A diatom (Neo-Latin diatoma) is any member of a large group comprising several genera of algae, specifically microalgae, found in the oceans, waterways and soils of the world. Living diatoms make up a significant portion of Earth's biomass. They generate about 20 to 50 percent of the oxygen produced on the planet each year, take in over 6.7 billion tonnes of silicon each year from the waters in which they live, and constitute nearly half of the organic material found in the oceans. The shells of dead diatoms are a significant component of marine sediment, and the entire Amazon basin is fertilized annually by 27 million tons of diatom shell dust transported by transatlantic winds from the African Sahara, much of it from the Bodélé Depression, which was once made up of a system of fresh-water lakes. Diatoms are unicellular organisms: they occur either as solitary cells or in colonies, which can take the shape of ribbons, fans, zigzags, or stars. Individual cells range in size from 2 to 2000 micrometers. In the presence of adequate nutrients and sunlight, an assemblage of living diatoms doubles approximately every 24 hours by asexual multiple fission; the maximum life span of individual cells is about six days. Diatoms have two distinct shapes: a few (centric diatoms) are radially symmetric, while most (pennate diatoms) are broadly bilaterally symmetric. About 12,000 species of diatoms have been documented, but it is thought many more have not been documented. The unique feature of diatoms is that they are surrounded by a cell wall made of silica (hydrated silicon dioxide), called a frustule. These frustules produce structural coloration, prompting them to be described as "jewels of the sea" and "living opals". Movement in diatoms primarily occurs passively as a result of both ocean currents and wind-induced water turbulence; however, male gametes of centric diatoms have flagella, permitting active movement to seek female gametes. Similar to plants, diatoms convert light energy to chemical energy by photosynthesis, but their chloroplasts were acquired in different ways. Unusually for autotrophic organisms, diatoms possess a urea cycle, a feature that they share with animals, although this cycle is used to different metabolic ends in diatoms. The family Rhopalodiaceae also possess a cyanobacterial endosymbiont called a spheroid body. This endosymbiont has lost its photosynthetic properties, but has kept its ability to perform nitrogen fixation, allowing the diatom to fix atmospheric nitrogen. Other diatoms in symbiosis with nitrogen-fixing cyanobacteria are among the genera Hemiaulus, Rhizosolenia and Chaetoceros. Dinotoms are dinoflagellates that have diatom endosymbionts inside them. Research on the dinoflagellates Durinskia baltica and Glenodinium foliaceum has shown that the endosymbiont event happened so recently, evolutionarily speaking, that their organelles and genome are still intact with minimal to no gene loss. The main difference between these and free living diatoms is that they have lost their cell wall of silica, making them the only known shell-less diatoms. The study of diatoms is a branch of phycology. Diatoms are classified as eukaryotes, organisms with a nuclear envelope-bound cell nucleus, that separates them from the prokaryotes archaea and bacteria. Diatoms are a type of plankton called phytoplankton, the most common of the plankton types. Diatoms also grow attached to benthic substrates, floating debris, and on macrophytes. They comprise an integral component of the periphyton community. Another classification divides plankton into eight types based on size: in this scheme, diatoms are classed as microalgae. Several systems for classifying the individual diatom species exist. Fossil evidence suggests that diatoms originated during or before the early Jurassic period, which was about 150 to 200 million years ago. The oldest fossil evidence for diatoms is a specimen of extant genus Hemiaulus in Late Jurassic aged amber from Thailand. Diatoms are used to monitor past and present environmental conditions, and are commonly used in studies of water quality. Diatomaceous earth (diatomite) is a collection of diatom shells found in the Earth's crust. They are soft, silica-containing sedimentary rocks which are easily crumbled into a fine powder and typically have a particle size of 10 to 200 μm. Diatomaceous earth is used for a variety of purposes including for water filtration, as a mild abrasive, in cat litter, and as a dynamite stabilizer.
Overview Diatoms are protists that form massive annual spring and fall blooms in aquatic environments and are estimated to be responsible for about half of photosynthesis in the global oceans. This predictable annual bloom dynamic fuels higher trophic levels and initiates delivery of carbon into the deep ocean biome. Diatoms have complex life history strategies that are presumed to have contributed to their rapid genetic diversification into ~200,000 species that are distributed between the two major diatom groups: centrics and pennates.
