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Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the
subphylum In zoological nomenclature, a subphylum is a taxonomic rank below the rank of phylum. The taxonomic rank of " subdivision" in fungi and plant taxonomy is equivalent to "subphylum" in zoological taxonomy. Some plant taxonomists have also used t ...
Medusozoa, a major part of the
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclatu ...
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter. Their distinguishing feature is cnidocyt ...
. Jellyfish are mainly free-swimming marine
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s with umbrella-shaped bells and trailing
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
s, although a few are anchored to the seabed by stalks rather than being mobile. The bell can pulsate to provide propulsion for highly efficient locomotion. The tentacles are armed with stinging cells and may be used to capture prey and defend against predators. Jellyfish have a complex
life cycle Life cycle, life-cycle, or lifecycle may refer to: Science and academia * Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring * Life-cycle hypothesi ...
; the medusa is normally the sexual phase, which produces
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, whi ...
larvae that disperse widely and enter a sedentary polyp phase before reaching sexual maturity. Jellyfish are found all over the world, from surface waters to the deep sea. Scyphozoans (the "true jellyfish") are exclusively marine, but some
hydrozoa Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specializ ...
ns with a similar appearance live in
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does in ...
. Large, often colorful, jellyfish are common in coastal zones worldwide. The medusae of most species are fast-growing, and mature within a few months then die soon after breeding, but the polyp stage, attached to the seabed, may be much more long-lived. Jellyfish have been in existence for at least 500 million years, and possibly 700 million years or more, making them the oldest multi-organ animal group. Jellyfish are eaten by humans in certain cultures. They are considered a delicacy in some Asian countries, where species in the Rhizostomae
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
are pressed and salted to remove excess water. Australian researchers have described them as a "perfect food", sustainable, and protein-rich but relatively low in
food energy Food energy is chemical energy that animals (including humans) derive from their food to sustain their metabolism, including their muscular activity. Most animals derive most of their energy from aerobic respiration, namely combining the carbohy ...
. They are also used in research, where the
green fluorescent protein The green fluorescent protein (GFP) is a protein that exhibits bright green fluorescence when exposed to light in the blue to ultraviolet range. The label ''GFP'' traditionally refers to the protein first isolated from the jellyfish '' Aeq ...
used by some species to cause
bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some ...
has been adapted as a fluorescent
marker The term Marker may refer to: Common uses * Marker (linguistics), a morpheme that indicates some grammatical function * Marker (telecommunications), a special-purpose computer * Boundary marker, an object that identifies a land boundary * Marke ...
for genes inserted into other cells or organisms. The stinging cells used by jellyfish to subdue their prey can injure humans. Thousands of swimmers worldwide are stung every year, with effects ranging from mild discomfort to serious injury or even death. When conditions are favourable, jellyfish can form vast swarms, which can be responsible for damage to fishing gear by filling fishing nets, and sometimes clog the cooling systems of power and
desalination plants Desalination is a process that takes away mineral components from saline water. More generally, desalination refers to the removal of salts and minerals from a target substance, as in soil desalination, which is an issue for agriculture. Salt ...
which draw their water from the sea.


Names

The name jellyfish, in use since 1796, has traditionally been applied to medusae and all similar animals including the comb jellies ( ctenophores, another phylum). The term ''jellies'' or ''sea jellies'' is more recent, having been introduced by public aquaria in an effort to avoid use of the word "fish" with its modern connotation of an animal with a backbone, though
shellfish Shellfish is a colloquial and fisheries term for exoskeleton-bearing aquatic invertebrates used as food, including various species of molluscs, crustaceans, and echinoderms. Although most kinds of shellfish are harvested from saltwater env ...
,
cuttlefish Cuttlefish or cuttles are marine molluscs of the order Sepiida. They belong to the class Cephalopoda which also includes squid, octopuses, and nautiluses. Cuttlefish have a unique internal shell, the cuttlebone, which is used for control ...
and starfish are not vertebrates either. In scientific literature, "jelly" and "jellyfish" have been used interchangeably. Many sources refer to only scyphozoans as "true jellyfish". A group of jellyfish is called a "smack".


Mapping to taxonomic groups


Phylogeny


Definition

The term jellyfish broadly corresponds to medusae, that is, a life-cycle stage in the Medusozoa. The American evolutionary biologist Paulyn Cartwright gives the following general definition: The Merriam-Webster dictionary defines jellyfish as follows: Given that jellyfish is a common name, its mapping to biological groups is inexact. Some authorities have called the
comb jellies Ctenophora (; ctenophore ; ) comprise a phylum of marine invertebrates, commonly known as comb jellies, that inhabit sea waters worldwide. They are notable for the groups of cilia they use for swimming (commonly referred to as "combs"), and ...
and certain
salp A salp (plural salps, also known colloquially as “sea grape”) or salpa (plural salpae or salpas) is a barrel-shaped, planktic tunicate. It moves by contracting, thereby pumping water through its gelatinous body, one of the most efficient ...
s jellyfish, though other authorities state that neither of these are jellyfish, which they consider should be limited to certain groups within the medusozoa. The non-medusozoan clades called jellyfish by some but not all authorities (both agreeing and disagreeing citations are given in each case) are indicated with "???" on the following cladogram of the animal kingdom:


Medusozoan jellyfish

Jellyfish are not a clade, as they include most of the Medusozoa, barring some of the Hydrozoa. The medusozoan groups included by authorities are indicated on the following phylogenetic tree by the presence of citations. Names of included jellyfish, in English where possible, are shown in boldface; the presence of a named and cited example indicates that at least that species within its group has been called a jellyfish.


Taxonomy

The subphylum Medusozoa includes all cnidarians with a medusa stage in their life cycle. The basic cycle is egg,
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, whi ...
larva, polyp, medusa, with the medusa being the sexual stage. The polyp stage is sometimes secondarily lost. The subphylum include the major taxa, Scyphozoa (large jellyfish), Cubozoa (box jellyfish) and
Hydrozoa Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specializ ...
(small jellyfish), and excludes
Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of t ...
(corals and sea anemones). This suggests that the medusa form evolved after the polyps.Cnidaria
, Tree of Life.
Medusozoans have tetramerous symmetry, with parts in fours or multiples of four. The four major classes of medusozoan Cnidaria are: * Scyphozoa are sometimes called true jellyfish, though they are no more truly jellyfish than the others listed here. They have tetra-radial symmetry. Most have tentacles around the outer margin of the bowl-shaped bell, and long, oral arms around the mouth in the center of the subumbrella. * Cubozoa (box jellyfish) have a (rounded) box-shaped bell, and their velarium assists them to swim more quickly. Box jellyfish may be related more closely to scyphozoan jellyfish than either are to the Hydrozoa. *
Hydrozoa Hydrozoa (hydrozoans; ) are a taxonomic class of individually very small, predatory animals, some solitary and some colonial, most of which inhabit saline water. The colonies of the colonial species can be large, and in some cases the specializ ...
medusae also have tetra-radial symmetry, nearly always have a velum (diaphragm used in swimming) attached just inside the bell margin, do not have oral arms, but a much smaller central stalk-like structure, the manubrium, with terminal mouth opening, and are distinguished by the absence of cells in the mesoglea. Hydrozoa show great diversity of lifestyle; some species maintain the polyp form for their entire life and do not form medusae at all (such as
Hydra Hydra generally refers to: * Lernaean Hydra, a many-headed serpent in Greek mythology * ''Hydra'' (genus), a genus of simple freshwater animals belonging to the phylum Cnidaria Hydra or The Hydra may also refer to: Astronomy * Hydra (constel ...
, which is hence not considered a jellyfish), and a few are entirely medusal and have no polyp form. *
Staurozoa Staurozoa is a class of Medusozoa, jellyfishes and hydrozoans. It has one extant order: Stauromedusae (stalked jellyfishes) with a total of 50 known species. A fossil group called Conulariida has been proposed as a second order, although this ...
(stalked jellyfish) are characterized by a medusa form that is generally sessile, oriented upside down and with a stalk emerging from the apex of the "calyx" (bell), which attaches to the substrate. At least some Staurozoa also have a polyp form that alternates with the medusoid portion of the life cycle. Until recently, Staurozoa were classified within the Scyphozoa. There are over 200 species of Scyphozoa, about 50 species of Staurozoa, about 20 species of Cubozoa, and the Hydrozoa includes about 1000–1500 species that produce medusae, but many more species that do not.


