expressed by some measure. Measurements of productivity are often expressed as a ratio of an aggregate output to a single input or an aggregate input used in a production proce ...
than shallower regions. Also, animals in the pelagic environment are sparse and food doesn't come along frequently. Because of this, organisms need adaptations that allow them to survive. Some have long feelers to help them locate prey or attract mates in the pitch black of the deep ocean. The deep-sea angler fish in particular has a long fishing-rod-like adaptation protruding from its face, on the end of which is a bioluminescent piece of skin that wriggles like a worm to lure its prey. Some must consume other fish that are the same size or larger than them and they need adaptations to help digest them efficiently. Great sharp teeth, hinged jaws, disproportionately large mouths, and expandable bodies are a few of the characteristics that deep-sea fishes have for this purpose. The gulper eel is one example of an organism that displays these characteristics. Fish in the different pelagic and deep-water benthic zones are physically structured, and behave in ways, that differ markedly from each other. Groups of coexisting species within each zone all seem to operate in similar ways, such as the small mesopelagic vertically migrating plankton-feeders, the bathypelagic
anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...
es, and the deep-water benthic rattails. Ray finned species, with spiny fins, are rare among deep-sea fishes, which suggests that deep-sea fish are ancient and so well adapted to their environment that invasions by more modern fishes have been unsuccessful. The few ray fins that do exist are mainly in the
Beryciformes The Beryciformes are a poorly-understood Order (biology), order of carnivorous ray-finned fishes consisting of 7 families, 30 genera, and 161 species. They feed on small fish and invertebrates. Beyond this, little is known about the biology of m ...
and
Lampriformes Lampriformes is an order (biology), order of Actinopterygii, ray-finned fish. Members are collectively called lamprids (which is more properly used for the Opah, Lampridae) or lampriforms, and unite such open-ocean and partially Deep sea, deep-s ...
, which are also ancient forms. Most deep-sea pelagic fishes belong to their own orders, suggesting a long evolution in deep-sea environments. In contrast, deep-water benthic species, are in orders that include many related shallow-water fishes.