Morphology
Diatoms are generally 20 to 200 micrometers in size, with a few larger species. Their yellowish-brown chloroplasts, the site of photosynthesis, are typical of heterokonts, having four cell membranes and containing pigments such as the carotenoid fucoxanthin. Individuals usually lack flagella, but they are present in male gametes of the centric diatoms and have the usual heterokont structure, including the hairs (mastigonemes) characteristic in other groups. Diatoms are often referred as "jewels of the sea" or "living opals" due to their optical properties. The biological function of this structural coloration is not clear, but it is speculated that it may be related to communication, camouflage, thermal exchange and/or UV protection. Diatoms build intricate hard but porous cell walls called frustules composed primarily of silica. This siliceous wall can be highly patterned with a variety of pores, ribs, minute spines, marginal ridges and elevations; all of which can be used to delineate genera and species. The cell itself consists of two halves, each containing an essentially flat plate, or valve, and marginal connecting, or girdle band. One half, the hypotheca, is slightly smaller than the other half, the epitheca. Diatom morphology varies. Although the shape of the cell is typically circular, some cells may be triangular, square, or elliptical. Their distinguishing feature is a hard mineral shell or frustule composed of opal (hydrated, polymerized silicic acid). Diatoms are divided into two groups that are distinguished by the shape of the frustule: the centric diatoms and the pennate diatoms. Pennate diatoms are bilaterally symmetric. Each one of their valves have openings that are slits along the raphes and their shells are typically elongated parallel to these raphes. They generate cell movement through cytoplasm that streams along the raphes, always moving along solid surfaces. Centric diatoms are radially symmetric. They are composed of upper and lower valves – epitheca and hypotheca – each consisting of a valve and a girdle band that can easily slide underneath each other and expand to increase cell content over the diatoms progression. The cytoplasm of the centric diatom is located along the inner surface of the shell and provides a hollow lining around the large vacuole located in the center of the cell. This large, central vacuole is filled by a fluid known as "cell sap" which is similar to seawater but varies with specific ion content. The cytoplasmic layer is home to several organelles, like the chloroplasts and mitochondria. Before the centric diatom begins to expand, its nucleus is at the center of one of the valves and begins to move towards the center of the cytoplasmic layer before division is complete. Centric diatoms have a variety of shapes and sizes, depending on from which axis the shell extends, and if spines are present.
Silicification
Diatom cells are contained within a unique silica cell wall known as a frustule made up of two valves called thecae, that typically overlap one another. The biogenic silica composing the cell wall is synthesised intracellularly by the polymerisation of silicic acid monomers. This material is then extruded to the cell exterior and added to the wall. In most species, when a diatom divides to produce two daughter cells, each cell keeps one of the two valves and grows a smaller valve within it. As a result, after each division cycle, the average size of diatom cells in the population gets smaller. Once such cells reach a certain minimum size, rather than simply divide, they reverse this decline by forming an auxospore, usually through meiosis and sexual reproduction, but exceptions exist. The auxospore expands in size to give rise to a much larger cell, which then returns to size-diminishing divisions.