Fossil history

Since jellyfish have no hard parts, fossils are rare. The oldest conulariid scyphozoans appeared between 635 and 577 mya in the
Neoproterozoic The Neoproterozoic Era is the unit of geologic time from 1 billion to 538.8 million years ago. It is the last era of the Precambrian Supereon and the Proterozoic Eon; it is subdivided into the Tonian, Cryogenian, and Ediacaran periods. It is ...
of the
Lantian Formation The Lantian Formation is a 150-meter-thick sequence of rocks deposited in southern China during a 90-million-year epoch in the Ediacaran period. Its algal macrofossils (which have alternatively been interpreted as putative metazoans) are the old ...
in China; others are found in the youngest
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and t ...
rocks of the Tamengo Formation of Brazil, c. 505 mya, through to the
Triassic The Triassic ( ) is a geologic period and system which spans 50.6 million years from the end of the Permian Period 251.902 million years ago (Mya), to the beginning of the Jurassic Period 201.36 Mya. The Triassic is the first and shortest period ...
. Cubozoans and hydrozoans appeared in the
Cambrian The Cambrian Period ( ; sometimes symbolized Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the preceding Ediacaran Period 538.8 million years ag ...
of the
Marjum Formation The Marjum Formation is a Cambrian The Cambrian Period ( ; sometimes symbolized C with bar, Ꞓ) was the first geological period of the Paleozoic Era, and of the Phanerozoic Eon. The Cambrian lasted 53.4 million years from the end of the prece ...
in Utah, USA, c. 540 mya.


Anatomy

The main feature of a true jellyfish is the umbrella-shaped bell. This is a hollow structure consisting of a mass of transparent jelly-like matter known as
mesoglea Mesoglea refers to the extracellular matrix found in cnidarians like coral or jellyfish that functions as a hydrostatic skeleton. It is related to but distinct from mesohyl, which generally refers to extracellular material found in sponges. Desc ...
, which forms the
hydrostatic skeleton A hydrostatic skeleton, or hydroskeleton, is a flexible skeleton supported by fluid pressure. Hydrostatic skeletons are common among simple invertebrate organisms. While more advanced organisms can be considered hydrostatic, they are sometimes refe ...
of the animal. 95% or more of the mesogloea consists of water, but it also contains collagen and other fibrous proteins, as well as wandering amoebocytes which can engulf debris and bacteria. The mesogloea is bordered by the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the ...
on the outside and the gastrodermis on the inside. The edge of the bell is often divided into rounded lobes known as lappets, which allow the bell to flex. In the gaps or niches between the lappets are dangling rudimentary sense organs known as rhopalia, and the margin of the bell often bears tentacles. On the underside of the bell is the manubrium, a stalk-like structure hanging down from the centre, with the mouth, which also functions as the anus, at its tip. There are often four oral arms connected to the manubrium, streaming away into the water below. The mouth opens into the
gastrovascular cavity The gastrovascular cavity is the primary organ of digestion and circulation in two major animal phyla: the Coelenterates or cnidarians (including jellyfish and corals) and Platyhelminthes (flatworms). The cavity may be extensively branched into a ...
, where digestion takes place and nutrients are absorbed. This is subdivided by four thick
septa The Southeastern Pennsylvania Transportation Authority (SEPTA) is a regional public transportation authority that operates transit bus, bus, rapid transit, commuter rail, light rail, and electric trolleybus services for nearly 4 million people ...
into a central stomach and four gastric pockets. The four pairs of gonads are attached to the septa, and close to them four septal funnels open to the exterior, perhaps supplying good oxygenation to the gonads. Near the free edges of the septa, gastric filaments extend into the gastric cavity; these are armed with nematocysts and enzyme-producing cells and play a role in subduing and digesting the prey. In some scyphozoans, the gastric cavity is joined to radial canals which branch extensively and may join a marginal ring canal. Cilia in these canals circulate the fluid in a regular direction. The box jellyfish is largely similar in structure. It has a squarish, box-like bell. A short pedalium or stalk hangs from each of the four lower corners. One or more long, slender tentacles are attached to each pedalium. The rim of the bell is folded inwards to form a shelf known as a velarium which restricts the bell's aperture and creates a powerful jet when the bell pulsates, allowing box jellyfish to swim faster than true jellyfish. Hydrozoans are also similar, usually with just four tentacles at the edge of the bell, although many hydrozoans are colonial and may not have a free-living medusal stage. In some species, a non-detachable bud known as a gonophore is formed that contains a gonad but is missing many other medusal features such as tentacles and rhopalia. Stalked jellyfish are attached to a solid surface by a basal disk, and resemble a polyp, the oral end of which has partially developed into a medusa with tentacle-bearing lobes and a central manubrium with four-sided mouth. Most jellyfish do not have specialized systems for
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 ...
, respiration and
circulation Circulation may refer to: Science and technology * Atmospheric circulation, the large-scale movement of air * Circulation (physics), the path integral of the fluid velocity around a closed curve in a fluid flow field * Circulatory system, a bio ...
, and do not have a
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
. Nematocysts, which deliver the sting, are located mostly on the tentacles; true jellyfish also have them around the mouth and stomach. Jellyfish do not need a respiratory system because sufficient oxygen diffuses through the epidermis. They have limited control over their movement, but can navigate with the pulsations of the bell-like body; some species are active swimmers most of the time, while others largely drift. The rhopalia contain rudimentary sense organs which are able to detect light, water-borne vibrations, odour and orientation. A loose network of nerves called a "
nerve net A nerve net consists of interconnected neurons lacking a brain or any form of cephalization. While organisms with bilateral body symmetry are normally associated with a condensation of neurons or, in more advanced forms, a central nervous syst ...
" is located in the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and Subcutaneous tissue, hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the ...
. Although traditionally thought not to have a
central nervous system The central nervous system (CNS) is the part of the nervous system consisting primarily of the brain and spinal cord. The CNS is so named because the brain integrates the received information and coordinates and influences the activity of all p ...
, nerve net concentration and ganglion-like structures could be considered to constitute one in most species. A jellyfish detects stimuli, and transmits impulses both throughout the nerve net and around a circular nerve ring, to other nerve cells. The rhopalial ganglia contain pacemaker neurones which control swimming rate and direction. In many species of jellyfish, the rhopalia include
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-le ...
, light-sensitive organs able to tell light from dark. These are generally pigment spot ocelli, which have some of their cells pigmented. The rhopalia are suspended on stalks with heavy
crystal A crystal or crystalline solid is a solid material whose constituents (such as atoms, molecules, or ions) are arranged in a highly ordered microscopic structure, forming a crystal lattice that extends in all directions. In addition, macr ...
s at one end, acting like gyroscopes to orient the eyes skyward. Certain jellyfish look upward at the mangrove canopy while making a daily migration from
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in several ...
swamps into the open lagoon, where they feed, and back again.
Box jellyfish Box jellyfish (class Cubozoa) are cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latt ...
have more advanced vision than the other groups. Each individual has 24
eye Eyes are organs of the visual system. They provide living organisms with vision, the ability to receive and process visual detail, as well as enabling several photo response functions that are independent of vision. Eyes detect light and conv ...
s, two of which are capable of seeing colour, and four parallel information processing areas that act in competition, supposedly making them one of the few kinds of animal to have a 360-degree view of its environment.


