Ochrophytes Temporal range: Middle Proterozoic [1] | |
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Dense kelp forest with understory at Partridge Point near Dave's Caves, Cape Peninsula | |
Scientific classification | |
Domain: | Eukaryota |
Clade: | Diaphoretickes |
Clade: | SAR |
Clade: | Stramenopiles |
Phylum: | Gyrista |
Subphylum: | Ochrophytina Cavalier-Smith 1986 emend. 1996 [2] |
Type genus | |
Fucus Linnaeus, 1753 | |
Classes [3] | |
Diversity | |
23,314 described species [5] >100,000 estimated species [6] | |
Synonyms | |
Ochrophytes, also known as heterokontophytes or stramenochromes, are a group of algae. They are the photosynthetic stramenopiles, a group of eukaryotes, organisms with a cell nucleus, characterized by the presence of two unequal flagella, one of which has tripartite hairs called mastigonemes. In particular, they are characterized by photosynthetic organelles or plastids enclosed by four membranes, with membrane-bound compartments called thylakoids organized in piles of three, chlorophyll a and c as their photosynthetic pigments, and additional pigments such as β-carotene and xanthophylls. Ochrophytes are one of the most diverse lineages of eukaryotes, containing ecologically important algae such as brown algae and diatoms. They are classified either as phylum Ochrophyta or Heterokontophyta, or as subphylum Ochrophytina within phylum Gyrista. Their plastids are of red algal origin.
Ochrophytes are eukaryotic organisms composed of cells that are either naked or covered by scales, lorica or a cell wall. They can be single-celled, colonial, coenocytic or multicellular. Some Phaeophyceae develop as large multicellular thalli with differentiated tissues. [7] All ochrophytes uniformly have tubular mitochondrial cristae. [10] This is a common trait shared with their relatives, heterotrophic stramenopiles, as well as other closely related groups such as Rhizaria, Telonemia and Alveolata. [11] [12] As primarily photosynthetic eukaryotes, they are considered algae, distinguished from other groups of algae by specific morphological and ultrastructural traits, such as their flagella, chloroplasts and pigments. [10]
As stramenopiles (=heterokonts), their swimming cells frequently display two markedly unequal flagella: an anterior flagellum ("tinsel") with straw-like hollow tripartite hairs called mastigonemes, and an immature posterior smooth flagellum ("whiplash") lacking these hairs. [13] [10] The ciliary transition zone of the flagellum generally has a transitional helix. [7]
The ochrophytes are mostly photosynthetic. As such, they may possess one or more photosynthetic plastids (chloroplasts) per cell. [14] Some groups contain species with leucoplasts, chloroplasts that have lost photosynthetic capacity and pigments but presumably continue to play a role in the synthesis of amino acids, lipids and heme groups. [10] Ochrophytes have a distinct plastid ultrastructure in comparison to other algal groups. [14] Their chloroplasts originate from an event of secondary endosymbiosis from a red alga, which lead to four [lower-alpha 1] surrounding membranes: two inner membranes, corresponding to the primary plastid membranes; a third membrane, corresponding to the plasma membrane of the red alga; and an outermost layer, corresponding to the phagosome membrane. [17] This characteristic differentiates them from archaeplastid algae (glaucophytes, red algae and green algae), whose chloroplasts have only two membranes. [18] The two outer layers of ochrophyte plastids are continguous with the endoplasmic reticulum (ER), together composing the chloroplast endoplasmic reticulum (CER), [14] also known as the periplastidial endoplasmic reticulum (PER), which is often connected to the nuclear envelope. The tripartite flagellar hairs, characteristic of stramenopiles, are produced within either the PER or the nuclear envelope. [10]
The periplastid compartment (PC), between the second and third layers, is a separate region that in other algal groups (i.e. cryptomonads and chlorarachniophytes) contains a nucleomorph, the vestigial nucleus of the secondary endosymbiont; however, no nucleomorphs are known within the ochrophytes. Instead, other structures have been observed within the PC, similarly to those seen in haptophytes and chromerid algae: [14] "blob-like structures" where PC proteins are localized, and a vesicular network. [17] Within the CER, there is a prominent region of tight direct contacts between the periplastid membrane and the inner nuclear envelope, where lipid transfers might occur, and perhaps exchange of other molecules. [17]