The exact mechanism of transferring silica absorbed by the diatom to the cell wall is unknown. Much of the sequencing of diatom genes comes from the search for the mechanism of silica uptake and deposition in nano-scale patterns in the frustule. The most success in this area has come from two species, Thalassiosira pseudonana, which has become the model species, as the whole genome was sequenced and methods for genetic control were established, and Cylindrotheca fusiformis, in which the important silica deposition proteins silaffins were first discovered. Silaffins, sets of polycationic peptides, were found in C. fusiformis cell walls and can generate intricate silica structures. These structures demonstrated pores of sizes characteristic to diatom patterns. When T. pseudonana underwent genome analysis it was found that it encoded a urea cycle, including a higher number of polyamines than most genomes, as well as three distinct silica transport genes. In a phylogenetic study on silica transport genes from 8 diverse groups of diatoms, silica transport was found to generally group with species. This study also found structural differences between the silica transporters of pennate (bilateral symmetry) and centric (radial symmetry) diatoms. The sequences compared in this study were used to create a diverse background in order to identify residues that differentiate function in the silica deposition process. Additionally, the same study found that a number of the regions were conserved within species, likely the base structure of silica transport. These silica transport proteins are unique to diatoms, with no homologs found in other species, such as sponges or rice. The divergence of these silica transport genes is also indicative of the structure of the protein evolving from two repeated units composed of five membrane bound segments, which indicates either gene duplication or dimerization. The silica deposition that takes place from the membrane bound vesicle in diatoms has been hypothesized to be a result of the activity of silaffins and long chain polyamines. This Silica Deposition Vesicle (SDV) has been characterized as an acidic compartment fused with Golgi-derived vesicles. These two protein structures have been shown to create sheets of patterned silica in-vivo with irregular pores on the scale of diatom frustules. One hypothesis as to how these proteins work to create complex structure is that residues are conserved within the SDV's, which is unfortunately difficult to identify or observe due to the limited number of diverse sequences available. Though the exact mechanism of the highly uniform deposition of silica is as yet unknown, the Thalassiosira pseudonana genes linked to silaffins are being looked to as targets for genetic control of nanoscale silica deposition. The ability of diatoms to make silica-based cell walls has been the subject of fascination for centuries. It started with a microscopic observation by an anonymous English country nobleman in 1703, who observed an object that looked like a chain of regular parallelograms and debated whether it was just crystals of salt, or a plant. The viewer decided that it was a plant because the parallelograms didn't separate upon agitation, nor did they vary in appearance when dried or subjected to warm water (in an attempt to dissolve the "salt"). Unknowingly, the viewer's confusion captured the essence of diatoms—mineral utilizing plants. It is not clear when it was determined that diatom cell walls are made of silica, but in 1939 a seminal reference characterized the material as silicic acid in a "subcolloidal" state Identification of the main chemical component of the cell wall spurred investigations into how it was made. These investigations have involved, and been propelled by, diverse approaches including, microscopy, chemistry, biochemistry, material characterisation, molecular biology, 'omics, and transgenic approaches. The results from this work have given a better understanding of cell wall formation processes, establishing fundamental knowledge which can be used to create models that contextualise current findings and clarify how the process works. The process of building a mineral-based cell wall inside the cell, then exporting it outside, is a massive event that must involve large numbers of genes and their protein products. The act of building and exocytosing this large structural object in a short time period, synched with cell cycle progression, necessitates substantial physical movements within the cell as well as dedication of a significant proportion of the cell's biosynthetic capacities. The first characterisations of the biochemical processes and components involved in diatom silicification were made in the late 1990s. These were followed by insights into how higher order assembly of silica structures might occur. More recent reports describe the identification of novel components involved in higher order processes, the dynamics documented through real-time imaging, and the genetic manipulation of silica structure. The approaches established in these recent works provide practical avenues to not only identify the components involved in silica cell wall formation but to elucidate their interactions and spatio-temporal dynamics. This type of holistic understanding will be necessary to achieve a more complete understanding of cell wall synthesis.
Behaviour
Most centric and araphid pennate diatoms are nonmotile, and their relatively dense cell walls cause them to readily sink. Planktonic forms in open water usually rely on turbulent mixing of the upper layers of the oceanic waters by the wind to keep them suspended in sunlit surface waters. Many planktonic diatoms have also evolved features that slow their sinking rate, such as spines or the ability to grow in colonial chains. These adaptations increase their surface area to volume ratio and drag, allowing them to stay suspended in the water column longer. Individual cells may regulate buoyancy via an ionic pump. Some pennate diatoms are capable of a type of locomotion called "gliding", which allows them to move across surfaces via adhesive mucilage secreted through a seamlike structure called the raphe. In order for a diatom cell to glide, it must have a solid substrate for the mucilage to adhere to. Cells are solitary or united into colonies of various kinds, which may be linked by siliceous structures; mucilage pads, stalks or tubes; amorphous masses of mucilage; or by threads of chitin (polysaccharide), which are secreted through strutted processes of the cell.
Phytochromes Even though light is a crucial part of how diatoms create oxygen for the planet, the organism faces some difficulties when it comes to detecting its energy source. The intensity of light in water lessens as depth increases. Light penetration also greatly differs between coastal and open waters and during the changing seasons. Such factors result in a less efficient photosynthetic conversion, similar to the process of plant photosynthesis, as the light becomes dimmer, photosynthesis slows down. However, diatoms possess photoreceptors, which are light-activated proteins, that aid them in sensing different light wavelengths, such as red light and far-red light, for detecting light in the ocean.
