71-451: A protist ( / ˈ p r oʊ t ɪ s t / PROH -tist ) or protoctist is any eukaryotic organism that is not an animal , land plant , or fungus . Protists do not form a natural group, or clade , but are a polyphyletic grouping of several independent clades that evolved from the last eukaryotic common ancestor . Protists were historically regarded as a separate taxonomic kingdom known as Protista or Protoctista . With
142-405: 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,
213-495: A model organism ). Free-living ciliates are usually the top heterotrophs and predators in microbial food webs, feeding on bacteria and smaller eukaryotes, present in a variety of ecosystems, although a few species are kleptoplastic . Others are parasitic of numerous animals. Ciliates have a basal position in the evolution of alveolates, together with a few species of heterotrophic flagellates with two cilia collectively known as colponemids . The remaining alveolates are grouped under
284-518: 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,
355-417: A big portion of the oxygen produced worldwide, and comprising much of the marine phytoplankton ; the brown algae , filamentous or 'truly' multicellular (with differentiated tissues) macroalgae that constitute the basis of many temperate and cold marine ecosystems, such as kelp forests ; and the golden algae , unicellular or colonial flagellates that are mostly present in freshwater habitats. Inside Gyrista,
426-430: 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
497-479: 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
568-475: A collection of amoebae, flagellates and amoeboflagellates with complex life cycles, among which are some slime molds ( acrasids ). The two clades Euglenozoa and Percolozoa are sister taxa, united under the name Discicristata , in reference to their mitochondrial cristae shaped like discs. The species Tsukubamonas globosa is a free-living flagellate whose precise position within Discoba is not yet settled, but
639-544: A diverse group (>1,000 living species) of amoebae, often bearing delicate and intricate siliceous skeletons. The forams (Foraminifera) are also diverse (>6,700 living species), and most of them are encased in multichambered tests constructed from calcium carbonate or agglutinated mineral particles. Both groups have a rich fossil record, with tens of thousands of described fossil species. Eukaryotic The eukaryotes ( / j uː ˈ k ær i oʊ t s , - ə t s / yoo- KARR -ee-ohts, -əts ) constitute
710-460: A large (>6,000 species) and highly specialized group of obligate parasites who have all secondarily lost their photosynthetic ability (e.g., Plasmodium falciparum , cause of malaria ). Their adult stages absorb nutrients from the host through the cell membrane, and they reproduce between hosts via sporozoites, which exhibit an organelle complex (the apicoplast ) evolved from non-photosynthetic chloroplasts. The other branch of Myzozoa contains
781-596: A major cause of harmful algal blooms due to their toxicity; some live as symbionts of corals, allowing the creation of coral reefs. Dinoflagellates exhibit a diversity of cellular structures, such as complex eyelike ocelli, specialized vacuoles, bioluminescent organelles, and a wall surrounding the cell known as the theca . Rhizaria is a lineage of morphologically diverse organisms, composed almost entirely of unicellular heterotrophic amoebae, flagellates and amoeboflagellates, commonly with reticulose (net-like) or filose (thread-like) pseudopodia for feeding and locomotion. It
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#1732772284626852-406: 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
923-471: A small group (3 species) of freshwater or marine suspension-feeding bacterivorous flagellates with typical excavate appearance, closely resembling Jakobida and some metamonads but not phylogenetically close to either in most analyses. Diaphoretickes includes nearly all photosynthetic eukaryotes. Within this clade, the TSAR supergroup gathers a colossal diversity of protists. The most basal branching member of
994-510: 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
1065-417: 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
1136-473: A variety of algae. In addition, two smaller groups, Haptista and Cryptista , also belong to Diaphoretickes. The Stramenopiles, also known as Heterokonta, are characterized by the presence of two cilia, one of which bears many short, straw-like hairs ( mastigonemes ). They include one clade of phototrophs and numerous clades of heterotrophs, present in virtually all habitats. Stramenopiles include two usually well-supported clades, Bigyra and Gyrista , although
1207-416: A wide variety of animals – which act as secondary or intermediate host – but can undergo sexual reproduction only in the primary or definitive host (for example: felids such as domestic cats in this case). Some species, for example Plasmodium falciparum , have extremely complex life cycles that involve multiple forms of the organism, some of which reproduce sexually and others asexually. However, it
1278-567: Is paraphyletic , with some analyses placing the root of the eukaryote tree within Metamonada. Discoba includes three major groups: Jakobida , Euglenozoa and Percolozoa . Jakobida are a small group (~20 species) of free-living heterotrophic flagellates, with two cilia, that primarily eat bacteria through suspension feeding; most are aquatic aerobes, with some anaerobic species, found in marine, brackish or fresh water. They are best known for their bacterial-like mitochondrial genomes. Euglenozoa
1349-436: Is a rich (>2,000 species) group of flagellates with very different lifestyles, including: the free-living heterotrophic (both osmo- and phagotrophic) and photosynthetic euglenids (e.g., the euglenophytes , with chloroplasts originated from green algae); the free-living and parasitic kinetoplastids (such as the trypanosomes ); the deep-sea anaerobic symbiontids ; and the elusive diplonemids . Percolozoa (~150 species) are
1420-610: 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
1491-643: Is much larger than that of prokaryotes. The eukaryotes seemingly emerged within 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
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#17327722846261562-418: Is probably more closely related to Discicristata than to Jakobida. The metamonads (Metamonada) are a phylum of completely anaerobic or microaerophilic protozoa, primarily flagellates . Some are gut symbionts of animals such as termites , others are free-living, and others are parasitic. They include three main clades: Fornicata , Parabasalia and Preaxostyla . Fornicata (>140 species) encompasses