Commonly, within the plastid stroma, three stacked thylakoids differentiate into the "girdle lamella", which runs around the periphery of the plastid, beneath the innermost membrane. [14] The remaining thylakoids are arranged in stacks of three. [10] In synchromophytes and aurearenophytes, a consortium of several plastids, each surrounded by two or three inner membranes respectively, is enveloped by a shared outer membrane. [14]
Ochrophyte chloroplasts contain chlorophylls a and c as photosynthetic pigments, in addition to fucoxanthin. [13] Chlorophyll a binds to thylakoids, while the c pigment is present in the stroma. [10] The most frequent accessory pigment in ochrophytes is the yellow β-carotene. The golden-brown or brown pigmentation in diatoms, brown algae, golden algae and others is conferred by the xanthophyll fucoxanthin. In the yellow-green or yellow-brown raphidophyceans, eustigmatophyceans and xanthophyceans, vaucheriaxanthin is dominant instead. These pigment combinations extend their photosynthetic ability beyond chlorophyll a alone. Additionally, xanthophylls protect the photosystems from high intensity light. [10]
Ochrophyte algae accumulate chrysolaminarin, a carbohydrate consisting of short chains of β-1,3-linked glucose molecules, as a storage product. [10] [19] It is stored in vesicles located within the cytoplasm, outside plastids, unlike other algae. [13] Cytoplasmic lipid droplets are also common. [10] They lack starch, which is the common storage product in green algae and plants. [7]
Ochrophytes are capable of asexual reproduction by fragmentation, propagules, vegetative cell division, sporogenesis or zoosporogenesis. In addition, they are capable of sexual reproduction through gametes, by three different modes: isogamy, anisogamy or oogamy. [7]
Ochrophytes are present in nearly all environments. [19] Some classes are more common in marine habitats, while others are more frequent in freshwater or soil. [10] Among the ochrophyte lineages are the diatoms, the most abundant photosynthetic eukaryotes worldwide in marine habitats; multicellular seaweeds, such as brown algae (e.g., kelp) and golden algae; and an array of microscopic single-celled lineages that are also abundant, as evidenced by environmental sequencing. [14] Regarding nutrition, various ochrophytes are mixotrophic, usually through phagocytosis. [19]
Several classes of heterokont algae are exclusively known from marine habitats, such as Bolidophyceae, Pelagophyceae, Pinguiophyceae and Schizocladiophyceae. The brown algae (Phaeophyceae) are almost exclusively marine, with very few freshwater genera. [19]
Chrysophyceae, Phaeothamniophyceae and Xanthophyceae are predominantly freshwater classes. In lotic habitats (rivers, streams), golden algae (Chrysophyceae) and yellow-green algae (Xanthophyceae) are common and occasionally abundant. The golden algal genus Hydrurus , in particular, can be widespread in some drainage basins and is common in cold, clear, fast-flowing mountain streams, where it attaches to a firm substrate. Xanthophycean genera commonly found in rivers include Vaucheria , Tribonema and Bumilleria , either freely floating or attached to filamentous algae and plants. [20] Diatoms are more diverse, with more than 60 genera commonly found in rivers. Many river diatoms have developed different strategies to attach to the substrate to avoid being displaced by water currents. The most basic strategy is to produce extracellular polymeric substances, varied carbohydrate structures formed from the cell membrane. In faster-flowing waters, some diatoms (e.g., Cocconeis ) grow directly attached to the substrate through adhesive films. Others (e.g., Eunotia , Nitzschia ) grow stalks or colonial tubes capable of reaching higher into the water column to acquire more nutrients. [21] Brown algae (Phaeophyceae), although highly diversified, contain only seven species present in rivers. These lack any complex multicellular thalli, and instead exist as benthic filamentous forms that have evolved independently from marine ancestors. [22]
Two main lineages of photosynthetic stramenopiles include many toxic species. Within the class Raphidophyceae, strains of Heterosigma and Chattonella at high concentrations are responsible for fish mortality, although the nature and action of their toxins is not resolved. Freshwater Gonyostomum species are capable of mucilage secretion at high amounts detrimental to fish gills. Within the diatoms (Bacillariophyta), harmful effects can be due to physical damage or to toxin production. Centric diatoms like Chaetoceros live as colonial chains of cells with long spines (setae) that can clog fish gills, causing their death. Among diatoms, the only toxin producers have been found among pennate diatoms, almost entirely within the genus Pseudonitzschia . More than a dozen species of Pseudonitzschia are capable of producing a neurotoxin, domoic acid, the cause of amnesiac shellfish poisoning. [23]