It has been demonstrated that diatoms use photoreceptors called phytochromes to determine the water's depth to respond to light signals. Phytochromes can sense red and far-red light and are widely known to be found in both plants and phytoplankton. These proteins switch between two states called red-light and far red-light so that the organism can detect and respond to any changes in the perceived underwater light intensity and spectrum. Since red and far-red light is known to diminish with increasing water depth, many questioned the importance of the phytochromes' role when it comes to marine life. Analysis of environmental DNA sequences taken from the Tara Oceans expedition, as well as the genome data from cultured diatoms which demonstrated that the phytochrome-encoding genes were mostly found in diatoms living in temperate and polar regions in mid-to-high latitudes but such diatom phytochrome genes were not found in diatoms living in tropical waters. Laboratory experiments with the diatom Phaeodactylum tricornutum demonstrated how the diatom phytochromes react to light. Using a yellow fluorescent protein gene controlled by phytochromes inserted into the diatom enabled its activity in simulations with deep-water conditions to be tracked. This showed that diatoms had developed a reduced sensitivity to far-red light, as well as an increased sensitivity to low-intensity blue and green light, which are more present at greater depths. Removal of the phytochrome gene from the diatom Thalassiosira pseudonana grown in a similar deep-water simulation demonstrated that the mutant diatom had a lower photosynthetic efficiency, as well as a reduced photoprotection, compared to the wild-type diatoms with the phytochrome gene, and that when both the mutant and wild-type diatoms were exposed to high white light, there was no difference in reactions. With these findings, the authors found that diatom phytochromes respond more to blue and green light in low intensities, unlike the plant phytochromes that respond to red and far-red light. They suggest that the diatom phytochromes went through an evolutionary adaptation to acclimate to the violent waters in the open waters of the temperate and polar regions. Since the function of diatom phytochromes is to sense the water depth, it provides the diatoms information that is very advantageous in regions with differing seasons. These photoreceptors play a critical role in helping phytoplankton adjust to environments with limited light, specifically the deep-water environments.
Life cycle
Reproduction and cell size Reproduction among these organisms is asexual by binary fission, during which the diatom divides into two parts, producing two "new" diatoms with identical genes. Each new organism receives one of the two frustules – one larger, the other smaller – possessed by the parent, which is now called the epitheca; and is used to construct a second, smaller frustule, the hypotheca. The diatom that received the larger frustule becomes the same size as its parent, but the diatom that received the smaller frustule remains smaller than its parent. This causes the average cell size of this diatom population to decrease. It has been observed, however, that certain taxa have the ability to divide without causing a reduction in cell size. Nonetheless, in order to restore the cell size of a diatom population for those that do endure size reduction, sexual reproduction and auxospore formation must occur.
Cell division Vegetative cells of diatoms are diploid (2N) and so meiosis can take place, producing male and female gametes which then fuse to form the zygote. The zygote sheds its silica theca and grows into a large sphere covered by an organic membrane, the auxospore. A new diatom cell of maximum size, the initial cell, forms within the auxospore thus beginning a new generation. Resting spores may also be formed as a response to unfavourable environmental conditions with germination occurring when conditions improve. A defining characteristic of all diatoms is their restrictive and bipartite silica cell wall that causes them to progressively shrink during asexual cell division. At a critically small cell size and under certain conditions, auxosporulation restitutes cell size and prevents clonal death. The entire lifecycles of only a few diatoms have been described and rarely have sexual events been captured in the environment.
Sexual reproduction Most eukaryotes are capable of sexual reproduction involving meiosis. Sexual reproduction appears to be an obligatory phase in the life cycle of diatoms, particularly as cell size decreases with successive vegetative divisions. Sexual reproduction involves production of gametes and the fusion of gametes to form a zygote in which maximal cell size is restored. The signaling that triggers the sexual phase is favored when cells accumulate together, so that the distance between them is reduced and the contacts and/or the perception of chemical cues is facilitated. An exploration of the genomes of five diatoms and one diatom transcriptome led to the identification of 42 genes potentially involved in meiosis. Thus a meiotic toolkit appears to be conserved in these six diatom species, indicating a central role of meiosis in diatoms as in other eukaryotes.