Box jellyfish eye

The study of jellyfish eye evolution is an intermediary to a better understanding of how visual systems evolved on Earth. Jellyfish exhibit immense variation in visual systems ranging from photoreceptive cell patches seen in simple photoreceptive systems to more derived complex eyes seen in box jellyfish. Major topics of jellyfish visual system research (with an emphasis on box jellyfish) include: the evolution of jellyfish vision from simple to complex visual systems), the eye morphology and molecular structures of box jellyfish (including comparisons to vertebrate eyes), and various uses of vision including task guided behaviors and niche specialization.


Evolution

Experimental evidence for photosensitivity and photoreception in cnidarians antecedes the mid 1900s, and a rich body of research has since covered evolution of visual systems in jellyfish. Jellyfish visual systems range from simple photoreceptive cells to complex image-forming eyes. More ancestral visual systems incorporate extraocular vision (vision without eyes) that encompass numerous receptors dedicated to single function behaviors. More derived visual systems comprise perception that is capable of multiple task guided behaviors. Although they lack a true brain, cnidarian jellyfish have a "ring" nervous system that plays a significant role in motor and sensory activity. This net of nerves is responsible for muscle contraction and movement, and culminates the emergence of photosensitive structures. Across
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latter. Their distinguishing feature is cnidocyt ...
, there is large variation in the systems that underlie photosensitivity. Photosensitive structures range from non-specialized groups of cells, to more "conventional" eyes similar to those of vertebrates. The general evolutionary steps to develop complex vision include (from more ancestral to more derived states): non-directional photoreception, directional photoreception, low-resolution vision, and high-resolution vision. Increased habitat and task complexity has favored the high-resolution visual systems common in derived cnidarians such as box jellyfish. Basal visual systems observed in various cnidarians exhibit photosensitivity representative of a single task or behavior. Extraocular photoreception (a form of non-directional photoreception), is the most basic form of light sensitivity and guides a variety of behaviors among cnidarians. It can function to regulate
circadian rhythm A circadian rhythm (), or circadian cycle, is a natural, internal process that regulates the sleep–wake cycle and repeats roughly every 24 hours. It can refer to any process that originates within an organism (i.e., endogenous) and responds to ...
(as seen in eyeless hydrozoans) and other light-guided behaviors responsive to the intensity and spectrum of light. Extraocular photoreception can function additionally in positive phototaxis (in planula larvae of hydrozoans), as well as in avoiding harmful amounts of UV radiation via negative phototaxis. Directional photoreception (the ability to perceive direction of incoming light) allows for more complex phototactic responses to light, and likely evolved by means of membrane stacking. The resulting behavioral responses can range from guided spawning events timed by moonlight to shadow responses for potential predator avoidance. Light-guided behaviors are observed in numerous scyphozoans including the common moon jelly, '' Aurelia aurita'', which migrates in response to changes in ambient light and solar position even though they lack proper eyes. The low-resolution visual system of box jellyfish is more derived than directional photoreception, and thus box jellyfish vision represents the most basic form of true vision in which multiple directional photoreceptors combine to create the first imaging and spatial resolution. This is different from the high-resolution vision that is observed in camera or compound eyes of vertebrates and cephalopods that rely on focusing optics. Critically, the visual systems of box jellyfish are responsible for guiding multiple tasks or behaviors in contrast to less derived visual systems in other jellyfish that guide single behavioral functions. These behaviors include phototaxis based on sunlight (positive) or shadows (negative), obstacle avoidance, and control of swim-pulse rate. Box jellyfish possess "proper eyes" (similar to vertebrates) that allow them to inhabit environments that lesser derived medusae cannot. In fact, they are considered the only class in the clade '' Medusozoa'' that have behaviors necessitating spatial resolution and genuine vision. However, the
lens A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements'' ...
in their eyes are more functionally similar to cup-eyes exhibited in low-resolution organisms, and have very little to no focusing capability. The lack of the ability to focus is due to the focal length exceeding the distance to the retina, thus generating unfocused images and limiting spatial resolution. The visual system is still sufficient for box jellyfish to produce an image to help with tasks such as object avoidance.


Utility as a model organism

Box jellyfish eyes are a visual system that is sophisticated in numerous ways. These intricacies include the considerable variation within the morphology of box jellyfishes' eyes (including their task/behavior specification), and the molecular makeup of their eyes including: photoreceptors,
opsin Animal opsins are G-protein-coupled receptors and a group of proteins made light-sensitive via a chromophore, typically retinal. When bound to retinal, opsins become Retinylidene proteins, but are usually still called opsins regardless. Most pro ...
s, lenses, and
synapse In the nervous system, a synapse is a structure that permits a neuron (or nerve cell) to pass an electrical or chemical signal to another neuron or to the target effector cell. Synapses are essential to the transmission of nervous impulses fr ...
s. The comparison of these attributes to more derived visual systems can allow for a further understanding of how the evolution of more derived visual systems may have occurred, and puts into perspective how box jellyfish can play the role as an evolutionary/developmental model for all visual systems.


= Characteristics

= Box jellyfish visual systems are both diverse and complex, comprising multiple photosystems. There is likely considerable variation in visual properties between species of box jellyfish given the significant inter-species morphological and physiological variation. Eyes tend to differ in size and shape, along with number of receptors (including opsins), and physiology across species of box jellyfish. Box jellyfish have a series of intricate lensed eyes that are similar to those of more derived multicellular organisms such as vertebrates. Their 24 eyes fit into four different morphological categories. These categories consist of two large, morphologically different medial eyes (a lower and upper lensed eye) containing spherical lenses, a lateral pair of pigment slit eyes, and a lateral pair of pigment pit eyes. The eyes are situated on rhopalia (small sensory structures) which serve sensory functions of the box jellyfish and arise from the cavities of the exumbrella (the surface of the body) on the side of the bells of the jellyfish. The two large eyes are located on the mid-line of the club and are considered complex because they contain lenses. The four remaining eyes lie laterally on either side of each rhopalia and are considered simple. The simple eyes are observed as small invaginated cups of epithelium that have developed pigmentation. The larger of the complex eyes contains a cellular
cornea The cornea is the transparent front part of the eye that covers the iris, pupil, and anterior chamber. Along with the anterior chamber and lens, the cornea refracts light, accounting for approximately two-thirds of the eye's total optical ...
created by a monociliated epithelium, cellular lens, homogenous capsule to the lens, vitreous body with prismatic elements, and a
retina The retina (from la, rete "net") is the innermost, light-sensitive layer of tissue of the eye of most vertebrates and some molluscs. The optics of the eye create a focused two-dimensional image of the visual world on the retina, which then ...
of pigmented cells. The smaller of the complex eyes is said to be slightly less complex given that it lacks a capsule but otherwise contains the same structure as the larger eye. Box jellyfish have multiple photosystems that comprise different sets of eyes. Evidence includes immunocytochemical and molecular data that show photopigment differences among the different morphological eye types, and physiological experiments done on box jellyfish to suggest behavioral differences among photosystems. Each individual eye type constitutes photosystems that work collectively to control visually guided behaviors. Box jellyfish eyes primarily use c-PRCs (ciliary photoreceptor cells) similar to that of vertebrate eyes. These cells undergo phototransduction cascades (process of light absorption by photoreceptors) that are triggered by c-opsins. Available opsin sequences suggest that there are two types of opsins possessed by all cnidarians including an ancient
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
opsin, and a sister ciliary opsin to the c-opsins group. Box jellyfish could have both ciliary and cnidops (cnidarian opsins), which is something not previously believed to appear in the same retina. Nevertheless, it is not entirely evident whether cnidarians possess multiple opsins that are capable of having distinctive spectral sensitivities.