The ochrophytes constitute a highly diverse clade within Stramenopila, a eukaryotic supergroup that also includes several heterotrophic lineages of protists such as oomycetes, hyphochytrids, labyrinthuleans, opalines and bicosoecids. [24] [3] [25] This lineage of stramenopiles originated from an event of secondary endosymbiosis where a red alga was transformed into the chloroplast of the common ancestor of ochrophytes. [3] [26] [27]
The total group of ochrophytes is estimated to have evolved between 874 and 543 million years ago (Ma) through molecular clock inference. However, the earliest fossil remains, assigned to the billion-year-old xanthophyte Palaeovaucheria , [1] suggest that ochrophytes had appeared by 1000 Ma. Other early putative representatives of photosynthetic stramenopiles are Jacutianema (750 Ma), Germinosphaera (750–700 Ma) and the brown alga Miaohephyton (600–550 Ma). Scales similar to modern chrysophyte scales, and valves resembling the modern centric diatom valves, have been found in 800–700 million-years-old sediments. [28]
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Evolutionary relationships between all ochrophyte classes based on the latest phylogenetic analyses, [30] [27] [29] [4] and the approximate number of species in each class. [3] |
Relationships among many classes of ochrophytes remain unresolved, but three main clades (called SI, SII and SIII) are supported in most phylogenetic analyses. The SI lineage, containing the diverse and multicellular class Phaeophyceae, or brown algae, experienced an evolutionary radiation during the late Paleozoic (around 310 million years ago). The class Schizocladiophyceae is the sister lineage to brown algae, followed by a clade of closely related classes Xanthophyceae, Phaeosacciophyceae [29] and Chrysoparadoxophyceae. [15] This is in turn the sister lineage to a clade containing Aurearenophyceae and Phaeothamniophyceae, [3] which are sometimes treated as one class Aurophyceae. [27] The Raphidophyceae are the most basal within the SI. The SII lineage contains the golden algae or Chrysophyceae, as well as smaller classes Synurophyceae, Eustigmatophyceae, Pinguiophyceae and Picophagea (also known as Synchromophyceae). Both clades, SI and SII, compose the Chrysista lineage. The remaining classes are grouped within the sister lineage Diatomista, equivalent to the SIII lineage; these are the diatoms or Bacillariophyceae, Bolidophyceae, Dictyochophyceae (including the silicoflagellates) and Pelagophyceae. [3] A new class of algae, Olisthodiscophyceae, was described in 2021 and recovered as part of the SII lineage. [4]
One group of heterotrophic heliozoan protists, Actinophryida, [31] is included in some classifications as the sister lineage to the raphidophytes, and both groups are treated as one class Raphidomonadea on the basis of 18S rDNA phylogenetic analyses. [32] However, a recent phylogenomic study places one actinophryid, Actinophrys sol , as the probable sister group to ochrophytes. Although it lacks chloroplasts, plastidial genes have been found in the nuclear genome of this actinophryid, implying that its common ancestor with ochrophytes may have already begun domesticating plastids. [33]
In hierarchical classifications, where taxonomic ranks (kingdom, phylum, class, order...) are utilized, the heterokont algae are commonly regarded as an entire phylum (or division in botanical nomenclature) by the name of Ochrophyta, within the Stramenopila or Heterokonta. [24] The phylum was first described by protozoologist Thomas Cavalier-Smith in 1986, as Ochrista, later renamed to Ochrophyta in 1996 in accordance to recommendations of the International Code of Nomenclature for algae, fungi, and plants (ICN). [2] [34] It remained a phylum-level taxon until 2017, when the same author lowered it to subphylum level and modified the name to Ochrophytina to match the -phytina suffix in botanical nomenclature, which corresponds to subdivisions. The phylum to which ochrophytes belong in his classification system is Gyrista, a clade that also contains some heterotrophic heterokonts, namely the Pseudofungi and the Bigyromonada. [27] Gyrista and Bigyra compose the two main branches of stramenopiles, which are regarded as the superphylum Heterokonta within the kingdom Chromista. However, this classification system is in disuse due to the kingdom's non-monophyletic nature. [25]