Mesopelagic fish

Below the epipelagic zone, conditions change rapidly. Between 200 m and about 1000 m, light continues to fade until there is almost none. Temperatures fall through a
thermocline A thermocline (also known as the thermal layer or the metalimnion in lakes) is a distinct layer based on temperature within a large body of fluid (e.g. water, as in an ocean or lake; or air, e.g. an atmosphere) with a high gradient of distinct te ...
to temperatures between . This is the twilight or
mesopelagic The mesopelagic zone (Greek language, Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light ...
zone. Pressure continues to increase, at the rate of one atm (0.1 MPa) every , while nutrient concentrations fall, along with dissolved oxygen and the rate at which the water circulates.
Sonar Sonar (sound navigation and ranging or sonic navigation and ranging) is a technique that uses sound propagation (usually underwater, as in submarine navigation) to navigate, measure distances ( ranging), communicate with or detect objects o ...
operators, using the newly developed sonar technology during , were puzzled by what appeared to be a false sea floor deep by day, and less deep at night. This turned out to be due to millions of marine organisms, most particularly small mesopelagic fish, with swim bladders that reflected the sonar. These organisms migrate up into shallower water at dusk to feed on plankton. The layer is deeper when the moon is out, and can become shallower when clouds pass over the moon. This phenomenon has come to be known as the deep scattering layer.Ryan P.
"Deep-sea creatures: The mesopelagic zone"
''Te Ara – the Encyclopedia of New Zealand''. Updated 21 September 2007.
Most mesopelagic fish make daily vertical migrations, moving at night into the epipelagic zone, often following similar migrations of zooplankton, and returning to the depths for safety during the day.Bone & Moore 2008, p. 38. These vertical migrations often occur over large vertical distances, and are undertaken with the assistance of a
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 ...
.Haedrich R. L. (1996
"Deep-water fishes: evolution and adaptation in the earth's largest living spaces"
''Journal of Fish Biology'', 49(sA):40–53.
The swim bladder is inflated when the fish wants to move up, and, given the high pressures in the messoplegic zone, this requires significant energy. As the fish ascends, the pressure in the swim bladder must adjust to prevent it from bursting. When the fish wants to return to the depths, the swim bladder is deflated. Some mesopelagic fishes make daily migrations through the
thermocline A thermocline (also known as the thermal layer or the metalimnion in lakes) is a distinct layer based on temperature within a large body of fluid (e.g. water, as in an ocean or lake; or air, e.g. an atmosphere) with a high gradient of distinct te ...
, where the temperature changes between , thus displaying considerable tolerances for temperature change. These fish have muscular bodies, ossified bones, scales, well developed gills and central nervous systems, and large hearts and kidneys. Mesopelagic plankton feeders have small mouths with fine
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
s, while the
piscivore A piscivore () is a carnivorous animal that primarily eats fish. Fish were the diet of early tetrapod evolution (via water-bound amphibians during the Devonian period); insectivory came next; then in time, the more terrestrially adapted repti ...
s have larger mouths and coarser gill rakers. Mesopelagic fish are adapted for an active life under low light conditions. Most of them are visual predators with large eyes. Some of the deeper water fish have tubular eyes with big lenses and only
rod cell Rod cells are photoreceptor cells in the retina of the eye that can function in lower light better than the other type of visual photoreceptor, cone cells. Rods are usually found concentrated at the outer edges of the retina and are used in ...
s that look upwards. These give binocular vision and great sensitivity to small light signals. This adaptation gives improved terminal vision at the expense of lateral vision, and allows the predator to pick out
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
,
cuttlefish Cuttlefish, or cuttles, are Marine (ocean), marine Mollusca, molluscs of the order (biology), suborder Sepiina. They belong to the class (biology), class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique ...
, and smaller fish that are silhouetted against the gloom above them. Mesopelagic fish usually lack defensive spines, and use colour to
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
themselves from other fish.
Ambush predator Ambush predators or sit-and-wait predators are carnivorous animals that capture their prey via stealth, luring or by (typically instinctive) strategies utilizing an element of surprise. Unlike pursuit predators, who chase to capture prey u ...
s are dark, black or red. Since the longer, red, wavelengths of light do not reach the deep sea, red effectively functions the same as black. Migratory forms use countershaded silvery colours. On their bellies, they often display
photophore A photophore is a specialized anatomical structure found in a variety of organisms that emits light through the process of boluminescence. This light may be produced endogenously by the organism itself (symbiotic) or generated through a mut ...
s producing low grade light. For a predator from below, looking upwards, this
bioluminescence Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorgani ...
camouflages the silhouette of the fish. However, some of these predators have yellow lenses that filter the (red deficient) ambient light, leaving the bioluminescence visible. The brownsnout spookfish, a species of barreleye, is the only vertebrate known to employ a mirror, as opposed to a lens, to focus an image in its eyes.Smith, L. (Jan. 8, 2009)
"Fish with four eyes can see through the deep sea gloom"
''Times Online''. Times Newspapers Ltd. Retrieved on March 14, 2009.
Sampling by deep
trawling Trawling is an industrial method of fishing that involves pulling a fishing net through the water behind one or more boats. The net used for trawling is called a trawl. This principle requires netting bags which are towed through water to catch di ...
indicates that
lanternfish Lanternfish (or myctophids, from the Greek language, Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family (biology), family Myctophidae. One of two families in the order Myctophiformes, ...
account for as much as 65% of all deep-sea fish
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
. Indeed, lanternfish are among the most widely distributed, populous, and diverse of all
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, playing an important
ecological Ecology () is the natural science of the relationships among living organisms and their environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere levels. Ecology overlaps with the closely re ...
role as prey for larger organisms. The estimated global biomass of lanternfish is 550–660 million
tonne The tonne ( or ; symbol: t) is a unit of mass equal to 1,000  kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton in the United States to distinguish it from the non-metric units of the s ...
s, several times the entire world fisheries catch. Lanternfish also account for much of the biomass responsible for the deep scattering layer of the world's oceans.
Bigeye tuna The bigeye tuna (''Thunnus obesus'') is a species of true tuna of the genus ''Thunnus'', belonging to the wider mackerel family (biology), family Scombridae. In Hawaiian language, Hawaiian, it is one of two species known as ahi, the other being t ...
are an epipelagic/mesopelagic species that eats other fish. Satellite tagging has shown that bigeye tuna often spend prolonged periods cruising deep below the surface during the daytime, sometimes making dives as deep as . These movements are thought to be in response to the vertical migrations of prey organisms in the deep scattering layer. File:Malacosteus niger.jpg, The stoplight loosejaw has a
lower jaw In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
one-quarter as long as its body. The jaw has no floor and is attached only by a hinge and a modified tongue bone. Large fang-like teeth in the front are followed by many small barbed teeth. File:Malacosteus.JPG, The stoplight loosejaw is also one of the few fishes that produce red
bioluminescence Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorgani ...
. As most of their prey cannot perceive red light, this allows it to hunt with an essentially invisible beam of light. File:Longnoselancetfish.jpg, Long-snouted lancetfish. Lancetfish are ambush predators which spend all their time in the mesopelagic zone. They are among the largest mesopelagic fishes (up to . File:Daggertooth.PNG, The daggertooth paralyses other mesopelagic fish when it bites them with its dagger-like teeth