Sperm motility Diatoms are mostly non-motile; however, sperm found in some species can be flagellated, though motility is usually limited to a gliding motion. In centric diatoms, the small male gametes have one flagellum while the female gametes are large and non-motile (oogamous). Conversely, in pennate diatoms both gametes lack flagella (isogamous). Certain araphid species, that is pennate diatoms without a raphe (seam), have been documented as anisogamous and are, therefore, considered to represent a transitional stage between centric and raphid pennate diatoms, diatoms with a raphe.
Degradation by microbes Certain species of bacteria in oceans and lakes can accelerate the rate of dissolution of silica in dead and living diatoms by using hydrolytic enzymes to break down the organic algal material.
Ecology
Distribution Diatoms are a widespread group and can be found in the oceans, in fresh water, in soils, and on damp surfaces. They are one of the dominant components of phytoplankton in nutrient-rich coastal waters and during oceanic spring blooms, since they can divide more rapidly than other groups of phytoplankton. Most live pelagically in open water, although some live as surface films at the water-sediment interface (benthic), or even under damp atmospheric conditions. They are especially important in oceans, where a 2003 study found that they contribute an estimated 45% of the total oceanic primary production of organic material. However, a more recent 2016 study estimates that the number is closer to 20%. Spatial distribution of marine phytoplankton species is restricted both horizontally and vertically.
Growth Planktonic diatoms in freshwater and marine environments typically exhibit a "boom and bust" (or "bloom and bust") lifestyle. When conditions in the upper mixed layer (nutrients and light) are favourable (as at the spring), their competitive edge and rapid growth rate enables them to dominate phytoplankton communities ("boom" or "bloom"). As such they are often classed as opportunistic r-strategists (i.e. those organisms whose ecology is defined by a high growth rate, r).
Impact The freshwater diatom Didymosphenia geminata, commonly known as Didymo, causes severe environmental degradation in water-courses where it blooms, producing large quantities of a brown jelly-like material called "brown snot" or "rock snot". This diatom is native to Europe and is an invasive species both in the antipodes and in parts of North America. The problem is most frequently recorded from Australia and New Zealand. When conditions turn unfavourable, usually upon depletion of nutrients, diatom cells typically increase in sinking rate and exit the upper mixed layer ("bust"). This sinking is induced by either a loss of buoyancy control, the synthesis of mucilage that sticks diatoms cells together, or the production of heavy resting spores. Sinking out of the upper mixed layer removes diatoms from conditions unfavourable to growth, including grazer populations and higher temperatures (which would otherwise increase cell metabolism). Cells reaching deeper water or the shallow seafloor can then rest until conditions become more favourable again. In the open ocean, many sinking cells are lost to the deep, but refuge populations can persist near the thermocline. Ultimately, diatom cells in these resting populations re-enter the upper mixed layer when vertical mixing entrains them. In most circumstances, this mixing also replenishes nutrients in the upper mixed layer, setting the scene for the next round of diatom blooms. In the open ocean (away from areas of continuous upwelling), this cycle of bloom, bust, then return to pre-bloom conditions typically occurs over an annual cycle, with diatoms only being prevalent during the spring and early summer. In some locations, however, an autumn bloom may occur, caused by the breakdown of summer stratification and the entrainment of nutrients while light levels are still sufficient for growth. Since vertical mixing is increasing, and light levels are falling as winter approaches, these blooms are smaller and shorter-lived than their spring equivalents. In the open ocean, the diatom (spring) bloom is typically ended by a shortage of silicon. Unlike other minerals, the requirement for silicon is unique to diatoms and it is not regenerated in the plankton ecosystem as efficiently as, for instance, nitrogen or phosphorus nutrients. This can be seen in maps of surface nutrient concentrations – as nutrients decline along gradients, silicon is usually the first to be exhausted (followed normally by nitrogen then phosphorus). Because of this bloom-and-bust cycle, diatoms are believed to play a disproportionately important role in the export of carbon from oceanic surface waters (see also the biological pump). Significantly, they also play a key role in the regulation of the biogeochemical cycle of silicon in the modern ocean.