= Comparison with other organisms

= Comparative research on genetic and molecular makeup of box jellyfishes' eyes versus more derived eyes seen in vertebrates and cephalopods focuses on: lenses and crystallin composition, synapses, and Pax genes and their implied evidence for shared primordial (ancestral) genes in eye evolution. Box jellyfish eyes are said to be an evolutionary/developmental model of all eyes based on their evolutionary recruitment of crystallins and Pax genes. Research done on box jellyfish including ''Tripedalia cystophora'' has suggested that they possess a single Pax gene, PaxB. PaxB functions by binding to crystallin promoters and activating them. PaxB
in situ hybridization ''In situ'' hybridization (ISH) is a type of hybridization that uses a labeled complementary DNA, RNA or modified nucleic acids strand (i.e., probe) to localize a specific DNA or RNA sequence in a portion or section of tissue (''in situ'') or ...
resulted in PaxB expression in the lens, retina, and statocysts. These results and the rejection of the prior hypothesis that Pax6 was an ancestral Pax gene in eyes has led to the conclusion that PaxB was a primordial gene in eye evolution, and that it is highly likely that eyes of all organisms share a common ancestor. The lens structure of box jellyfish appears very similar to those of other organisms, but the crystallins are distinct in both function and appearance. Weak reactions were seen within the sera and there were very weak sequence similarities within the crystallins among vertebrate and invertebrate lenses. This is likely due to differences in lower molecular weight proteins and the subsequent lack of immunological reactions with antisera that other organisms' lenses exhibit. All four of the visual systems of box jellyfish species investigated with detail (''Carybdea marsupialis, Chiropsalmus quadrumanus, Tamoya haplonema and Tripedalia cystophora'') have invaginated synapses, but only in the upper and lower lensed eyes. Different densities were found between the upper and lower lenses, and between species. Four types of chemical synapses have been discovered within the rhopalia which could help in understanding neural organization including: clear unidirectional, dense-core unidirectional, clear bidirectional, and clear and dense-core bidirectional. The synapses of the lensed eyes could be useful as markers to learn more about the neural circuit in box jellyfish retinal areas.


Evolution as a response to natural stimuli

The primary adaptive responses to environmental variation observed in box jellyfish eyes include pupillary constriction speeds in response to light environments, as well as photoreceptor tuning and lens adaptations to better respond to shifts between light environments and darkness. Interestingly, some box jellyfish species' eyes appear to have evolved more focused vision in response to their habitat. Pupillary contraction appears to have evolved in response to variation in the light environment across
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (fo ...
s across three species of box jellyfish ('' Chironex fleckeri, Chiropsella bronzie, and Carukia barnesi''). Behavioral studies suggest that faster pupil contraction rates allow for greater object avoidance, and in fact, species with more complex habitats exhibit faster rates. ''Ch. bronzie'' inhabit shallow beach fronts that have low visibility and very few obstacles, thus, faster pupil contraction in response to objects in their environment is not important. ''Ca. barnesi'' and ''Ch. fleckeri'' are found in more three dimensionally complex environments like
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in several ...
s with an abundance of natural obstacles, where faster pupil contraction is more adaptive. Behavioral studies support the idea that faster pupillary contraction rates assist with obstacle avoidance as well as depth adjustments in response to differing light intensities. Light/dark adaptation via pupillary light reflexes is an additional form of an evolutionary response to the light environment. This relates to the pupil's response to shifts between light intensity (generally from sunlight to darkness). In the process of light/dark adaptation, the upper and lower lens eyes of different box jellyfish species vary in specific function. The lower lens-eyes contain pigmented photoreceptors and long pigment-cells with dark pigments that migrate on light/dark adaptation, while the upper-lens eyes play a concentrated role in light direction and phototaxis given that they face upward towards the water surface (towards the sun or moon). The upper lens of ''Ch. bronzie'' does not exhibit any considerable optical power while ''Tr. cystophora'' (a box jellyfish species that tends to live in mangroves) does. The ability to use light to visually guide behavior is not of as much importance to ''Ch. bronzie'' as it is to species in more obstacle filled environments. Differences in visually guided behavior serve as evidence that species that share the same number and structure of eyes can exhibit differences in how they control behavior.


Largest and smallest

Jellyfish range from about one millimeter in bell height and diameter, to nearly in bell height and diameter; the tentacles and mouth parts usually extend beyond this bell dimension. The smallest jellyfish are the peculiar creeping jellyfish in the genera ''
Staurocladia ''Staurocladia'' is a genus of anthomedusan hydrozoans. Its former family Eleutheriidae is now included in the Cladonematidae. Species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as we ...
'' and '' Eleutheria'', which have bell disks from to a few millimeters in diameter, with short tentacles that extend out beyond this, which these jellyfish use to move across the surface of seaweed or the bottoms of rocky pools; many of these tiny creeping jellyfish cannot be seen in the field without a hand lens or microscope. They can reproduce asexually by
fission Fission, a splitting of something into two or more parts, may refer to: * Fission (biology), the division of a single entity into two or more parts and the regeneration of those parts into separate entities resembling the original * Nuclear fissio ...
(splitting in half). Other very small jellyfish, which have bells about one millimeter, are the hydromedusae of many species that have just been released from their parent polyps; some of these live only a few minutes before shedding their gametes in the plankton and then dying, while others will grow in the plankton for weeks or months. The hydromedusae ''Cladonema radiatum'' and ''Cladonema californicum'' are also very small, living for months, yet never growing beyond a few mm in bell height and diameter. The lion's mane jellyfish, '' Cyanea capillata'', was long-cited as the largest jellyfish, and arguably the longest animal in the world, with fine, thread-like tentacles that may extend up to long (though most are nowhere near that large). They have a moderately painful, but rarely fatal, sting. The increasingly common giant Nomura's jellyfish, '' Nemopilema nomurai'', found in some, but not all years in the waters of Japan, Korea and China in summer and autumn is another candidate for "largest jellyfish", in terms of diameter and weight, since the largest Nomura's jellyfish in late autumn can reach in bell (body) diameter and about in weight, with average specimens frequently reaching in bell diameter and about in weight. The large bell mass of the giant Nomura's jellyfish can dwarf a diver and is nearly always much greater than the Lion's Mane, whose bell diameter can reach . The rarely encountered deep-sea jellyfish ''
Stygiomedusa gigantea ''Stygiomedusa gigantea'', commonly known as the giant phantom jelly is a part of the monotypic genus of deep sea jellyfish, ''Stygiomedusa''. This is in the Ulmaridae family. With only around 110 sightings in 110 years, it is a jellyfish that is ...
'' is another candidate for "largest jellyfish", with its thick, massive bell up to wide, and four thick, "strap-like" oral arms extending up to in length, very different from the typical fine, threadlike tentacles that rim the umbrella of more-typical-looking jellyfish, including the Lion's Mane.


Life history and behavior


Life cycle

Jellyfish have a complex life cycle which includes both sexual and asexual phases, with the medusa being the sexual stage in most instances. Sperm fertilize eggs, which develop into larval planulae, become polyps, bud into ephyrae and then transform into adult medusae. In some species certain stages may be skipped. Upon reaching adult size, jellyfish spawn regularly if there is a sufficient supply of food. In most species, spawning is controlled by light, with all individuals spawning at about the same time of day; in many instances this is at dawn or dusk. Jellyfish are usually either male or female (with occasional
hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have ...
s). In most cases, adults release sperm and eggs into the surrounding water, where the unprotected eggs are fertilized and develop into larvae. In a few species, the sperm swim into the female's mouth, fertilizing the eggs within her body, where they remain during early development stages. In moon jellies, the eggs lodge in pits on the oral arms, which form a temporary brood chamber for the developing
planula A planula is the free-swimming, flattened, ciliated, bilaterally symmetric larval form of various cnidarian species and also in some species of Ctenophores. Some groups of Nemerteans also produce larvae that are very similar to the planula, whi ...
larvae. The planula is a small
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. Th ...
covered with
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
. When sufficiently developed, it settles onto a firm surface and develops into a polyp. The polyp generally consists of a small stalk topped by a mouth that is ringed by upward-facing tentacles. The polyps resemble those of closely related
anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of t ...
ns, such as sea anemones and
coral Corals are marine invertebrates within the class Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important reef builders that inhabit tropical oceans and secre ...
s. The jellyfish polyp may be sessile, living on the bottom, boat hulls or other substrates, or it may be free-floating or attached to tiny bits of free-living plankton or rarely, fish or other invertebrates. Polyps may be solitary or colonial. Most polyps are only millimetres in diameter and feed continuously. The polyp stage may last for years. After an interval and stimulated by seasonal or hormonal changes, the polyp may begin reproducing asexually by
budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...
and, in the Scyphozoa, is called a segmenting polyp, or a scyphistoma. Budding produces more scyphistomae and also ephyrae. Budding sites vary by species; from the
tentacle In zoology, a tentacle is a flexible, mobile, and elongated organ present in some species of animals, most of them invertebrates. In animal anatomy, tentacles usually occur in one or more pairs. Anatomically, the tentacles of animals work main ...
bulbs, the manubrium (above the mouth), or the
gonad A gonad, sex gland, or reproductive gland is a mixed gland 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, produces ...
s of hydromedusae. In a process known as strobilation, the polyp's tentacles are reabsorbed and the body starts to narrow, forming transverse constrictions, in several places near the upper extremity of the polyp. These deepen as the constriction sites migrate down the body, and separate segments known as ephyra detach. These are free-swimming precursors of the adult medusa stage, which is the life stage that is typically identified as a jellyfish. The ephyrae, usually only a millimeter or two across initially, swim away from the polyp and grow. Limnomedusae polyps can asexually produce a creeping ''frustule'' larval form, which crawls away before developing into another polyp. A few species can produce new medusae by budding directly from the medusan stage. Some hydromedusae reproduce by fission.