Bathypelagic fish

Below the mesopelagic zone it is pitch dark. This is the midnight (or
bathypelagic zone The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
), extending from to the bottom deep-water
benthic zone The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from the Ancient Greek word (), meaning "the depths". ...
. If the water is exceptionally deep, the pelagic zone below is sometimes called the lower midnight (or abyssopelagic zone). Temperatures in this zone range from and it is completely aphotic. Conditions are somewhat uniform throughout these zones; the darkness is complete, the pressure is crushing, and temperatures, nutrients and dissolved oxygen levels are all low. Bathypelagic fish have special
adaptation In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
s to cope with these conditions – they have slow
metabolism Metabolism (, from ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run cellular processes; the co ...
s and unspecialized diets, being willing to eat anything that comes along. They prefer to sit and wait for food rather than waste energy searching for it. The behaviour of bathypelagic fish can be contrasted with the behaviour of mesopelagic fish. Mesopelagic fish are often highly mobile, whereas bathypelagic fish are almost all lie-in-wait predators, normally expending little energy in movement. The dominant bathypelagic fishes are small bristlemouth and
anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...
; fangtooth,
viperfish A viperfish is any species of marine fish in the genus ''Chauliodus''. Viperfish are mostly found in the mesopelagic zone and are characterized by long, needle-like teeth and hinged lower jaws. A typical viperfish grows to lengths of . Viperfis ...
, daggertooth and
barracudina Barracudinas are any member of the marine mesopelagic fish family Paralepididae: 50 or so extant species are found almost worldwide in deep waters. Several genera, including '' Holosteus'' and '' Drimys'', are known only from fossils dating b ...
are also common. These fishes are small, many about long, and not many longer than . They spend most of their time waiting patiently in the water column for prey to appear or to be lured by their phosphors. What little energy is available in the bathypelagic zone filters from above in the form of detritus, faecal material, and the occasional invertebrate or mesopelagic fish. About 20 percent of the food that has its origins in the epipelagic zone falls down to the mesopelagic zone, but only about 5 percent filters down to the bathypelagic zone.Ryan
"Deep-sea creatures: The bathypelagic zone"
''Te Ara - the Encyclopedia of New Zealand''. Updated 21 September 2007.
Bathypelagic fish are sedentary, adapted to outputting minimum energy in a habitat with very little food or available energy, not even sunlight, only bioluminescence. Their bodies are elongated with weak, watery muscles and
skeletal A skeleton is the structural frame that supports the body of most animals. There are several types of skeletons, including the exoskeleton, which is a rigid outer shell that holds up an organism's shape; the endoskeleton, a rigid internal fram ...
structures. Since so much of the fish is water, they are not compressed by the great pressures at these depths. They often have extensible, hinged jaws with recurved teeth. They are slimy, without scales. The central nervous system is confined to the lateral line and olfactory systems, the eyes are small and may not function, and
gill A gill () is a respiration organ, respiratory organ that many aquatic ecosystem, 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 r ...
s, kidneys and hearts, and
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 ...
s are small or missing. These are the same features found in fish
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
, which suggests that during their evolution, bathypelagic fish have acquired these features through
neoteny Neoteny (), also called juvenilization,Montagu, A. (1989). Growing Young. Bergin & Garvey: CT. is the delaying or slowing of the Physiology, physiological, or Somatic (biology), somatic, development of an organism, typically an animal. Neoteny i ...
. As with larvae, these features allow the fish to remain suspended in the water with little expenditure of energy.Marshall (1984) "Progenetic tendencies in deep-sea fishes", pp. 91-101 in Potts GW and Wootton RJ (eds.) (1984) ''Fish reproduction: strategies and tactics'' Fisheries Society of the British Isles. Despite their ferocious appearance, these forms are mostly miniature fish with weak muscles, and are too small to represent any threat to humans. An exception to the generally small size of bathypelagic fish is the Yokozuna slickhead (''Narcetes shonanmaruae''), described in 2021, which is the largest known entirely bathypelagic
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