Reason for success Diatoms are ecologically successful, and occur in virtually every environment that contains water – not only oceans, seas, lakes, and streams, but also soil and wetlands. The use of silicon by diatoms is believed by many researchers to be the key to this ecological success. Raven (1983) noted that, relative to organic cell walls, silica frustules require less energy to synthesize (approximately 8% of a comparable organic wall), potentially a significant saving on the overall cell energy budget. In a now classic study, Egge and Aksnes (1992) found that diatom dominance of mesocosm communities was directly related to the availability of silicic acid – when concentrations were greater than 2 μmol m−3, they found that diatoms typically represented more than 70% of the phytoplankton community. Other researchers have suggested that the biogenic silica in diatom cell walls acts as an effective pH buffering agent, facilitating the conversion of bicarbonate to dissolved CO2 (which is more readily assimilated). More generally, whether or not by reason of these possible advantages conferred by their use of silicon, diatoms typically have higher growth rates than other algae of the same corresponding size.
Sources for collection
Diatoms can be obtained from multiple sources. Marine diatoms can be collected by direct water sampling, and benthic forms can be secured by scraping barnacles, oyster and other shells. Diatoms are frequently present as a brown, slippery coating on submerged stones and sticks, and may be seen to "stream" with river current. The surface mud of a pond, ditch, or lagoon will almost always yield some diatoms. Living diatoms are often found clinging in great numbers to filamentous algae, or forming gelatinous masses on various submerged plants. Cladophora is frequently covered with Cocconeis, an elliptically shaped diatom; Vaucheria is often covered with small forms. Since diatoms form an important part of the food of molluscs, tunicates, and fishes, the alimentary tracts of these animals often yield forms that are not easily secured in other ways. Diatoms can be made to emerge by filling a jar with water and mud, wrapping it in black paper and letting direct sunlight fall on the surface of the water. Within a day, the diatoms will come to the top in a scum and can be isolated.
Biogeochemistry
Silica cycle
The diagram shows the major fluxes of silicon in the current ocean. Most biogenic silica in the ocean (silica produced by biological activity) comes from diatoms. Diatoms extract dissolved silicic acid from surface waters as they grow, and return it to the water column when they die. Inputs of silicon arrive from above via aeolian dust, from the coasts via rivers, and from below via seafloor sediment recycling, weathering, and hydrothermal activity. Although diatoms may have existed since the Triassic, the timing of their ascendancy and "take-over" of the silicon cycle occurred more recently. Prior to the Phanerozoic (before 544 Ma), it is believed that microbial or inorganic processes weakly regulated the ocean's silicon cycle. Subsequently, the cycle appears dominated (and more strongly regulated) by the radiolarians and siliceous sponges, the former as zooplankton, the latter as sedentary filter-feeders primarily on the continental shelves. Within the last 100 My, it is thought that the silicon cycle has come under even tighter control, and that this derives from the ecological ascendancy of the diatoms. However, the precise timing of the "take-over" remains unclear, and different authors have conflicting interpretations of the fossil record. Some evidence, such as the displacement of siliceous sponges from the shelves, suggests that this takeover began in the Cretaceous (146 Ma to 66 Ma), while evidence from radiolarians suggests "take-over" did not begin until the Cenozoic (66 Ma to present).
Carbon cycle The diagram depicts some mechanisms by which marine diatoms contribute to the biological carbon pump and influence the ocean carbon cycle. The anthropogenic CO2 emission to the atmosphere (mainly generated by fossil fuel burning and deforestation) is nearly 11 gigatonne carbon (GtC) per year, of which almost 2.5 GtC is taken up by the surface ocean. In surface seawater (pH 8.1–8.4), bicarbonate (HCO−3) and carbonate ions (CO2−3) constitute nearly 90 and <10% of dissolved inorganic carbon (DIC) respectively, while dissolved CO2 (CO2 aqueous) contributes <1%. Despite this low level of CO2 in the ocean and its slow diffusion rate in water, diatoms fix 10–20 GtC annually via photosynthesis thanks to their carbon dioxide concentrating mechanisms, allowing them to sustain marine food chains. In addition, 0.1–1% of this organic material produced in the euphotic layer sinks down as particles, thus transferring the surface carbon toward the deep ocean and sequestering atmospheric CO2 for thousands of years or longer. The remaining organic matter is remineralized through respiration. Thus, diatoms are one of the main players in this biological carbon pump, which is arguably the most important biological mechanism in the Earth System allowing CO2 to be removed from the carbon cycle for very long period.
