Corals are colonial marine invertebrates within the subphylum Anthozoa of the phylum Cnidaria. They typically form compact colonies of many identical individual polyps. Coral species include the important Coral reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.
A coral "group" is a colony of very many cloning polyps. Each polyp is a sac-like animal typically only a few millimeters in diameter and a few centimeters in height. A set of surround a central mouth opening. Each polyp excretes an exoskeleton near the base. Over many generations, the colony thus creates a skeleton characteristic of the species which can measure up to several meters in size. Individual colonies grow by asexual reproduction of polyps. Corals also breed sexually by spawning: polyps of the same species release simultaneously overnight, often around a full moon. Fertilized eggs form planulae, a mobile early form of the coral polyp which, when mature, settles to form a new colony.
Although some corals are able to catch plankton and small fish using Cnidocyte on their tentacles, most corals obtain the majority of their energy and nutrients from photosynthetic unicellular of the genus Symbiodinium that live within their tissues. These are commonly known as zooxanthellae and give the coral color. Such corals require sunlight and grow in clear, shallow water, typically at depths less than , but corals in the genus Leptoseris have been found as deep as . Corals are major contributors to the physical structure of the that develop in tropical and subtropical waters, such as the Great Barrier Reef off the coast of Australia. These corals are increasingly at risk of coral bleaching events where polyps expel the zooxanthellae in response to stress such as high water temperature or toxins.
Other corals do not rely on zooxanthellae and can live globally in much deeper water, such as the cold-water genus Lophelia which can survive as deep as . Some have been found as far north as the Darwin Mounds, northwest of Cape Wrath, Scotland, and others off the coast of Washington state and the Aleutian Islands.
The Persian polymath Al-Biruni (d.1048) classified sponges and corals as animals, arguing that they respond to touch. Nevertheless, people believed corals to be plants until the eighteenth century when William Herschel used a microscope to establish that coral had the characteristic thin cell membranes of an animal.
Presently, corals are classified as species of animals within the sub-classes Hexacorallia and Octocorallia of the class Anthozoa in the phylum Cnidaria. Hexacorallia includes the stony corals and these groups have polyps that generally have a 6-fold symmetry. Octocorallia includes blue coral and Octocorallia and species of Octocorallia have polyps with an eightfold symmetry, each polyp having eight tentacles and eight mesenteries. The group of corals is paraphyletic because the are also in the sub-class Hexacorallia.
There are two main classifications for corals: hard coral (scleractinian and stony coral) which form reefs by a calcium carbonate base, with polyps that bear six stiff tentacles, and soft coral (Alcyonacea and ahermatypic coral) which are pliable and formed by a colony of polyps with eight feather-like tentacles. These two classifications arose from differentiation in gene expressions in their branch tips and bases that arose through developmental signaling pathways such as Hox gene, Hedgehog, Wnt, and BMP.
Scientists typically select Acropora as research models since they are the most diverse genus of hard coral, having over 120 species. Most species within this genus have polyps which are dimorphic:
Systematic studies of soft coral species have faced challenges due to a lack of taxonomic knowledge. Researchers have not found enough variability within the genus to confidently delineate similar species, due to a low rate in mutation of mitochondrial DNA.
Environmental factors, such as the rise of temperatures and acid levels in our oceans account for some speciation of corals in the form of Species loss. Various coral species have heat shock proteins (HSP) that are also in the category of DE across species. These HSPs help corals combat the increased temperatures they are facing which lead to protein denaturing, growth loss, and eventually coral death. Approximately 33% of coral species are on the International Union for Conservation of Nature's endangered species list and at risk of species loss. Ocean acidification (falling pH levels in the oceans) is threatening the continued species growth and differentiation of corals. Mutation rates of Vibrio shilonii, the reef pathogen responsible for coral bleaching, heavily outweigh the typical reproduction rates of coral colonies when pH levels fall. Thus, corals are unable to mutate their HSPs and other climate change preventative genes to combat the increase in temperature and decrease in pH at a competitive rate to these pathogens responsible for coral bleaching, resulting in species loss.
The body of the polyp may be roughly compared in a structure to a , the wall of which is composed of two layers of cells. The outer layer is known technically as the germ layer, the inner layer as the germ layer. Between ectoderm and endoderm is a supporting layer of gelatinous substance termed mesoglea, secreted by the cell layers of the body wall. The mesoglea can contain endoskeleton elements derived from cells cell migration from the ectoderm.