at over in length. The swim bladders of deep-sea fish are either absent or scarcely operational, and bathypelagic fish do not normally undertake vertical migrations. Filling bladders at such great pressures incurs huge energy costs. Some deep-sea fishes have swim bladders which function while they are young and inhabit the upper epipelagic zone, but they wither or fill with fat when the fish move down to their adult habitat. A couple of known exceptions are the cusk-eel ('' Holcomycteronus profundissimus''), retrieved from 7160 meters deep, and the rough abyssal grenadier ('' Coryphaenoides yaquinae''), found at 7259 meters depth. These species still have functional swim bladders, which allows them to maintain high bone density and strong jaws. The most important sensory systems are usually the
inner ear The inner ear (internal ear, auris interna) is the innermost part of the vertebrate ear. In vertebrates, the inner ear is mainly responsible for sound detection and balance. In mammals, it consists of the bony labyrinth, a hollow cavity in the ...
, which responds to sound, and the
lateral line 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 epithelia ...
, which responds to changes in water pressure. The
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
system can also be important for males who find females by smell. Bathypelagic fish are black, or sometimes red, with few
photophore A photophore is a specialized anatomical structure found in a variety of organisms that emits light through the process of boluminescence. This light may be produced endogenously by the organism itself (symbiotic) or generated through a mut ...
s. When photophores are used, it is usually to entice prey or attract a mate. Because food is so scarce, bathypelagic predators are not selective in their feeding habits, but grab whatever comes close enough. They accomplish this by having a large mouth with sharp teeth for grabbing large prey and overlapping
gill raker Gill rakers in fish are bony or cartilaginous processes that project from the branchial arch (gill arch) and are involved with suspension feeding tiny prey. They are not to be confused with the gill filaments that compose the fleshy part of th ...
s which prevent small prey that have been swallowed from escaping. It is not easy finding a mate in this zone. Some species depend on
bioluminescence Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some Fungus, fungi, microorgani ...
, where bioluminescent patterns are unique to specific species. Others are
hermaphrodite A hermaphrodite () is a sexually reproducing organism that produces both male and female gametes. Animal species in which individuals are either male or female are gonochoric, which is the opposite of hermaphroditic. The individuals of many ...
s, which doubles their chances of producing both eggs and sperm when an encounter occurs. The female anglerfish releases
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s to attract tiny males. When a male finds her, he bites on to her and never lets go. When a male of the anglerfish species '' Haplophryne mollis'' bites into the skin of a female, he releases an
enzyme An enzyme () is a protein that acts as a biological catalyst by accelerating chemical reactions. The molecules upon which enzymes may act are called substrate (chemistry), substrates, and the enzyme converts the substrates into different mol ...
that digests the skin of his mouth and her body, fusing the pair to the point where the two circulatory systems join up. The male then atrophies into nothing more than a pair of
gonads A gonad, sex gland, or reproductive gland is a mixed gland and sex organ that produces the gametes and sex hormones of an organism. Female reproductive cells are egg cells, and male reproductive cells are sperm. The male gonad, the testicle, ...
. This extreme
sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
ensures that, when the female is ready to spawn, she has a mate immediately available. Many forms other than fish live in the bathypelagic zone, such as squid, large whales, octopuses, sponges,
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear e ...
s, sea stars, and echinoids, but this zone is difficult for fish to live in. File:Eurypharynx pelecanoides.jpg, The pelican eel uses its large mouth like a net by opening its jaws and swimming towards prey. It has a luminescent organ at the tip of its tail to attract prey. File:Chiasmodon niger.jpg, The black swallower, with its distensible stomach, is notable for its ability to swallow, whole,
bony fish Osteichthyes ( ; ), also known as osteichthyans or commonly referred to as the bony fish, is a Biodiversity, diverse clade of vertebrate animals that have endoskeletons primarily composed of bone tissue. They can be contrasted with the Chondricht ...
es ten times its mass. File:Hamol u0.gif, Female '' Haplophryne mollis'' anglerfish trailing attached males which have atrophied into a pair of gonads, for use when the female is ready to spawn. File:Narcetes shonanmaruae a frozen specimen.jpg, The Yokozuna slickhead is the largest known bathypelagic bony fish, at over in length.