While Ochrophyta is the preferred name by general protistologists and protozoologists, the name Heterokontophyta is considered more familiar among phycologists. [7] The origin of this name is the class Heterokontæ , introduced by Finnish biologist Alexander Ferdinand Luther in 1899 [35] to include the orders Chloromonadales and Confervales, later separated into Xanthophyceae and Raphidophyceae. This name referenced, among other traits, the two unequal flagella characteristic of all Stramenopiles, also known as heterokonts. After several electron microscopy discoveries, Christiaan van den Hoek introduced in 1978 the division Heterokontophyta for five algal classes: Chrysophyceae, Xanthophyceae, Bacillariophyceae, Phaeophyceae, and Chloromonadophyceae. [36] Several other names were used to group heterokont algae with ambiregnal organisms, such as Chromophyta, Heterokonta and lastly Stramenopiles, which is not a validly published name under the ICN. [7] Several phycologists currently advocate the use of Heterokontophyta as the phylum name for heterokont algae. However, the original use by Hoek in 1978 did not provide a Latin description, which was a requirement for valid publication under the ICN until 2011. Phycologists Michael Guiry, Øjvind Moestrup and Robert Andersen validly published Heterokontophyta as a phylum in 2023. [7]
As opposed to the hierarchical classification, the cladistic classification only recognizes clades as valid groups, rejecting the use of paraphyletic or polyphyletic groups. This method of classification is preferred among protistologists. The latest revision of the International Society of Protistologists, in 2019, recognizes Ochrophyta as a valid taxon within the higher Stramenopiles group, within the SAR supergroup. [25] The subdivision of ochrophytes between Chrysista and Diatomista is fully accepted by the scientific community and backed up by phylogenetic analyses. [25]
As of 2024, ochrophytes amount to 23,314 described species, with 490 species of uncertain position. [5] However, it is estimated that they amount to more than 100,000 species, of which the majority are diatoms. [6] Below is the present classification of ochrophytes according to the 2019 revision of eukaryotic classification, [25] with the inclusion of classes of algae described in posterior years [15] [29] [4] as well as the number of described species for each class. [5] According to the aforementioned 2019 revision by protistologists, the diatoms (Diatomeae) do not form a single class Bacillariophyceae, but numerous classes to reflect the phylogenetic advances over the previous decade. [25]
Multicellular seaweeds, in the class Phaeophyceae, were described in early Chinese (around 3000 BC), Greek (300 BC, such as Theophrastus) and Japanese (ca. 500 AD) writings. Knowledge of them likely predates recorded history, being used as food, dyes, and for medicinal purposes. The first formal description of a stramenopile alga was that of the genus Fucus , by Linnaeus in his 1753 work Species Plantarum . Shortly after, unicellular chrysophytes were described by Otto Friedrich Müller. During this first era of scientific discovery, brown algae were described as plants, while microscopic algae were treated as animals under the name of infusoria. [19]
During the 20th century, evolutionary and phylogenetic discussions began including heterokont algae. Transmission electron microscopy and molecular phylogenetic analysis led to the description of many new groups and several classes well into the 21st century. The sequencing of the first ochrophyte genome, belonging to Thalassiosira pseudonana , began in 2002. [19]
A chloroplast is a type of organelle known as a plastid that conducts photosynthesis mostly in plant and algal cells. Chloroplasts have a high concentration of chlorophyll pigments which capture the energy from sunlight and convert it to chemical energy and release oxygen. The chemical energy created is then used to make sugar and other organic molecules from carbon dioxide in a process called the Calvin cycle. Chloroplasts carry out a number of other functions, including fatty acid synthesis, amino acid synthesis, and the immune response in plants. The number of chloroplasts per cell varies from one, in some unicellular algae, up to 100 in plants like Arabidopsis and wheat.
