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Cercozoa

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39-447: Cercozoa (now synonymised with Filosa ) is a phylum of diverse single-celled eukaryotes . They lack shared morphological characteristics at the microscopic level, and are instead united by molecular phylogenies of rRNA and actin or polyubiquitin . They were the first major eukaryotic group to be recognized mainly through molecular phylogenies . They are the natural predators of many species of bacteria. They are closely related to

78-405: A microtubular spindle during nuclear division, in the distinctively eukaryotic process of mitosis . Eukaryotes differ from prokaryotes in multiple ways, with unique biochemical pathways such as sterane synthesis. The eukaryotic signature proteins have no homology to proteins in other domains of life, but appear to be universal among eukaryotes. They include the proteins of the cytoskeleton,

117-518: A "symbiosis-based phylogeny", giving the description "Eukarya (symbiosis-derived nucleated organisms)". By 2014, a rough consensus started to emerge from the phylogenomic studies of the previous two decades. The majority of eukaryotes can be placed in one of two large clades dubbed Amorphea (similar in composition to the unikont hypothesis) and the Diphoda (formerly bikonts), which includes plants and most algal lineages. A third major grouping,

156-430: A body, with its cells dividing by mitosis , and at some stage produce haploid gametes through meiosis , a division that reduces the number of chromosomes and creates genetic variability . There is considerable variation in this pattern. Plants have both haploid and diploid multicellular phases . Eukaryotes have lower metabolic rates and longer generation times than prokaryotes, because they are larger and therefore have

195-479: A bundle of microtubules arising from a centriole , characteristically arranged as nine doublets surrounding two singlets. Flagella may have hairs ( mastigonemes ), as in many stramenopiles . Their interior is continuous with the cell's cytoplasm . Centrioles are often present, even in cells and groups that do not have flagella, but conifers and flowering plants have neither. They generally occur in groups that give rise to various microtubular roots. These form

234-569: A membrane-bound nucleus . All animals , plants , fungi , and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes : the Bacteria and the Archaea . Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass is much larger than that of prokaryotes. The eukaryotes seemingly emerged within

273-406: A primary component of the cytoskeleton, and are often assembled over the course of several cell divisions, with one flagellum retained from the parent and the other derived from it. Centrioles produce the spindle during nuclear division. The cells of plants, algae, fungi and most chromalveolates , but not animals, are surrounded by a cell wall. This is a layer outside the cell membrane , providing

312-568: A sister relationship between Filosa (=Cercozoa) and Endomyxa once again, although the modern classification of eukaryotes retains Endomyxa , Cercozoa and Retaria as separate phyla within Rhizaria . The phylum Cercozoa previously contained both Filosa and Endomyxa , but in the latest classifications Endomyxa has been excluded, and Cercozoa is now synonymous with Filosa. It is composed of two subphyla : Monadofilosa and Reticulofilosa . According to multigene phylogenetic analyses, Monadofilosa

351-510: A smaller surface area to volume ratio. The evolution of sexual reproduction may be a primordial characteristic of eukaryotes. Based on a phylogenetic analysis, Dacks and Roger have proposed that facultative sex was present in the group's common ancestor. A core set of genes that function in meiosis is present in both Trichomonas vaginalis and Giardia intestinalis , two organisms previously thought to be asexual. Since these two species are descendants of lineages that diverged early from

390-417: A system of domains rather than kingdoms as top level rank being put forward by Carl Woese , Otto Kandler , and Mark Wheelis in 1990, uniting all the eukaryote kingdoms in the domain "Eucarya", stating, however, that " 'eukaryotes' will continue to be an acceptable common synonym". In 1996, the evolutionary biologist Lynn Margulis proposed to replace kingdoms and domains with "inclusive" names to create

429-414: Is a robust clade , in which the deepest branching group is Metromonadea , followed by Helkesea as the second group (together forming the paraphyletic Eoglissa ) before the divergence of the clade Ventrifilosa ( Imbricatea , Sarcomonadea and Thecofilosea ). On the other hand, Reticulofilosa is probably paraphyletic, with Granofilosea diverging earlier than Chlorarachnea , which makes Chlorarachnea