The sac-like body built up in this way is attached to a hard surface, which in hard corals are cup-shaped depressions in the skeleton known as . At the center of the upper end of the sac lies the only opening called the mouth, surrounded by a circle of which resemble glove fingers. The tentacles are organs which serve both for tactile sense and for the capture of food. Polyps extend their tentacles, particularly at night, often containing coiled stinging cells () which pierce, poison and firmly hold living prey paralyzing or killing them. Polyp prey includes plankton such as copepods and fish larvae. Longitudinal muscular fibers formed from the cells of the ectoderm allow tentacles to contract to convey the food to the mouth. Similarly, circularly disposed muscular fibers formed from the endoderm permit tentacles to be protracted or thrust out once they are contracted. In both stony and soft corals, the polyps can be retracted by contracting muscle fibers, with stony corals relying on their hard skeleton and cnidocytes for defense. Soft corals generally secrete terpenoid toxins to ward off predators.
In most corals, the tentacles are retracted by day and spread out at night to catch plankton and other small organisms. Shallow-water species of both stony and soft corals can be , the corals supplementing their plankton diet with the products of photosynthesis produced by these symbionts. The polyps interconnect by a complex and well-developed system of gastrovascular canals, allowing significant sharing of nutrients and symbionts.
The external form of the polyp varies greatly. The column may be long and slender, or may be so short in the axial direction that the body becomes disk-like. The tentacles may number many hundreds or may be very few, in rare cases only one or two. They may be simple and unbranched, or feathery in pattern. The mouth may be level with the surface of the peristome, or may be projecting and trumpet-shaped.
Soft corals vary considerably in form, and most are colonial. A few soft corals are , but the polyps of most are connected by sheets of tissue called coenosarc, and in some species these sheets are thick and the polyps deeply embedded in them. Some soft corals encrust other sea objects or form lobes. Others are tree-like or whip-like and have a central axial skeleton embedded at their base in the matrix of the supporting branch. These branches are composed of a fibrous protein called gorgonin or of a calcified material.
Coral skeletons are biocomposites (mineral + organics) of calcium carbonate, in the form of calcite or aragonite. In scleractinian corals, "centers of calcification" and fibers are clearly distinct structures differing with respect to both morphology and chemical compositions of the crystalline units. The organic matrices extracted from diverse species are acidic, and comprise proteins, sulphated sugars and lipids; they are species specific. The soluble organic matrices of the skeletons allow to differentiate zooxanthellae and non-zooxanthellae specimens.
The zooxanthellae benefit from a safe place to live and consume the polyp's carbon dioxide, phosphate and nitrogenous waste. Stressed corals will eject their zooxanthellae, a process that is becoming increasingly common due to strain placed on coral by rising ocean temperatures. Mass ejections are known as coral bleaching because the algae contribute to coral coloration; some colors, however, are due to host coral pigments, such as green fluorescent proteins (GFPs). Ejection increases the polyp's chance of surviving short-term stress and if the stress subsides they can regain algae, possibly of a different species, at a later time. If the stressful conditions persist, the polyp eventually dies. Zooxanthellae are located within the coral cytoplasm and due to the algae's photosynthetic activity the internal pH of the coral can be raised; this behavior indicates that the zooxanthellae are responsible to some extent for the metabolism of their host corals. Stony Coral Tissue Loss Disease has been associated with the breakdown of host-zooxanthellae physiology.Landsberg et al., "Stony Coral Tissue Loss Disease in Florida Is Associated With Disruption of Host–Zooxanthellae Physiology". Moreover, Vibrio bacterium are known to have virulence traits used for host coral tissue damage and photoinhibition of algal symbionts.de O Santos et al., "Genomic and Proteomic Analyses of the Coral Pathogen Vibrio Coralliilyticus Reveal a Diverse Virulence Repertoire". Therefore, both coral and their symbiotic microorganisms could have evolved to harbour traits resistant to disease and transmission.
Environmental cues that influence the release of gametes into the water vary from species to species. The cues involve temperature change, lunar cycle, day length, and possibly chemical signalling.
Mass coral spawning often occurs at night on days following a full moon. A full moon is equivalent to four to six hours of continuous dim light exposure, which can cause light-dependent reactions in protein. Corals contain light-sensitive cryptochromes, proteins whose light-absorbing flavin structures are sensitive to different types of light. This allows corals such as Dipsastraea speciosa to detect and respond to changes in sunlight and moonlight.
Moonlight itself may actually suppress coral spawning. The most immediate cue to cause spawning appears to be the dark portion of the night between sunset and moonrise.