Lifespan

Little is known of the life histories of many jellyfish as the places on the seabed where the benthic forms of those species live have not been found. However, an asexually reproducing strobila form can sometimes live for several years, producing new medusae (ephyra larvae) each year. An unusual species, '' Turritopsis dohrnii'', formerly classified as ''
Turritopsis nutricula ''Turritopsis nutricula'' is a small hydrozoan that once reaching adulthood, can transfer its cells back to childhood. This adaptive trait likely evolved in order to extend the life of the individual. Several different species of the genus ''Tur ...
'', might be effectively immortal because of its ability under certain circumstances to transform from medusa back to the polyp stage, thereby escaping the death that typically awaits medusae post-reproduction if they have not otherwise been eaten by some other organism. So far this reversal has been observed only in the laboratory.


Locomotion

Using the moon jelly '' Aurelia aurita'' as an example, jellyfish have been shown to be the most energy efficient swimmers of all animals. They move through the water by radially expanding and contracting their bell-shaped bodies to push water behind them. They pause between the contraction and expansion phases to create two
vortex In fluid dynamics, a vortex ( : vortices or vortexes) is a region in a fluid in which the flow revolves around an axis line, which may be straight or curved. Vortices form in stirred fluids, and may be observed in smoke rings, whirlpools in t ...
rings. Muscles are used for the contraction of the body, which creates the first vortex and pushes the animal forward, but the mesoglea is so elastic that the expansion is powered exclusively by relaxing the bell, which releases the energy stored from the contraction. Meanwhile, the second vortex ring starts to spin faster, sucking water into the bell and pushing against the centre of the body, giving a secondary and "free" boost forward. The mechanism, called passive energy recapture, only works in relatively small jellyfish moving at low speeds, allowing the animal to travel 30 percent farther on each swimming cycle. Jellyfish achieved a 48 percent lower cost of transport (food and oxygen intake versus energy spent in movement) than other animals in similar studies. One reason for this is that most of the gelatinous tissue of the bell is inactive, using no energy during swimming.


Ecology


Diet

Jellyfish are like other cnidarians generally carnivorous (or parasitic), feeding on planktonic organisms, crustaceans, small fish, fish eggs and larvae, and other jellyfish, ingesting food and voiding undigested waste through the mouth. They hunt passively using their tentacles as drift lines, or sink through the water with their tentacles spread widely; the tentacles, which contain nematocysts to stun or kill the
prey Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
, may then flex to help bring it to the mouth. Their swimming technique also helps them to capture prey; when their bell expands it sucks in water which brings more potential prey within reach of the tentacles. A few species such as ''
Aglaura hemistoma ''Aglaura'' is a monotypic genus of deep-sea hydrozoan in the family Rhopalonematidae. It is represented by the species ''Aglaura hemistoma''. It has a cosmopolitan distribution In biogeography, cosmopolitan distribution is the term for th ...
'' are omnivorous, feeding on microplankton which is a mixture of
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
and
phytoplankton Phytoplankton () are the autotrophic (self-feeding) components of the plankton community and a key part of ocean and freshwater ecosystems. The name comes from the Greek words (), meaning 'plant', and (), meaning 'wanderer' or 'drifter'. P ...
(microscopic plants) such as
dinoflagellate The dinoflagellates (Greek δῖνος ''dinos'' "whirling" and Latin ''flagellum'' "whip, scourge") are a monophyletic group of single-celled eukaryotes constituting the phylum Dinoflagellata and are usually considered algae. Dinoflagellates are ...
s. Others harbour mutualistic algae (
Zooxanthellae Zooxanthellae is a colloquial term for single-celled dinoflagellates that are able to live in symbiosis with diverse marine invertebrates including demosponges, corals, jellyfish, and nudibranchs. Most known zooxanthellae are in the genus '' ...
) in their tissues; the spotted jellyfish (''
Mastigias papua The spotted jelly (''Mastigias papua''), lagoon jelly, golden medusa, or Papuan jellyfish, is a species of jellyfish from the Indo-Pacific oceans. Like corals, sea anemones, and other sea jellies, it belongs to the phylum Cnidaria. ''Mastigias p ...
'') is typical of these, deriving part of its nutrition from the products of
photosynthesis Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
, and part from captured zooplankton. The upside-down jellyfish (''Cassiopea andromeda'') also has a symbiotic relationship with
microalgae Microalgae or microphytes are microscopic algae invisible to the naked eye. They are phytoplankton typically found in freshwater and marine systems, living in both the water column and sediment. They are unicellular species which exist indi ...
, but captures tiny animals to supplement their diet. This is done by releasing tiny balls of living cells composed of
mesoglea Mesoglea refers to the extracellular matrix found in cnidarians like coral or jellyfish that functions as a hydrostatic skeleton. It is related to but distinct from mesohyl, which generally refers to extracellular material found in sponges. Desc ...
. These use cilia to drive them through water and stinging cells which stun the prey. The blobs also seems to have digestive capabilities.


Predation

Other species of jellyfish are among the most common and important jellyfish predators. Sea anemones may eat jellyfish that drift into their range. Other predators include
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae ( mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max le ...
s, sharks,
swordfish Swordfish (''Xiphias gladius''), also known as broadbills in some countries, are large, highly migratory predatory fish characterized by a long, flat, pointed bill. They are a popular sport fish of the billfish category, though elusive. Swordfi ...
, sea turtles and penguins. Jellyfish washed up on the beach are consumed by foxes, other terrestrial mammals and birds. In general however, few animals prey on jellyfish; they can broadly be considered to be top predators in the food chain. Once jellyfish have become dominant in an ecosystem, for example through overfishing which removes predators of jellyfish larvae, there may be no obvious way for the previous balance to be restored: they eat fish eggs and juvenile fish, and compete with fish for food, preventing fish stocks from recovering.


Symbiosis

Some small fish are immune to the stings of the jellyfish and live among the tentacles, serving as bait in a fish trap; they are safe from potential predators and are able to share the fish caught by the jellyfish. The
cannonball jellyfish The cannonball jellyfish (''Stomolophus meleagris''), also known as the cabbagehead jellyfish, is a species of jellyfish in the family Stomolophidae. Its common name derives from its similarity to a cannonball in shape and size. Its dome-shaped b ...
has a symbiotic relationship with ten different species of fish, and with the longnose spider crab, which lives inside the bell, sharing the jellyfish's food and nibbling its tissues.