Urea cycle A feature of diatoms is the urea cycle, which is a feature shared with animals. In 2011, Allen et al. established that diatoms have a functioning urea cycle. This result was significant, since prior to this, the urea cycle was thought to have originated with the metazoans, which appeared several hundreds of millions of years before the diatoms. Their study demonstrated that while diatoms and animals use the urea cycle for different ends, they are seen to share this feature that more closely related organisms such as plants lack. While often overlooked in photosynthetic organisms, the mitochondria also play critical roles in energy balance. Two nitrogen-related pathways are relevant and they may also change under ammonium (NH+4) nutrition compared with nitrate (NO−3) nutrition. First, in diatoms, and likely some other algae, there is a urea cycle. The long-known function of the urea cycle in animals is to excrete excess nitrogen produced by amino acid Catabolism; like photorespiration, the urea cycle had long been considered a waste pathway. However, in diatoms the urea cycle appears to play a role in exchange of nutrients between the mitochondria and the cytoplasm, and potentially the plastid and may help to regulate ammonium metabolism. Because of this cycle, marine diatoms, in contrast to chlorophytes, also have acquired a mitochondrial urea transporter and, in fact, based on bioinformatics, a complete mitochondrial GS-GOGAT cycle has been hypothesised.
Other Diatoms are mainly photosynthetic; however a few are obligate heterotrophs and can live in the absence of light provided an appropriate organic carbon source is available. Photosynthetic diatoms that find themselves in an environment absent of oxygen and/or sunlight can switch to anaerobic respiration known as nitrate respiration (DNRA), and stay dormant for up till months and decades. Major pigments of diatoms are chlorophylls a and c, beta-carotene, fucoxanthin, diatoxanthin and diadinoxanthin.
Taxonomy
Diatoms belong to a large group of protists, many of which contain plastids rich in chlorophylls a and c. The group has been variously referred to as heterokonts, chrysophytes, chromists or stramenopiles. Many are autotrophs such as golden algae and kelp; and heterotrophs such as water moulds, opalinids, and actinophryid heliozoa. The classification of this area of protists is still unsettled. In terms of rank, they have been treated as a division, phylum, kingdom, or something intermediate to those. Consequently, diatoms are ranked anywhere from a class, usually called Diatomophyceae or Bacillariophyceae, to a division (=phylum), usually called Bacillariophyta, with corresponding changes in the ranks of their subgroups.
Genera and species An estimated 20,000 extant diatom species are believed to exist, of which around 12,000 have been named to date according to Guiry, 2012 (other sources give a wider range of estimates). Around 1,000–1,300 diatom genera have been described, both extant and fossil, of which some 250–300 exist only as fossils.