Adaptation to high pressure

As a fish moves deeper into the sea, the weight of the water overhead exerts increasing
hydrostatic pressure Hydrostatics is the branch of fluid mechanics that studies fluids at hydrostatic equilibrium and "the pressure in a fluid or exerted by a fluid on an immersed body". The word "hydrostatics" is sometimes used to refer specifically to water and o ...
on the fish. This increased pressure amounts to about one atm (0.1 MPa) for every in depth. For a fish at the bottom of the
bathypelagic zone The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...
, this pressure amounts to about 400 atm (40 MPa, 6000 psi). Deep-sea organisms possess adaptations at cellular and physiological levels that allow them to survive in environments of great pressure. Not having these adaptations limits the depths at which shallow-water species can operate. High levels of external pressure affects how metabolic processes and biochemical reactions proceed. The equilibrium of many chemical reactions is disturbed by pressure, and pressure can inhibit processes which result in an increase in volume. Water, a key component in many biological processes, is very susceptible to volume changes, mainly because constituents of cellular fluid have an effect on water structure. Thus, enzymatic reactions that induce changes in water organization effectively change the system's volume. Proteins responsible for catalyzing reactions are typically held together by weak bonds and the reactions usually involve volume increases. Species that can tolerate these depths have evolved changes in their protein structure and reaction criteria to withstand pressure, in order to perform reactions in these conditions. In high pressure environments, bilayer cellular membranes experience a loss of fluidity. Deep-sea cellular membranes favor
phospholipid bilayer The lipid bilayer (or phospholipid bilayer) is a thin polar membrane made of two layers of lipid molecules. These membranes form a continuous barrier around all cells. The cell membranes of almost all organisms and many viruses are made of a l ...
s with a higher proportion of
unsaturated fatty acid In chemistry, in particular in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
s, which induce a higher fluidity than their sea-level counterparts. Ten orders, thirteen families and about 200 known species of deep-sea fish have evolved a gelatinous layer below the skin or around the spine, which is used for buoyancy, low-cost growth and to increase swimming efficiency by reducing drag. Deep-sea species exhibit lower changes of
entropy Entropy is a scientific concept, most commonly associated with states of disorder, randomness, or uncertainty. The term and the concept are used in diverse fields, from classical thermodynamics, where it was first recognized, to the micros ...
and
enthalpy Enthalpy () is the sum of a thermodynamic system's internal energy and the product of its pressure and volume. It is a state function in thermodynamics used in many measurements in chemical, biological, and physical systems at a constant extern ...
compared to surface level organisms, since a high pressure and low temperature environment favors negative enthalpy changes and reduced dependence on entropy-driven reactions. From a structural standpoint,
globular protein In biochemistry, globular proteins or spheroproteins are spherical ("globe-like") proteins and are one of the common protein types (the others being fibrous, disordered and membrane proteins). Globular proteins are somewhat water-soluble (form ...
s of deep-sea fish due to the tertiary structure of G-actin are relatively rigid compared to those of surface level fish. The fact that proteins in deep-sea fish are structurally different from surface fish is apparent from the observation that actin from the muscle fibers of deep-sea fish are extremely heat-resistant; an observation similar to what is found in lizards. These proteins are structurally strengthened by modification of the bonds in the tertiary structure of the protein which also happens to induce high levels of thermal stability. Proteins are structurally strengthened to resist pressure by modification of bonds in the tertiary structure. Therefore, high levels of hydrostatic pressure, similar to high body temperatures of thermophilic desert reptiles, favor rigid protein structures. Na+/K+ -ATPase is a