Blooms

Jellyfish form large masses or blooms in certain environmental conditions of
ocean current An ocean current is a continuous, directed movement of sea water generated by a number of forces acting upon the water, including wind, the Coriolis effect, breaking waves, cabbeling, and temperature and salinity differences. Depth contou ...
s,
nutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excr ...
s, sunshine, temperature, season, prey availability, reduced predation and
oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as we ...
concentration. Currents collect jellyfish together, especially in years with unusually high populations. Jellyfish can detect
marine current An ocean current is a continuous, directed movement of sea water generated by a number of forces acting upon the water, including wind, the Coriolis effect, breaking waves, cabbeling, and temperature and salinity differences. Depth contou ...
s and swim against the current to congregate in blooms. Jellyfish are better able to survive in nutrient-rich, oxygen-poor water than competitors, and thus can feast on plankton without competition. Jellyfish may also benefit from saltier waters, as saltier waters contain more
iodine Iodine is a chemical element with the Symbol (chemistry), symbol I and atomic number 53. The heaviest of the stable halogens, it exists as a semi-lustrous, non-metallic solid at standard conditions that melts to form a deep violet liquid at , ...
, which is necessary for polyps to turn into jellyfish. Rising sea temperatures caused by
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...
may also contribute to jellyfish blooms, because many species of jellyfish are able to survive in warmer waters. Increased nutrients from agricultural or urban runoff with nutrients including nitrogen and phosphorus compounds increase the growth of phytoplankton, causing
eutrophication Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytopla ...
and
algal bloom An algal bloom or algae bloom is a rapid increase or accumulation in the population of algae in freshwater or marine water systems. It is often recognized by the discoloration in the water from the algae's pigments. The term ''algae'' encompas ...
s. When the phytoplankton die, they may create dead zones, so called because they are hypoxic (low in oxygen). This in turn kills fish and other animals, but not jellyfish, allowing them to bloom.''
The Washington Post ''The Washington Post'' (also known as the ''Post'' and, informally, ''WaPo'') is an American daily newspaper published in Washington, D.C. It is the most widely circulated newspaper within the Washington metropolitan area and has a large n ...
'', republished in the European Cetacean Bycatch Campaign
''Jellyfish "blooms" could be sign of ailing seas''
, 6 May 2002. Retrieved 25 November 2007.
Jellyfish populations may be expanding globally as a result of land runoff and
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in the ...
of their natural predators. Jellyfish are well placed to benefit from disturbance of marine ecosystems. They reproduce rapidly; they prey upon many species, while few species prey on them; and they feed via touch rather than visually, so they can feed effectively at night and in turbid waters. It may be difficult for
fish stocks Fish stocks are subpopulations of a particular species of fish, for which intrinsic parameters (growth, recruitment, mortality and fishing mortality) are traditionally regarded as the significant factors determining the stock's population dyn ...
to re-establish themselves in marine ecosystems once they have become dominated by jellyfish, because jellyfish feed on plankton, which includes fish eggs and
larvae A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. T ...
. As suspected at the turn of this century, jellyfish blooms are increasing in frequency. Between 2013 and 2020 the Mediterranean Science Commission monitored on a weekly basis the frequency of such outbreaks in coastal waters from Morocco to the Black Sea, revealing a relatively high frequency of these blooms nearly all year round, with peaks observed from March to July and often again in the autumn. The blooms are caused by different jellyfish species, depending on their localisation within the Basin: one observes a clear dominance of ''
Pelagia noctiluca ''Pelagia noctiluca'' is a jellyfish in the family Pelagiidae and the only currently recognized species in the genus ''Pelagia''. It is typically known in English as the mauve stinger, but other common names are purple-striped jelly (causing pot ...
'' and '' Velella velella'' outbreaks in the western Mediterranean, of '' Rhizostoma pulmo'' and '' Rhopilema nomadica'' outbreaks in the eastern Mediterranean, and of '' Aurelia aurita'' and '' Mnemiopsis leidyi'' outbreaks in the Black Sea. Some jellyfish populations that have shown clear increases in the past few decades are
invasive species An invasive species otherwise known as an alien is an introduced organism that becomes overpopulated and harms its new environment. Although most introduced species are neutral or beneficial with respect to other species, invasive species adv ...
, newly arrived from other habitats: examples include the
Black Sea The Black Sea is a marginal mediterranean sea of the Atlantic Ocean lying between Europe and Asia, east of the Balkans, south of the East European Plain, west of the Caucasus, and north of Anatolia. It is bounded by Bulgaria, Georgia, ...
,
Caspian Sea The Caspian Sea is the world's largest inland body of water, often described as the List of lakes by area, world's largest lake or a full-fledged sea. An endorheic basin, it lies between Europe and Asia; east of the Caucasus, west of the broad s ...
,
Baltic Sea The Baltic Sea is an arm of the Atlantic Ocean that is enclosed by Denmark, Estonia, Finland, Germany, Latvia, Lithuania, Poland, Russia, Sweden and the North and Central European Plain. The sea stretches from 53°N to 66°N latitude and fr ...
, central and eastern
Mediterranean The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on th ...
,
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only ...
, and tropical and subtropical parts of the West Atlantic (including the Caribbean,
Gulf of Mexico The Gulf of Mexico ( es, Golfo de México) is an ocean basin and a marginal sea of the Atlantic Ocean, largely surrounded by the North American continent. It is bounded on the northeast, north and northwest by the Gulf Coast of the United S ...
and Brazil). Jellyfish blooms can have significant impact on community structure. Some carnivorous jellyfish species prey on zooplankton while others graze on primary producers. Reductions in zooplankton and ichthyoplankton due to a jellyfish bloom can ripple through the trophic levels. High density jellyfish populations can outcompete other predators and reduce fish recruitment. Increased grazing on primary producers by jellyfish can also interrupt energy transfer to higher trophic levels. During blooms, jellyfish significantly alter the nutrient availability in their environment. Blooms require large amounts of available organic nutrients in the water column to grow, limiting availability for other organisms. Some jellyfish have a symbiotic relationship with single-celled dinoflagellates, allowing them to assimilate inorganic carbon, phosphorus, and nitrogen creating competition for phytoplankton. Their large biomass makes them an important source of dissolved and particulate organic matter for microbial communities through excretion, mucus production, and decomposition. The microbes break down the organic matter into inorganic ammonium and phosphate. However, the low carbon availability shifts the process from production to respiration creating low oxygen areas making the dissolved inorganic nitrogen and phosphorus largely unavailable for primary production. These blooms have very real impacts on industries. Jellyfish can outcompete fish by utilizing open niches in over-fished fisheries. Catch of jellyfish can strain fishing gear and lead to expenses relating to damaged gear. Power plants have been shut down due to jellyfish blocking the flow of cooling water. Blooms have also been harmful for tourism, causing a rise in stings and sometimes the closure of beaches. Jellyfish form a component of jelly-falls, events where gelatinous
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
fall to the seafloor, providing food for 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 ancient Greek, βένθος (bénthos), meaning "t ...
organisms there. In temperate and subpolar regions, jelly-falls usually follow immediately after a bloom.