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468-610: Is closer in structure to bacterial RNA than to eukaryote RNA. Some eukaryotes, such as the metamonads Giardia and Trichomonas , and the amoebozoan Pelomyxa , appear to lack mitochondria, but all contain mitochondrion-derived organelles, like hydrogenosomes or mitosomes , having lost their mitochondria secondarily. They obtain energy by enzymatic action in the cytoplasm. Plants and various groups of algae have plastids as well as mitochondria. Plastids, like mitochondria, have their own DNA and are developed from endosymbionts , in this case cyanobacteria . They usually take

507-516: Is strongly supported by phylogenetic studies. Some cercozoans are grouped by whether they are "filose" or "reticulose" in the behavior of their cytoskeleton when moving: Other important ecological groups are: As well as being highly diverse in morphology and physiology , Cercozoa also shows high ecological diversity . The phylum Cercozoa includes many of the most abundant and ecologically significant protozoa in soil , marine and freshwater ecosystems. Soil-dwelling cercozoans are one of

546-536: Is that their cells have nuclei . This gives them their name, from the Greek εὖ ( eu , "well" or "good") and κάρυον ( karyon , "nut" or "kernel", here meaning "nucleus"). Eukaryotic cells have a variety of internal membrane-bound structures, called organelles , and a cytoskeleton which defines the cell's organization and shape. The nucleus stores the cell's DNA , which is divided into linear bundles called chromosomes ; these are separated into two matching sets by

585-684: The Asgard archaea , and are closely related to the Heimdallarchaeia . This implies that there are only two domains of life , Bacteria and Archaea, with eukaryotes incorporated among the Archaea. Eukaryotes first emerged during the Paleoproterozoic , likely as flagellated cells. The leading evolutionary theory is they were created by symbiogenesis between an anaerobic Asgard archaean and an aerobic proteobacterium , which formed

624-1178: The Excavata , has been abandoned as a formal group as it is paraphyletic . The proposed phylogeny below includes only one group of excavates ( Discoba ), and incorporates the 2021 proposal that picozoans are close relatives of rhodophytes. The Provora are a group of microbial predators discovered in 2022. Ancyromonadida [REDACTED] Malawimonada [REDACTED] CRuMs [REDACTED] Amoebozoa [REDACTED] Breviatea [REDACTED] Apusomonadida [REDACTED] Holomycota (inc. fungi) [REDACTED] Holozoa (inc. animals) [REDACTED] ? Metamonada [REDACTED] Discoba [REDACTED] Cryptista [REDACTED] Rhodophyta (red algae) [REDACTED] Picozoa [REDACTED] Glaucophyta [REDACTED] Viridiplantae (plants) [REDACTED] Hemimastigophora [REDACTED] Provora [REDACTED] Haptista [REDACTED] Telonemia [REDACTED] Rhizaria [REDACTED] Alveolata [REDACTED] Stramenopiles [REDACTED] [REDACTED] Endomyxa Endomyxa

663-518: The archaea —having a volume of around 10,000 times greater. Eukaryotes represent a small minority of the number of organisms , but, as many of them are much larger, their collective global biomass (468 gigatons) is far larger than that of prokaryotes (77 gigatons), with plants alone accounting for over 81% of the total biomass of Earth . The eukaryotes are a diverse lineage, consisting mainly of microscopic organisms . Multicellularity in some form has evolved independently at least 25 times within

702-583: The blue whale , weighing up to 190 tonnes and measuring up to 33.6 metres (110 ft) long, or plants like the coast redwood , up to 120 metres (390 ft) tall. Many eukaryotes are unicellular; the informal grouping called protists includes many of these, with some multicellular forms like the giant kelp up to 200 feet (61 m) long. The multicellular eukaryotes include the animals, plants, and fungi , but again, these groups too contain many unicellular species . Eukaryotic cells are typically much larger than those of prokaryotes —the bacteria and

741-744: The mitochondria . A second episode of symbiogenesis with a cyanobacterium created the plants, with chloroplasts . Eukaryotic cells contain membrane-bound organelles such as the nucleus , the endoplasmic reticulum , and the Golgi apparatus . Eukaryotes may be either unicellular or multicellular . In comparison, prokaryotes are typically unicellular. Unicellular eukaryotes are sometimes called protists . Eukaryotes can reproduce both asexually through mitosis and sexually through meiosis and gamete fusion ( fertilization ). Eukaryotes are organisms that range from microscopic single cells , such as picozoans under 3 micrometres across, to animals like