Over the lunar cycle, moonrise shifts progressively later, occurring after sunset on the day of the full moon. The resulting dark period between day-light and night-light removes the suppressive effect of moonlight and enables coral to spawn.
The spawning event can be visually dramatic, clouding the usually clear water with gametes. Once released, gametes fertilize at the water's surface and form a microscopic larva called a planula, typically pink and elliptical in shape. A typical coral colony needs to release several thousand larvae per year to overcome the odds against formation of a new colony.
Studies suggest that light pollution desynchronizes spawning in some coral species.
In areas such as the Red Sea, as many as 10 out of 50 species may be showing spawning asynchrony, compared to 30 years ago. The establishment of new corals in the area has decreased and in some cases ceased. The area was previously considered a refuge for corals because mass bleaching events due to climate change had not been observed there. Coral restoration techniques for coral reef management are being developed to increase fertilization rates, larval development, and settlement of new corals.
Budding involves splitting a smaller polyp from an adult. As the new polyp grows, it forms its body parts. The distance between the new and adult polyps grows, and with it, the coenosarc (the common body of the colony). Budding can be intratentacular, from its oral discs, producing same-sized polyps within the ring of tentacles, or extratentacular, from its base, producing a smaller polyp.
Division forms two polyps that each become as large as the original. Longitudinal division begins when a polyp broadens and then divides its coelenteron (body), effectively splitting along its length. The mouth divides and new tentacles form. The two polyps thus created then generate their missing body parts and exoskeleton. Transversal division occurs when polyps and the exoskeleton divide transversally into two parts. This means one has the basal disc (bottom) and the other has the oral disc (top); the new polyps must separately generate the missing pieces.
Asexual reproduction offers the benefits of high reproductive rate, delaying senescence, and replacement of dead modules, as well as geographical distribution.
Studies have also suggested that resident bacteria, archaea, and fungi additionally contribute to nutrient and organic matter cycling within the coral, with viruses also possibly playing a role in structuring the composition of these members, thus providing one of the first glimpses at a multi-domain marine animal symbiosis.Bourne, D.G., Morrow, K.M. and Webster, N.S. (2016) "Insights into the coral microbiome: underpinning the health and resilience of reef ecosystems". Annual Review of Microbiology, 70: 317–340. . The gammaproteobacterium Endozoicomonas is emerging as a central member of the coral's microbiome, with flexibility in its lifestyle.Neave, M.J., Apprill, A., Ferrier-Pagès, C. and Voolstra, C.R. (2016) "Diversity and function of prevalent symbiotic marine bacteria in the genus Endozoicomonas". Applied Microbiology and Biotechnology, 100(19): 8315–8324. .Neave, M.J., Michell, C.T., Apprill, A. and Voolstra, C.R. (2017) "Endozoicomonas genomes reveal functional adaptation and plasticity in bacterial strains symbiotically associated with diverse marine hosts". Scientific Reports, 7: 40579. . Given the recent mass bleaching occurring on reefs,Hughes, T.P., Kerry, J.T., Álvarez-Noriega, M., Álvarez-Romero, J.G., Anderson, K.D., Baird, A.H., Babcock, R.C., Beger, M., Bellwood, D.R., Berkelmans, R. and Bridge, T.C. (2017) "Global warming and recurrent mass bleaching of corals". Nature, 543(7645): 373–377. . corals will likely continue to be a useful and popular system for symbiosis and dysbiosis research.
Astrangia poculata, the northern star coral, is a temperate stony coral, widely documented along the eastern coast of the United States. The coral can live with and without zooxanthellae (algal symbionts), making it an ideal model organism to study microbial community interactions associated with symbiotic state. However, the ability to develop primers and Molecular probe to more specifically target key microbial groups has been hindered by the lack of full-length 16S rRNA sequences, since sequences produced by the Illumina platform are of insufficient length (approximately 250 base pairs) for the design of primers and probes. USGS scientists publish long-read microbiome sequences from temperate coral, providing community resource for probe and primer design, United States Geological Survey, 6 March 2019. Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License. In 2019, Goldsmith et al. demonstrated Sanger sequencing was capable of reproducing the biologically relevant diversity detected by deeper next-generation sequencing, while also producing longer sequences useful to the research community for probe and primer design (see diagram on right). Material was copied from this source, which is available under a Creative Commons Attribution 4.0 International License.