The Stramenopiles, also called Heterokonts, are a clade of organisms distinguished by the presence of stiff tripartite external hairs. In most species, the hairs are attached to flagella, in some they are attached to other areas of the cellular surface, and in some they have been secondarily lost. Stramenopiles represent one of the three major clades in the SAR supergroup, along with Alveolata and Rhizaria.
The alveolates are a group of protists, considered a major clade and superphylum within Eukarya. They are currently grouped with the stramenopiles and Rhizaria among the protists with tubulocristate mitochondria into the SAR supergroup.
The silicoflagellates are a small group of unicellular photosynthetic protists, or algae, belonging to the supergroup of eukaryotes known as Stramenopiles. They behave as plankton and are present in oceanic waters. They are well-known from harmful algal blooms that cause high mortality of fish. Additionally, they compose a rich fossil record represented by their silica skeletons.
Brown algae are a large group of multicellular algae comprising the class Phaeophyceae. They include many seaweeds located in colder waters of the Northern Hemisphere. Brown algae are the major seaweeds of the temperate and polar regions. Many brown algae, such as members of the order Fucales, commonly grow along rocky seashores. Most brown algae live in marine environments, where they play an important role both as food and as a potential habitat. For instance, Macrocystis, a kelp of the order Laminariales, may reach 60 m (200 ft) in length and forms prominent underwater kelp forests that contain a high level of biodiversity. Another example is Sargassum, which creates unique floating mats of seaweed in the tropical waters of the Sargasso Sea that serve as the habitats for many species. Some members of the class, such as kelps, are used by humans as food.
Chromista is a proposed but polyphyletic biological kingdom, refined from the Chromalveolata, consisting of single-celled and multicellular eukaryotic species that share similar features in their photosynthetic organelles (plastids). It includes all eukaryotes whose plastids contain chlorophyll c and are surrounded by four membranes. If the ancestor already possessed chloroplasts derived by endosymbiosis from red algae, all non-photosynthetic Chromista have secondarily lost the ability to photosynthesise. Its members might have arisen independently as separate evolutionary groups from the last eukaryotic common ancestor.
Labyrinthulomycetes (ICBN) or Labyrinthulea (ICZN) is a class of protists that produce a network of filaments or tubes, which serve as tracks for the cells to glide along and absorb nutrients for them. The two main groups are the labyrinthulids and thraustochytrids. They are mostly marine, commonly found as parasites on algae and seagrasses or as decomposers on dead plant material. They also include some parasites of marine invertebrates and mixotrophic species that live in a symbiotic relationship with zoochlorella.
Yellow-green algae or the Xanthophyceae (xanthophytes) are an important group of heterokont algae. Most live in fresh water, but some are found in marine and soil habitats. They vary from single-celled flagellates to simple colonial and filamentous forms. Xanthophyte chloroplasts contain the photosynthetic pigments chlorophyll a, chlorophyll c, β-carotene, and the carotenoid diadinoxanthin. Unlike other Stramenopiles (heterokonts), their chloroplasts do not contain fucoxanthin, which accounts for their lighter colour. Their storage polysaccharide is chrysolaminarin. Xanthophyte cell walls are produced of cellulose and hemicellulose. They appear to be the closest relatives of the brown algae.
Euglenophyceae (ICBN) or Euglenea (ICZN) is a group of single-celled algae belonging to the phylum Euglenozoa. They have chloroplasts originated from an event of secondary endosymbiosis with a green alga. They are distinguished from other algae by the presence of paramylon as a storage product and three membranes surrounding each chloroplast.
Chromalveolata was a eukaryote supergroup present in a major classification of 2005, then regarded as one of the six major groups within the eukaryotes. It was a refinement of the kingdom Chromista, first proposed by Thomas Cavalier-Smith in 1981. Chromalveolata was proposed to represent the organisms descended from a single secondary endosymbiosis involving a red alga and a bikont. The plastids in these organisms are those that contain chlorophyll c.