lipoprotein A lipoprotein is a biochemical assembly whose primary function is to transport hydrophobic lipid (also known as fat) molecules in water, as in blood plasma or other extracellular fluids. They consist of a triglyceride and cholesterol center, sur ...
enzyme that plays a prominent role in
osmoregulation Osmoregulation is the active regulation of the osmotic pressure of an organism's body fluids, detected by osmoreceptors, to maintain the homeostasis of the organism's water content; that is, it maintains the fluid balance and the concentration ...
and is heavily influenced by hydrostatic pressure. The inhibition of Na+/K+ -ATPase is due to increased compression due to pressure. The rate-limiting step of the Na+/K+ -ATPase reaction induces an expansion in the bilayer surrounding the protein, and therefore an increase in volume. An increase in volume makes Na+/K+ -ATPase reactivity susceptible to higher pressures. Even though the Na+/K+ -ATPase activity per gram of gill tissue is lower for deep-sea fishes, the Na+/K+ -ATPases of deep-sea fishes exhibit a much higher tolerance of hydrostatic pressure compared to their shallow-water counterparts. This is exemplified between the species ''C. acrolepis'' (around 2000 m deep) and its hadalpelagic counterpart '' C. armatus'' (around deep), where the Na+/K+ -ATPases of ''C. armatus'' are much less sensitive to pressure. This resistance to pressure can be explained by adaptations in the protein and lipid moieties of Na+/K+ -ATPase.


Lanternfish

Sampling via deep trawling indicates that
lanternfish Lanternfish (or myctophids, from the Greek language, Greek μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large family (biology), family Myctophidae. One of two families in the order Myctophiformes, ...
account for as much as 65% of all deep-sea fish
biomass Biomass is a term used in several contexts: in the context of ecology it means living organisms, and in the context of bioenergy it means matter from recently living (but now dead) organisms. In the latter context, there are variations in how ...
. Indeed, lanternfish are among the most widely distributed, populous, and diverse of all
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, playing an important ecological role as prey for larger organisms. With an estimated global biomass of 550–660 million
metric ton The tonne ( or ; symbol: t) is a unit of mass equal to 1,000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton in the United States to distinguish it from the non-metric units of the sh ...
s, several times the entire world fisheries catch, lanternfish also account for much of the biomass responsible for the deep scattering layer of the world's oceans. In the
Southern Ocean The Southern Ocean, also known as the Antarctic Ocean, comprises the southernmost waters of the world ocean, generally taken to be south of 60th parallel south, 60° S latitude and encircling Antarctica. With a size of , it is the seco ...
, Myctophids provide an alternative food resource to
krill Krill ''(Euphausiids)'' (: krill) are small and exclusively marine crustaceans of the order (biology), order Euphausiacea, found in all of the world's oceans. The name "krill" comes from the Norwegian language, Norwegian word ', meaning "small ...
for predators such as
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
and the
king penguin The king penguin (''Aptenodytes patagonicus'') is the second largest species of penguin, smaller than but somewhat similar in appearance to the emperor penguin. King penguins mainly eat lanternfish, squid, and krill. On foraging trips, king pen ...
. Although these fish are plentiful and prolific, currently only a few commercial lanternfish fisheries exist: these include limited operations off
South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...
, in the
sub-Antarctic The sub-Antarctic zone is a physiographic region in the Southern Hemisphere, located immediately north of the Antarctic region. This translates roughly to a latitude of between 46° and 60° south of the Equator. The subantarctic region inc ...
, and in the
Gulf of Oman The Gulf of Oman or Sea of Oman ( ''khalīj ʿumān''; ''daryâ-ye omân''), also known as Gulf of Makran or Sea of Makran ( ''khalīj makrān''; ''daryâ-ye makrān''), is a gulf in the Indian Ocean that connects the Arabian Sea with th ...
.