It is known that the coral's microbiome and symbiont influence host health, however, the historic influence of each member on others is not well understood. Scleractinian corals have been diversifying for longer than many other symbiotic systems, and their microbiomes are known to be partially species-specific. It has been suggested that Endozoicomonas, a commonly highly abundant bacterium in corals, has exhibited codiversification with its host. This hints at an intricate set of relationships between the members of the coral holobiont that have been developing as evolution of these members occurs.
A study published in 2018 revealed evidence of phylosymbiosis between corals and their tissue and skeleton microbiomes. The coral skeleton, which represents the most diverse of the three coral microbiomes, showed the strongest evidence of phylosymbiosis. Coral microbiome composition and Species richness were found to reflect coral phylogeny. For example, interactions between bacterial and eukaryotic coral phylogeny influence the abundance of Endozoicomonas, a highly abundant bacterium in the coral holobiont. However, host-microbial cophylogeny appears to influence only a subset of coral-associated bacteria.
There are various types of shallow-water coral reef, including fringing reefs, barrier reefs and atolls; most occur in tropical and subtropical seas. They are very slow-growing, adding perhaps one centimetre (0.4 in) in height each year. The Great Barrier Reef is thought to have been laid down about two million years ago. Over time, corals fragment and die, sand and rubble accumulates between the corals, and the shells of clams and other molluscs decay to form a gradually evolving calcium carbonate structure. Coral reefs are extremely diverse marine hosting over 4,000 species of fish, massive numbers of cnidarians, molluscs, , and many other animals.
Tabulate corals occur in and calcareous of the Ordovician period, with a gap in the fossil record due to extinction events at the end of the Ordovician. Corals reappeared some millions of years later during the Silurian period, and tabulate corals often form low cushions or branching masses of calcite alongside rugose corals. Tabulate coral numbers began to decline during the middle of the Silurian period.
Rugose or horn corals became dominant by the middle of the Silurian period, and during the Devonian, corals flourished with more than 200 genera. The rugose corals existed in solitary and colonial forms, and were also composed of calcite. Both rugose and tabulate corals became extinct in the Permian–Triassic extinction eventXiang-Dong Wang and Xiao-Juan Wang (2007). "Extinction patterns of Late Permian (Lopingian) corals in China", Palaeoworld, 16, No. 1–3, 31–38 (along with 85% of marine species), and there is a gap of tens of millions of years until new forms of coral evolved in the Triassic.
Approximately 10% of the world's coral reefs are dead.Save Our Seas, 1997 Summer Newsletter, Cindy Hunter and Alan Friedlander About 60% of the world's reefs are at risk due to human-related activities. The threat to reef health is particularly strong in Southeast Asia, where 80% of reefs are endangered. Over 50% of the world's may be destroyed by 2030; as a result, most nations protect them through environmental laws.
In the Caribbean and tropical Pacific, direct contact between ~40–70% of common seaweeds and coral causes bleaching and death to the coral via transfer of lipid-soluble . Seaweed and algae proliferate given adequate nutrients and limited grazing by such as parrotfish.
Water temperature changes of more than or salinity changes can kill some species of coral. Under such environmental stresses, corals expel their Symbiodinium; without them, coral tissues reveal the white of their skeletons, an event known as coral bleaching.
Submarine springs found along the coast of Mexico's Yucatán Peninsula produce water with a naturally low pH (relatively high acidity) providing conditions similar to those expected to become widespread as the oceans absorb carbon dioxide. Surveys discovered multiple species of live coral that appeared to tolerate the acidity. The colonies were small and patchily distributed and had not formed structurally complex reefs such as those that compose the nearby Mesoamerican Barrier Reef System.
Annual growth bands in some corals, such as the deep sea ( Isididae), may be among the first signs of the effects of ocean acidification on marine life. The growth rings allow to construct year-by-year chronologies, a form of incremental dating, which underlie high-resolution records of past paleoclimatology and paleoecology changes using geochemical techniques.
Certain species form communities called , which are colonies whose top is dead and mostly above the water line, but whose perimeter is mostly submerged and alive. Average tide level limits their height. By analyzing the various growth morphologies, microatolls offer a low-resolution record of sea level change. Fossilized microatolls can also be dated using radiocarbon dating. Such methods can help to reconstruct Holocene .
Though coral have large sexually-reproducing populations, their evolution can be slowed by abundant asexual reproduction. Gene flow is variable among coral species. According to the biogeography of coral species, gene flow cannot be counted on as a dependable source of adaptation as they are very stationary organisms. Also, coral longevity might factor into their adaptivity.