780-419: The ocean floor . Some cercozoa are coprophilic or coprozoic , meaning they use feces as a source of nutrients or as transport through animal hosts. The faecal habitat is an understudied reservoir of microbial eukaryotic diversity, dominated by amoeboflagellates from the phylum Cercozoa. Strongly coprophilic examples of cercozoa are the flagellates Cercomonas , Proleptomonas and Helkesimastix , and

819-480: The sorocarpic amoeba Guttulinopsis . Many new cercozoan lineages, especially among sarcomonads , have been discovered through phylogenetic sampling of feces because they appear preferentially in this medium. Cercozoan bacterivores (i.e. predators of bacteria ) are highly diverse and important in the plant phyllosphere , the leaf surfaces of plants. Particularly sarcomonads , with their ability to cyst , feed and multiply within hours, are perfectly adapted to

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858-514: The taxonomic rank of kingdom by Linnaeus in the 18th century. Though he included the fungi with plants with some reservations, it was later realized that they are quite distinct and warrant a separate kingdom. The various single-cell eukaryotes were originally placed with plants or animals when they became known. In 1818, the German biologist Georg A. Goldfuss coined the word Protozoa to refer to organisms such as ciliates , and this group

897-709: The cell to move, change shape, or transport materials. The motor structures are microfilaments of actin and actin-binding proteins , including α- actinin , fimbrin , and filamin are present in submembranous cortical layers and bundles. Motor proteins of microtubules, dynein and kinesin , and myosin of actin filaments, provide dynamic character of the network. Many eukaryotes have long slender motile cytoplasmic projections, called flagella , or multiple shorter structures called cilia . These organelles are variously involved in movement, feeding, and sensation. They are composed mainly of tubulin , and are entirely distinct from prokaryotic flagella. They are supported by

936-404: The cell with structural support, protection, and a filtering mechanism. The cell wall also prevents over-expansion when water enters the cell. The major polysaccharides making up the primary cell wall of land plants are cellulose , hemicellulose , and pectin . The cellulose microfibrils are linked together with hemicellulose, embedded in a pectin matrix. The most common hemicellulose in

975-427: The complex transcription machinery, the membrane-sorting systems, the nuclear pore , and some enzymes in the biochemical pathways. Eukaryote cells include a variety of membrane-bound structures, together forming the endomembrane system. Simple compartments, called vesicles and vacuoles , can form by budding off other membranes. Many cells ingest food and other materials through a process of endocytosis , where

1014-622: The cytoplasm. Mitochondria are organelles in eukaryotic cells. The mitochondrion is commonly called "the powerhouse of the cell", for its function providing energy by oxidising sugars or fats to produce the energy-storing molecule ATP . Mitochondria have two surrounding membranes , each a phospholipid bilayer , the inner of which is folded into invaginations called cristae where aerobic respiration takes place. Mitochondria contain their own DNA , which has close structural similarities to bacterial DNA , from which it originated, and which encodes rRNA and tRNA genes that produce RNA which

1053-399: The dominant groups of free-living eukaryotic microorganisms found in temperate soils, accounting for around 30% of identifiable protozoan DNA in arid or semi-arid soils and 15% in more humid soils. In transcriptomic analyses they account for 40-60% of all identifiable protozoan RNA found in forest and grassland soils. They also comprise 9-24% of all operational taxonomic units found in

1092-406: The eukaryotes. Complex multicellular organisms, not counting the aggregation of amoebae to form slime molds , have evolved within only six eukaryotic lineages: animals , symbiomycotan fungi , brown algae , red algae , green algae , and land plants . Eukaryotes are grouped by genomic similarities, so that groups often lack visible shared characteristics. The defining feature of eukaryotes

1131-484: The eukaryotic evolutionary tree, core meiotic genes, and hence sex, were likely present in the common ancestor of eukaryotes. Species once thought to be asexual, such as Leishmania parasites, have a sexual cycle. Amoebae, previously regarded as asexual, may be anciently sexual; while present-day asexual groups could have arisen recently. In antiquity , the two lineages of animals and plants were recognized by Aristotle and Theophrastus . The lineages were given