Habitats

Most jellyfish are marine animals, although a few hydromedusae inhabit
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does in ...
. The best known freshwater example is the cosmopolitan hydrozoan jellyfish, '' Craspedacusta sowerbii''. It is less than an inch (2.5 cm) in diameter, colorless and does not sting. Some jellyfish populations have become restricted to coastal saltwater lakes, such as
Jellyfish Lake Jellyfish Lake ( pau, Ongeim'l Tketau, lit=Fifth Lake) is a marine lake located on Eil Malk island in Palau. Eil Malk is part of the Rock Islands, a group of small, rocky, mostly uninhabited islands in Palau's Southern Lagoon, between Koror an ...
in
Palau Palau,, officially the Republic of Palau and historically ''Belau'', ''Palaos'' or ''Pelew'', is an island country and microstate in the western Pacific. The nation has approximately 340 islands and connects the western chain of the Ca ...
. Jellyfish Lake is a
marine lake A marine lake is a lake situated very close to the sea, which has a connection to this sea, via tunnels or cracks in the soil. Marine lakes are fed with seawater but are usually also fed with rainwater, meaning that the salt content of the water i ...
where millions of golden jellyfish (''Mastigias'' spp.) migrate horizontally across the lake daily. Although most jellyfish live well off the ocean floor and form part of the plankton, a few species are closely associated with the bottom for much of their lives and can be considered
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 ancient Greek, βένθος (bénthos), meaning "t ...
. The upside-down jellyfish in the genus '' Cassiopea'' typically lie on the bottom of shallow lagoons where they sometimes pulsate gently with their umbrella top facing down. Even some deep-sea species of hydromedusae and scyphomedusae are usually collected on or near the bottom. All of the stauromedusae are found attached to either seaweed or rocky or other firm material on the bottom. Some species explicitly adapt to tidal flux. In Roscoe Bay, jellyfish ride the current at ebb tide until they hit a gravel bar, and then descend below the current. They remain in still waters until the tide rises, ascending and allowing it to sweep them back into the bay. They also actively avoid fresh water from mountain snowmelt, diving until they find enough salt.


Parasites

Jellyfish are hosts to a wide variety of parasitic organisms. They act as intermediate hosts of endoparasitic
helminth Parasitic worms, also known as helminths, are large macroparasites Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adap ...
s, with the infection being transferred to the definitive host fish after
predation Predation is a biological interaction In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species (intraspecific interactions), or o ...
. Some digenean
trematode Trematoda is a Class (biology), class of flatworms known as flukes. They are obligate parasite, obligate internal Parasitism, parasites with a complex biological life cycle, life cycle requiring at least two Host_(biology), hosts. The intermedia ...
s, especially species in the family Lepocreadiidae, use jellyfish as their second intermediate hosts. Fish become infected by the trematodes when they feed on infected jellyfish.


Relation to humans


Fisheries

Jellyfish have long been eaten in some parts of the world. Fisheries have begun harvesting the American cannonball jellyfish, ''
Stomolophus meleagris The cannonball jellyfish (''Stomolophus meleagris''), also known as the cabbagehead jellyfish, is a species of jellyfish in the family Stomolophidae. Its common name derives from its similarity to a cannonball in shape and size. Its dome-shaped ...
'', along the southern Atlantic coast of the United States and in the Gulf of Mexico for export to Asia. Jellyfish are also harvested for their collagen, which is being investigated for use in a variety of applications including the treatment of
rheumatoid arthritis Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects synovial joint, joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and ...
.
Aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lotus ...
and fisheries of other species often suffer severe losses – and so losses of productivity – due to jellyfish.


Products

Aristotle Aristotle (; grc-gre, Ἀριστοτέλης ''Aristotélēs'', ; 384–322 BC) was a Greek philosopher and polymath during the Classical Greece, Classical period in Ancient Greece. Taught by Plato, he was the founder of the Peripatet ...
stated in the ''
Parts of Animals ''Parts of Animals'' (or ''On the Parts of Animals''; Greek Περὶ ζῴων μορίων; Latin ''De Partibus Animalium'') is one of Aristotle's major texts on biology. It was written around 350 BC. The whole work is roughly a study in animal ...
'' IV, 6 that jellyfish (sea-nettles) were eaten in winter time in a fish stew. In some countries, including China, Japan, and Korea, jellyfish are a delicacy. The jellyfish is dried to prevent spoiling. Only some 12 species of scyphozoan jellyfish belonging to the order Rhizostomeae are harvested for food, mostly in southeast Asia. Rhizostomes, especially '' Rhopilema esculentum'' in China ( ''hǎizhé'', 'sea stingers') and ''Stomolophus meleagris'' (cannonball jellyfish) in the United States, are favored because of their larger and more rigid bodies and because their toxins are harmless to humans. Traditional processing methods, carried out by a jellyfish master, involve a 20- to 40-day multi-phase procedure in which, after removing the gonads and
mucous membrane A mucous membrane or mucosa is a membrane that lines various cavities in the body of an organism and covers the surface of internal organs. It consists of one or more layers of epithelial cells overlying a layer of loose connective tissue. It i ...
s, the umbrella and oral arms are treated with a mixture of table salt and
alum An alum () is a type of chemical compound, usually a hydrated double sulfate salt of aluminium with the general formula , where is a monovalent cation such as potassium or ammonium. By itself, "alum" often refers to potassium alum, with t ...
, and compressed. Processing makes the jellyfish drier and more acidic, producing a crisp texture. Jellyfish prepared this way retain 7–10% of their original weight, and the processed product consists of approximately 94% water and 6% protein. Freshly processed jellyfish has a white, creamy color and turns yellow or brown during prolonged storage. In China, processed jellyfish are desalted by soaking in water overnight and eaten cooked or raw. The dish is often served shredded with a dressing of oil, soy sauce, vinegar and sugar, or as a salad with vegetables. In Japan, cured jellyfish are rinsed, cut into strips and served with vinegar as an appetizer. Desalted, ready-to-eat products are also available.


Biotechnology

Pliny the Elder Gaius Plinius Secundus (AD 23/2479), called Pliny the Elder (), was a Roman author, naturalist and natural philosopher, and naval and army commander of the early Roman Empire, and a friend of the emperor Vespasian. He wrote the encyclopedic ...
reported in his '' Natural History'' that the slime of the jellyfish "" produced light when rubbed on a walking stick. In 1961,
Osamu Shimomura was a Japanese organic chemist and marine biologist, and Professor Emeritus at Marine Biological Laboratory (MBL) in Woods Hole, Massachusetts and Boston University School of Medicine. He was awarded the Nobel Prize in Chemistry in 2008 for ...
extracted
green fluorescent protein The green fluorescent protein (GFP) is a protein that exhibits bright green fluorescence when exposed to light in the blue to ultraviolet range. The label ''GFP'' traditionally refers to the protein first isolated from the jellyfish '' Aeq ...
(GFP) and another bioluminescent protein, called aequorin, from the large and abundant hydromedusa '' Aequorea victoria'', while studying photoproteins that cause
bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some ...
in this species. Three decades later,
Douglas Prasher Douglas C. Prasher (born August 1951) is an American molecular biologist. He is known for his work to clone and sequence the genes for the photoprotein aequorin and green fluorescent protein (GFP) and for his proposal to use GFP as a tracer mole ...
sequenced and cloned the gene for GFP.
Martin Chalfie Martin Lee Chalfie (born January 15, 1947) is an American scientist. He is List of university professors at Columbia University, University Professor at Columbia University. He shared the 2008 Nobel Prize in Chemistry along with Osamu Shimomura ...
figured out how to use GFP as a fluorescent marker of genes inserted into other cells or organisms. Roger Tsien later chemically manipulated GFP to produce other fluorescent colors to use as markers. In 2008, Shimomura, Chalfie and Tsien won the
Nobel Prize in Chemistry ) , image = Nobel Prize.png , alt = A golden medallion with an embossed image of a bearded man facing left in profile. To the left of the man is the text "ALFR•" then "NOBEL", and on the right, the text (smaller) "NAT•" then "M ...
for their work with GFP. Man-made GFP became widely used as a fluorescent tag to show which cells or tissues express specific genes. The
genetic engineering Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including ...
technique fuses the
gene In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
of interest to the GFP gene. The fused DNA is then put into a cell, to generate either a cell line or (via IVF techniques) an entire animal bearing the gene. In the cell or animal, the
artificial gene Artificial gene synthesis, or simply gene synthesis, refers to a group of methods that are used in synthetic biology to construct and assemble genes from nucleotides ''De novo synthesis, de novo''. Unlike DNA synthesis in living cells, artificial ...
turns on in the same tissues and the same time as the normal gene, making a fusion of the normal protein with GFP attached to the end, illuminating the animal or cell reveals what tissues express that protein—or at what stage of development. The fluorescence shows where the gene is expressed.