Classes and orders For many years the diatoms—treated either as a class (Bacillariophyceae) or a phylum (Bacillariophyta)—were divided into just 2 orders, corresponding to the centric and the pennate diatoms (Centrales and Pennales). This classification was extensively overhauled by Round, Crawford and Mann in 1990 who treated the diatoms at a higher rank (division, corresponding to phylum in zoological classification), and promoted the major classification units to classes, maintaining the centric diatoms as a single class Coscinodiscophyceae, but splitting the former pennate diatoms into 2 separate classes, Fragilariophyceae and Bacillariophyceae (the latter older name retained but with an emended definition), between them encompassing 45 orders, the majority of them new. As of 2020, it is recognised that the 1990 system of Round et al. is in need of revision with the advent of newer molecular work, however the best system to replace it is unclear. Current systems in widespread use, such as AlgaeBase, the World Register of Marine Species, and its contributing database DiatomBase, and the system for "all life" represented in Ruggiero et al., 2015, all retain the Round et al. treatment as their basis, albeit with diatoms as a whole treated as a class rather than division/phylum, with Round et al.'s classes reduced to subclasses, for better agreement with the treatment of phylogenetically adjacent groups and their containing taxa. (For references refer the individual sections below). One proposal, by Linda Medlin and co-workers commencing in 2004, considers some of the centric diatoms more closely related to the pennates to be split off as a new class, Mediophyceae, itself more closely aligned with the pennate diatoms than the remaining centrics. This hypothesis—later designated the Coscinodiscophyceae-Mediophyceae-Bacillariophyceae, or Coscinodiscophyceae+(Mediophyceae+Bacillariophyceae) (CMB) hypothesis—has been accepted by D.G. Mann among others, who use it as the basis for the classification of diatoms as presented in Adl. et al.'s series of syntheses (2005, 2012, 2019), and also in the Bacillariophyta chapter of the 2017 Handbook of the Protists edited by Archibald et al., with some modifications reflecting the apparent non-monophyly of Medlin et al. original "Coscinodiscophyceae". Meanwhile, a group led by E.C. Theriot favours a different hypothesis of phylogeny, which has been termed the structural gradation hypothesis (SGH) and does not recognise the Mediophyceae as a monophyletic group. Another analysis, by Parks et al., 2018, finds that the radial centric diatoms (Medlin et al.'s Coscinodiscophyceae) are not monophyletic, but supports the monophyly of Mediophyceae minus Attheya, which is an anomalous genus. Discussion of the relative merits of these conflicting schemes continues by the various parties involved.
Adl et al., 2019 treatment In 2019, Adl et al. presented the following classification of diatoms, while noting: "This revision reflects numerous advances in the phylogeny of the diatoms over the last decade. Due to our poor taxon sampling outside of the Mediophyceae and pennate diatoms, and the known and anticipated diversity of all diatoms, many clades appear at a high classification level (and the higher level classification is rather flat)." This classification accepts the Mediophyceae of Medlin and co-workers and introduces new clades for a number of otherwise isolated genera.
Ochrophyta Cavalier-Smith 1986, emend. Cavalier-Smith & Chao 1996 (ochrophytes) Diatomista Derelle et al. 2016, emend. Cavalier-Smith 2017 (diatoms plus a subset of other ochrophyte groups) Diatomeae Dumortier 1821 [= Bacillariophyta Haeckel 1878] (diatoms) Leptocylindrophytina D.G. Mann in Adl et al. 2019 Leptocylindrophyceae D.G. Mann in Adl et al. 2019 (Leptocylindrus, Tenuicylindrus) Corethrophyceae D.G. Mann in Adl et al. 2019 (Corethron) Ellerbeckiophytina D.G. Mann in Adl et al. 2019 (Ellerbeckia) Probosciophytina D.G. Mann in Adl et al. 2019 (Proboscia) Melosirophytina D.G. Mann in Adl et al. 2019 (Aulacoseira, Melosira, Hyalodiscus, Stephanopyxis, Paralia, Endictya) Coscinodiscophytina Medlin & Kaczmarska 2004, emend. (Actinoptychus, Coscinodiscus, Actinocyclus, Asteromphalus, Aulacodiscus, Stellarima) Rhizosoleniophytina D.G. Mann in Adl et al. 2019 (Guinardia, Rhizosolenia, Pseudosolenia) Arachnoidiscophytina D.G. Mann in Adl et al. 2019 (Arachnoidiscus) Bacillariophytina Medlin & Kaczmarska 2004, emend. Mediophyceae Jouse & Proshkina-Lavrenko in Medlin & Kaczmarska 2004 Chaetocerotophycidae Round & R.M. Crawford in Round et al. 1990, emend. Lithodesmiophycidae Round & R.M. Crawford in Round et al. 1990, emend. Thalassiosirophycidae Round & R.M. Crawford in Round et al. 1990 Cymatosirophycidae Round & R.M. Crawford in Round et al. 1990 Odontellophycidae D.G. Mann in Adl et al. 2019 Chrysanthemodiscophycidae D.G. Mann in Adl et al. 2019 Biddulphiophyceae D.G. Mann in Adl et al. 2019 Biddulphiophycidae Round and R.M. Crawford in Round et al. 1990, emend. Attheya Bacillariophyceae Haeckel 1878, emend. Striatellaceae Urneidophycidae Medlin 2016