1170-696: The fluctuating environmental factors in the phyllosphere. Their predation causes shifts in the bacterial communities: they reduce populations of alphaproteobacteria and betaproteobacteria , which are less resistant to their grazing, in favour of other bacterial populations such as gammaproteobacteria . Reticulofilosa Monadofilosa Endomyxa Taxopodida Foraminifera Radiolaria Reticulofilosa Monadofilosa Endomyxa Lapot gusevi Foraminifera Polycystinea Acantharea Originally, Cercozoa contained both Filosa and Endomyxa , according to phylogenetic analyses using ribosomal RNA and tubulin . These analyses also confirmed Cercozoa as

1209-646: The form of chloroplasts which, like cyanobacteria, contain chlorophyll and produce organic compounds (such as glucose ) through photosynthesis . Others are involved in storing food. Although plastids probably had a single origin, not all plastid-containing groups are closely related. Instead, some eukaryotes have obtained them from others through secondary endosymbiosis or ingestion. The capture and sequestering of photosynthetic cells and chloroplasts, kleptoplasty , occurs in many types of modern eukaryotic organisms. The cytoskeleton provides stiffening structure and points of attachment for motor structures that enable

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1248-449: The outer membrane invaginates and then pinches off to form a vesicle. Some cell products can leave in a vesicle through exocytosis . The nucleus is surrounded by a double membrane known as the nuclear envelope , with nuclear pores that allow material to move in and out. Various tube- and sheet-like extensions of the nuclear membrane form the endoplasmic reticulum , which is involved in protein transport and maturation. It includes

1287-482: The phylum Retaria , comprising amoeboids that usually have complex shells, and together form a supergroup called Rhizaria . The group includes most amoeboids and flagellates that feed by means of filose pseudopods. These may be restricted to part of the cell surface, but there is never a true cytostome or mouth as found in many other protozoa. They show a variety of forms and have proven difficult to define in terms of structural characteristics, although their unity

1326-417: The primary cell wall is xyloglucan . Eukaryotes have a life cycle that involves sexual reproduction , alternating between a haploid phase, where only one copy of each chromosome is present in each cell, and a diploid phase, with two copies of each chromosome in each cell. The diploid phase is formed by fusion of two haploid gametes, such as eggs and spermatozoa , to form a zygote ; this may grow into

1365-498: The rough endoplasmic reticulum, covered in ribosomes which synthesize proteins; these enter the interior space or lumen. Subsequently, they generally enter vesicles, which bud off from the smooth endoplasmic reticulum. In most eukaryotes, these protein-carrying vesicles are released and further modified in stacks of flattened vesicles ( cisternae ), the Golgi apparatus . Vesicles may be specialized; for instance, lysosomes contain digestive enzymes that break down biomolecules in

1404-607: The sister group of Monadofilosa. Imbricatea Sarcomonadea Thecofilosea Helkesea Metromonadea Chlorarachnea Granofilosea A more recent phylogenomic analysis recovered both Monadofilosa and Reticulofilosa as monophyletic within the clade Filosa. In addition to the known Granofilosea , Chlorarachnea and Monadofilosa , a variety of clades inside Cercozoa have been discovered in other analyses and have slowly been described and named, such as Tremulida (previously known as Novel Clade 11) and Aquavolonida (Novel Clade 10), although their specific positions among

1443-431: The sister group of Retaria within the supergroup Rhizaria. However, the monophyly of the group was still uncertain. Posterior multigene phylogenetic analyses consistently found Cercozoa to be paraphyletic, because Endomyxa clustered next to Retaria instead of Filosa. Because of this, Endomyxa was excluded from Cercozoa, which became a synonym of Filosa. More recent phylogenomic analyses with better sampling recovered

1482-481: The two main cercozoan subphyla have yet to be refined. These two orders have been classified as the class Skiomonadea , within Reticulofilosa. The classification of Cercozoa was revised in 2018: Eukaryote The eukaryotes ( / j uː ˈ k ær i oʊ t s , - ə t s / yoo- KARR -ee-ohts, -⁠əts ) constitute the domain of Eukaryota or Eukarya , organisms whose cells have

1521-503: Was expanded until Ernst Haeckel made it a kingdom encompassing all single-celled eukaryotes, the Protista , in 1866. The eukaryotes thus came to be seen as four kingdoms: The protists were at that time thought to be "primitive forms", and thus an evolutionary grade , united by their primitive unicellular nature. Understanding of the oldest branchings in the tree of life only developed substantially with DNA sequencing , leading to

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