Aquarium display

Jellyfish are displayed in many
public aquarium A public aquarium (plural: ''public aquaria'' or ''public Water Zoo'') is the aquatic counterpart of a zoo, which houses living aquatic animal and plant specimens for public viewing. Most public aquariums feature tanks larger than those kept ...
s. Often the tank's background is blue and the animals are illuminated by side light, increasing the contrast between the animal and the background. In natural conditions, many jellies are so transparent that they are nearly invisible. Jellyfish are not adapted to closed spaces. They depend on currents to transport them from place to place. Professional exhibits as in the
Monterey Bay Aquarium Monterey Bay Aquarium is a Nonprofit organization, nonprofit public aquarium in Monterey, California, Monterey, California. Known for its regional focus on the marine habitats of Monterey Bay, it was the first to exhibit a living kelp forest w ...
feature precise water flows, typically in circular tanks to avoid trapping specimens in corners. The outflow is spread out over a large surface area and the inflow enters as a sheet of water in front of the outflow, so the jellyfish do not get sucked into it. As of 2009, jellyfish were becoming popular in home aquariums, where they require similar equipment.


Stings

Jellyfish are armed with nematocysts, a type of specialized stinging cell. Contact with a jellyfish tentacle can trigger millions of nematocysts to pierce the skin and inject
venom Venom or zootoxin is a type of toxin produced by an animal that is actively delivered through a wound by means of a bite, sting, or similar action. The toxin is delivered through a specially evolved ''venom apparatus'', such as fangs or a ...
, but only some species' venom causes an adverse reaction in humans. In a study published in
Communications Biology ''Communications Biology'' is a peer-reviewed open access scientific journal covering research in biology. It was established in 2018 and is published by Nature Portfolio. It is a sister journal to ''Communications Physics'' and '' Communications ...
, researchers found a jellyfish species called ''
Cassiopea xamachana ''Cassiopea xamachana'', commonly known as the upside-down jellyfish, is a species of jellyfish in the family Cassiopeidae. It is found in warm parts of the western Atlantic Ocean, the Caribbean Sea, and the Gulf of Mexico. It was first descr ...
'' which when triggered will release tiny balls of cells that swim around the jellyfish stinging everything in their path. Researchers described these as "self-propelling microscopic grenades" and named them cassiosomes. The effects of stings range from mild discomfort to extreme pain and death. Most jellyfish stings are not deadly, but stings of some box jellyfish ( Irukandji jellyfish), such as the
sea wasp ''Chironex fleckeri'', commonly known as the Australian box jelly, and nicknamed the sea wasp, is a species of extremely venomous box jellyfish found in coastal waters from northern Australia and New Guinea to Malaysia, the Philippines and Vietn ...
, can be deadly. Stings may cause
anaphylaxis Anaphylaxis is a serious, potentially fatal allergic reaction and medical emergency that is rapid in onset and requires immediate medical attention regardless of use of emergency medication on site. It typically causes more than one of the follo ...
(a form of shock), which can be fatal. Jellyfish kill 20 to 40 people a year in the Philippines alone. In 2006 the Spanish Red Cross treated 19,000 stung swimmers along the
Costa Brava The Costa Brava (, ; "Wild Coast" or "Rough Coast") is a coastal region of Catalonia in northeastern Spain. Whilst sources differ on the exact definition of the Costa Brava, it can be regarded as stretching from the town of Blanes, northeast o ...
.
Vinegar Vinegar is an aqueous solution of acetic acid and trace compounds that may include flavorings. Vinegar typically contains 5–8% acetic acid by volume. Usually, the acetic acid is produced by a double fermentation, converting simple sugars to ...
(3–10% aqueous
acetic acid Acetic acid , systematically named ethanoic acid , is an acidic, colourless liquid and organic compound with the chemical formula (also written as , , or ). Vinegar is at least 4% acetic acid by volume, making acetic acid the main componen ...
) may help with
box jellyfish Box jellyfish (class Cubozoa) are cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latt ...
stings but not the stings of the
Portuguese man o' war The Portuguese man o' war (''Physalia physalis''), also known as the man-of-war, is a marine hydrozoan found in the Atlantic Ocean and the Indian Ocean. It is considered to be the same species as the Pacific man o' war or blue bottle, which is ...
. There is low certainty evidence from one study that treating a Hawaiian box jellyfish sting with vinegar may make the skin appear worse. Salt water may help if vinegar is unavailable. Immersing the sting in hot water may be the most effective way to reduce the pain from a
Physalia ''Physalia'' is a genus of the order Siphonophorae, colonies of four specialized polyps and medusoids that drift on the surface of the Atlantic, Indian and Pacific oceans. Although these organisms look like a single multicellular organism, eac ...
sting. Covering it with an iced pack may help significantly relieve pain as well. Rubbing wounds, or using
alcohol Alcohol most commonly refers to: * Alcohol (chemistry), an organic compound in which a hydroxyl group is bound to a carbon atom * Alcohol (drug), an intoxicant found in alcoholic drinks Alcohol may also refer to: Chemicals * Ethanol, one of sev ...
,
ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogeno ...
, fresh water, or
urine Urine is a liquid by-product of metabolism in humans and in many other animals. Urine flows from the kidneys through the ureters to the urinary bladder. Urination results in urine being excreted from the body through the urethra. Cellular ...
is not advised, as they can encourage the release of more venom. Clearing the area of jelly and tentacles reduces nematocyst firing. Scraping the affected skin, such as with the edge of a credit card, may remove remaining nematocysts. Once the skin has been cleaned of nematocysts,
hydrocortisone Hydrocortisone is the name for the hormone cortisol when supplied as a medication. Uses include conditions such as adrenocortical insufficiency, adrenogenital syndrome, high blood calcium, thyroiditis, rheumatoid arthritis, dermatitis, a ...
cream applied locally reduces pain and inflammation. Antihistamines may help to control itching. Immunobased antivenins are used for serious box jellyfish stings. File:Irukandji-jellyfish-queensland-australia.jpg,
Box jellyfish Box jellyfish (class Cubozoa) are cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in Fresh water, freshwater and Marine habitats, marine environments, predominantly the latt ...
are small and venomous. File:Jellyfish dermatitis case1 abdominal skin lateral.jpg, Jellyfish dermatitis


Mechanical issues

Jellyfish in large quantities can fill and split fishing nets and crush captured fish. They can clog cooling equipment, having disabled
power station A power station, also referred to as a power plant and sometimes generating station or generating plant, is an industrial facility for the generation of electric power. Power stations are generally connected to an electrical grid. Many p ...
s in several countries; jellyfish caused a cascading blackout in the Philippines in 1999, as well as damaging the Diablo Canyon Power Plant in California in 2008. They can also stop
desalination plants Desalination is a process that takes away mineral components from saline water. More generally, desalination refers to the removal of salts and minerals from a target substance, as in soil desalination, which is an issue for agriculture. Salt ...
and ships' engines.


See also

* Jellyfish dermatitis *
List of prehistoric medusozoans This list of prehistoric medusozoan genera is an attempt to create a comprehensive listing of all genera that have ever been included in the subphylum Medusozoa, excluding purely vernacular terms. The list includes all commonly accepted genera, b ...
*
Ocean sunfish The ocean sunfish or common mola (''Mola mola'') is one of the largest bony fish in the world. It was misidentified as the heaviest bony fish, which was actually a different species, '' Mola alexandrini''. Adults typically weigh between . The sp ...
, a significant jellyfish predator


References


Further reading

*


External links


Jellyfish and Comb Jellies – Smithsonian Ocean Portal

Jellyfish Facts – Information on Jellyfish and Jellyfish Safety
*
There's no such thing as a jellyfish
from The MBARI YouTube channel
"Vicious beauties – Jellyfish"
– a documentary about jellyfish
They're Taking Over!
nybooks.com September 26, 2013. Tim Flannery ; Photos
Jellyfish Exhibition At National Aquarium, Baltimore, Maryland (USA) – Photo Gallery
{{Authority control Articles containing video clips . . . Extant Cambrian first appearances