The Archaeplastida are a major group of eukaryotes, comprising the photoautotrophic red algae (Rhodophyta), green algae, land plants, and the minor group glaucophytes. It also includes the non-photosynthetic lineage Rhodelphidia, a predatorial (eukaryotrophic) flagellate that is sister to the Rhodophyta, and probably the microscopic picozoans. The Archaeplastida have chloroplasts that are surrounded by two membranes, suggesting that they were acquired directly through a single endosymbiosis event by phagocytosis of a cyanobacterium. All other groups which have chloroplasts, besides the amoeboid genus Paulinella, have chloroplasts surrounded by three or four membranes, suggesting they were acquired secondarily from red or green algae. Unlike red and green algae, glaucophytes have never been involved in secondary endosymbiosis events.
In botany, a zoid or zoïd is a reproductive cell that possesses one or more flagella, and is capable of independent movement. Zoid can refer to either an asexually reproductive spore or a sexually reproductive gamete. In sexually reproductive gametes, zoids can be either male or female depending on the species. For example, some brown alga (Phaeophyceae) reproduce by producing multi-flagellated male and female gametes that recombine to form the diploid sporangia. Zoids are primarily found in some protists, diatoms, green alga, brown alga, non-vascular plants, and a few vascular plants. The most common classification group that produces zoids is the heterokonts or stramenopiles. These include green alga, brown alga, oomycetes, and some protists. The term is generally not used to describe motile, flagellated sperm found in animals. Zoid is also commonly confused for zooid which is a single organism that is part of a colonial animal.
SAR or Harosa is a highly diverse clade of eukaryotes, often considered a supergroup, that includes stramenopiles (heterokonts), alveolates, and rhizarians. It is a node-based taxon, including all descendants of the three groups' last common ancestor, and comprises most of the now-rejected Chromalveolata. Their sister group has been found to be telonemids, with which they make up the TSAR clade.
Bigyra is a phylum of microscopic eukaryotes that are found at the base of the Stramenopiles clade. It includes three well-known heterotrophic groups Bicosoecida, Opalinata and Labyrinthulomycetes, as well as several small clades initially discovered through environmental DNA samples: Nanomonadea, Placididea, Opalomonadea and Eogyrea. The classification of Bigyra has changed several times since its origin, and its monophyly remains unresolved.
Guillardia is a genus of marine biflagellate cryptomonad algae with a plastid obtained through secondary endosymbiosis of a red alga.
Vaucheria litorea is a species of yellow-green algae (Xanthophyceae). It grows in a filamentous fashion. V. litorea is a common intertidal species of coastal brackish waters and salt marshes of the Northern Atlantic, along the coasts of Europe, North America and New Zealand. It is also found in the Eastern Pacific coasts of Washington state. It is found to be able to tolerate a large range of salinities, making it euryhaline.
Picophagea, also known as Synchromophyceae, is a class of photosynthetic stramenopiles. The chloroplast of the Synchromophyceae are surrounded by two membranes and arranged in a way where they share the outer pair of membranes. The entire chloroplast complex is surrounded by an additional two outer membranes.
Gyrista is a phylum of heterokont protists containing three diverse groups: the mostly photosynthetic Ochrophyta, the parasitic Pseudofungi, and the recently described group of nanoflagellates known as Bigyromonada. Members of this phylum are characterized by the presence of a helix or a double helix/ring system in the ciliary transition region.
Synchroma is a genus of marine stramenopile algae containing two species with amoeboid morphology. They are grouped within the monotypic family Synchromaceae and order Synchromales as part of an independent clade of ochrophytes known as Picophagea or Synchromophyceae.
Ochromonadales is an order of single-celled algae belonging to the class Chrysophyceae, also known as golden algae. Initially it contained numerous groups of flagellates that were not closely related. During the late 20th century, advancements in molecular and ultrastructural studies allowed the transfer of many of these groups out of Ochromonadales, and the order was reduced to a single family Ochromonadaceae. They are aquatic single-celled flagellated algae, with two heterokont flagella each, some of which have secondarily lost their chloroplasts and appear colorless.