Endangered species

A 2006 study by Canadian scientists has found five species of deep-sea fish – blue hake, spiny eel – to be on the verge of extinction due to the shift of
commercial fishing Commercial fishing is the activity of catching fish and other seafood for Commerce, commercial Profit (economics), profit, mostly from wild fisheries. It provides a large quantity of food to many countries around the world, but those who practice ...
from
continental shelves A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
to the slopes of the
continental shelves A continental shelf is a portion of a continent that is submerged under an area of relatively shallow water, known as a shelf sea. Much of these shelves were exposed by drops in sea level during glacial periods. The shelf surrounding an island ...
, down to depths of . The slow reproduction of these fish – they reach
sexual maturity Sexual maturity is the capability of an organism to reproduce. In humans, it is related to both puberty and adulthood. ''Puberty'' is the biological process of sexual maturation, while ''adulthood'', the condition of being socially recognized ...
at about the same age as human beings – is one of the main reasons that they cannot recover from the excessive fishing.


See also

*
Census of Marine Life The Census of Marine Life was a scientific initiative involving a global network of researchers in more than 80 nations, engaged to assess and explain the diversity, distribution, and abundance of life in the oceans. The census cost US$650 milli ...
* Deep ocean water *
Deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...
* Deep-sea community *
Demersal fish Demersal fish, also known as groundfish, live and feed on or near the bottom of seas or lakes (the demersal zone).Walrond Carl . "Coastal fish - Fish of the open sea floor"Te Ara - the Encyclopedia of New Zealand. Updated 2 March 2009 They oc ...
*
Pelagic fish Pelagic fish live in the pelagic zone of ocean or lake waters—being neither close to the bottom nor near the shore—in contrast with demersal fish that live on or near the bottom, and reef fish that are associated with coral reefs. ...


Citations


References

* * * * * * *


Further reading

* Gordon J. D. M. (2001
"Deep-sea fishes"
In: John H. Steele, Steve A. Thorpe, Karl K. Turekian (eds.) ''Elements of Physical Oceanography'', pages 227–233, Academic Press. . * Hoar W. S., Randall D. J. and Farrell A. P. (eds.) (1997
''Deep-Sea Fishes''
Academic Press. . * Shotton, Ross (1995

In: ''Review of the state of world marine fishery resources'', FAO Fisheries technical paper 457, FAO, Rome. . * Tandstad M., Shotton R., Sanders J. and Carocci F. (2011
"Deep-sea Fisheries"
In: ''Review of the state of world marine fishery resources'', pages 265–278, FAO Fisheries technical paper 569, FAO, Rome. .


External links




Photo Gallery: Deep-Sea Creatures

Crash Course To Deep Sea Fishing
– articles, facts and images of deep-sea animals {{DEFAULTSORT:Deep Sea Fish Deep sea fish Articles containing video clips Oceanographical terminology Ichthyology