However, adaptation to climate change has been demonstrated in many cases, which is usually due to a shift in coral and zooxanthellae . These shifts in allele frequency have progressed toward more tolerant types of zooxanthellae. Scientists found that a certain zooxanthella is becoming more common where sea temperature is high. Symbionts able to tolerate warmer water seem to photosynthesise more slowly, implying an evolutionary trade-off.
In the Gulf of Mexico, where sea temperatures are rising, cold-sensitive Staghorn coral and elkhorn coral have shifted in location.
Not only have the symbionts and specific species been shown to shift, but there seems to be a certain growth rate favorable to selection. Slower-growing but more heat-tolerant corals have become more common. The changes in temperature and acclimation are complex. Some reefs in current shadows represent a refugium location that will help them adjust to the disparity in the environment even if eventually the temperatures may rise more quickly there than in other locations. This vicariance by climatic barriers causes a realized niche to shrink greatly in comparison to the old fundamental niche.
Geochemical analysis of skeletal coral can be linked to sea surface salinity (SSS) and sea surface temperature (SST), from El Nino 3.4 SSTA data, of tropical oceans to seawater δ18O ratio anomalies from corals. ENSO phenomenon can be related to variations in sea surface salinity (SSS) and sea surface temperature (SST) that can help model tropical climate activities.
Many governments now prohibit removal of coral from reefs, and inform coastal residents about reef protection and ecology. While local action such as habitat restoration and herbivore protection can reduce local damage, the longer-term threats of acidification, temperature change and sea-level rise remain a challenge.
Protecting networks of diverse and healthy reefs, not only climate refugia, helps ensure the greatest chance of genetic diversity, which is critical for coral to adapt to new climates. A variety of conservation methods applied across marine and terrestrial threatened ecosystems makes coral adaption more likely and effective.
To eliminate destruction of corals in their indigenous regions, projects have been started to grow corals in non-tropical countries. EcoDeco EcologicalTechnology . Ecodeco.nl. Retrieved on 2011-11-29. KoralenKAS project . Koraalwetenschap.nl. Retrieved on 2011-11-29.
Always considered a precious mineral, "the Chinese have long associated red coral with auspiciousness and longevity because of its color and its resemblance to deer antlers (so by association, virtue, long life, and high rank".Welch, Patricia Bjaaland, Chinese Art: A Guide to Motifs and Visual Imagery. Tokyo, Rutland and Singapore: Tuttle, 2008, p. 61 It reached its height of popularity during the Manchu or Qing Dynasty (1644–1911) when it was almost exclusively reserved for the emperor's use either in the form of coral beads (often combined with pearls) for court jewelry or as decorative Penjing (decorative miniature mineral trees). Coral was known as shanhu in Chinese. The "early-modern 'coral network' began the Mediterranean Sea and to Qing China via the English East India Company".Lacey, Pippa, "The Coral Network: The trade of red coral to the Qing imperial court in the eighteenth century" in The Global Lives of Things, ed. by Anne Gerritsen and Giorgio Aiello, London: Rutledge, 2016, p. 81 There were strict rules regarding its use in a code established by the Qianlong Emperor in 1759.
By the middle of the 19th century, “coral fisheries” existed in the Mediterranean, the Red Sea, Persian Gulf and elsewhere. An instrument used to retrieve the coral consisted of two beams of heavy wood attached to each other at right angles. Heavy stones were added to make the apparatus sink and netting was attached beneath the device. It is then lowered by a strong rope over an outcrop of coral and the boat trawls over the coral causing it to break off and be caught in the netting. The device is then drawn to the surface by the boat crew.
The most popular kind of coral kept is soft coral, especially and mushroom corals, which are especially easy to grow and propagate in a wide variety of conditions, because they originate in enclosed parts of reefs where water conditions vary and lighting may be less reliable and direct. Coral Reefs . Marinebio.org. Retrieved on 2016-06-13. More serious fishkeepers may keep small polyp stony coral, which is from open, brightly lit reef conditions and therefore much more demanding, while large polyp stony coral is a sort of compromise between the two.
Soft corals
Stony corals
Ecology
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Intracellular symbionts
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Sexual
Broadcasters
Brooders
Planulae
Asexual
Colony division
Coral microbiomes
Holobionts
Reefs
Evolution
Early corals
Modern corals
Status
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Coral health
Climate change impacts
Geochemistry
Strontium/calcium ratio anomaly
Oxygen isotope anomaly
Sea surface temperature and sea surface salinity
Limited climate research on current species
Protection
Relation to humans
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