1633-498: Is termed protistology . Protists are a diverse group of eukaryotes (organisms whose cells possess a nucleus ) that are primarily single-celled and microscopic but exhibit a wide variety of shapes and life strategies. They have different life cycles , trophic levels , modes of locomotion , and cellular structures . Although most protists are unicellular , there is a considerable range of multicellularity amongst them; some form colonies or multicellular structures visible to
1704-536: 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
1775-868: Is they were created by symbiogenesis between an anaerobic Asgard archaean and an aerobic proteobacterium , which formed 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
1846-401: Is unclear how frequently sexual reproduction causes genetic exchange between different strains of Plasmodium in nature and most populations of parasitic protists may be clonal lines that rarely exchange genes with other members of their species. The pathogenic parasitic protists of the genus Leishmania have been shown to be capable of a sexual cycle in the invertebrate vector, likened to
1917-408: Is widespread among multicellular eukaryotes, it seemed unlikely until recently, that sex could be a primordial and fundamental characteristic of eukaryotes. The main reason for this view was that sex appeared to be lacking in certain pathogenic protists whose ancestors branched off early from the eukaryotic family tree. However, several of these "early-branching" protists that were thought to predate
1988-1178: 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] Aphelida Aphelida
2059-518: 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
2130-498: The bicosoecids , phagotrophic flagellates that consume bacteria, and the closely related Placidozoa , which consists of several groups of heterotrophic flagellates (e.g., the deep-sea halophilic Placididea ) as well as the intestinal commensals known as Opalinata (e.g., the human parasite Blastocystis , and the highly unusual opalinids , composed of giant cells with numerous nuclei and cilia, originally misclassified as ciliates). The alveolates (Alveolata) are characterized by
2201-727: The bigyromonads , a group of bacterivorous or eukaryovorous phagotrophs. A small group of heliozoan-like heterotrophic amoebae, Actinophryida , has an uncertain position, either within or as the sister taxon of Ochrophyta. The little studied phylum Bigyra is an assemblage of exclusively heterotrophic organisms, most of which are free-living. It includes the Labyrinthulomycetes , among which are single-celled amoeboid phagotrophs, mixotrophs, and fungus-like filamentous heterotrophs that create slime networks to move and absorb nutrients, as well as some parasites. Also included in Bigyra are
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2272-583: 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
2343-544: The diplomonads , with two nuclei (e.g., Giardia , genus of well-known parasites of humans), and several smaller groups of free-living, commensal and parasitic protists (e.g., Carpediemonas , retortamonads ). Parabasalia (>460 species) is a varied group of anaerobic, mostly endobiotic organisms, ranging from small parasites (like Trichomonas vaginalis , another human pathogen) to giant intestinal symbionts with numerous flagella and nuclei found in wood-eating termites and cockroaches . Preaxostyla (~140 species) includes
2414-558: The domain of Eukaryota or Eukarya , organisms whose cells have 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
2485-528: The heterotrophic protists, known as protozoa , were considered part of the animal kingdom , while the phototrophic ones, called algae , were studied as part of the plant kingdom . Even after the creation of a separate protist kingdom, some minuscule animals (the myxozoans ) and 'lower' fungi (namely the aphelids , rozellids and microsporidians , collectively known as Opisthosporidia ) were studied as protists, and some algae (particularly red and green algae ) remained classified as plants. According to
2556-426: The monophyly of Bigyra is being questioned. Branching outside both Bigyra and Gyrista is a single species of enigmatic heterotrophic flagellates, Platysulcus tardus . Much of the diversity of heterotrophic stramenopiles is still uncharacterized, known almost entirely from lineages of genetic sequences known as MASTs (MArine STramenopiles), of which only a few species have been described. The phylum Gyrista includes
2627-480: The sequencing of entire genomes and transcriptomes , and electron microscopy studies of the flagellar apparatus and cytoskeleton . New major lineages of protists and novel biodiversity continue to be discovered, resulting in dramatic changes to the eukaryotic tree of life. The newest classification systems of eukaryotes do not recognize the formal taxonomic ranks (kingdom, phylum, class, order...) and instead only recognize clades of related organisms, making
2698-514: 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
2769-575: The TSAR clade is Telonemia , a small (7 species) phylum of obscure phagotrophic predatory flagellates, found in marine and freshwater environments. They share some cellular similarities with the remaining three clades: Rhizaria , Alveolata and Stramenopiles , collectively known as the SAR supergroup . Another highly diverse clade within Diaphoretickes is Archaeplastida , which houses land plants and
2840-623: The advent of phylogenetic analysis and electron microscopy studies, the use of Protista as a formal taxon was gradually abandoned. In modern classifications, protists are spread across several eukaryotic clades called supergroups , such as Archaeplastida ( photoautotrophs that includes land plants), SAR , Obazoa (which includes fungi and animals), Amoebozoa and Excavata . Protists represent an extremely large genetic and ecological diversity in all environments, including extreme habitats. Their diversity, larger than for all other eukaryotes, has only been discovered in recent decades through
2911-475: The anaerobic and endobiotic oxymonads , with modified mitochondria , and two genera of free-living microaerophilic bacterivorous flagellates Trimastix and Paratrimastix , with typical excavate morphology. Two genera of anaerobic flagellates of recent description and unique cell architecture, Barthelona and Skoliomonas , are closely related to the Fornicata. The malawimonads (Malawimonadida) are
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2982-447: The botanical ( ICN ) and the zoological ( ICZN ) codes of nomenclature . Protists display a wide range of distinct morphologies that have been used to classify them for practical purposes, although most of these categories do not represent evolutionary cohesive lineages or clades and have instead evolved independently several times. The most recognizable types are: In general, protists are typical eukaryotic cells that follow
3053-709: 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
3124-404: 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
3195-435: The clade Myzozoa , whose common ancestor acquired chloroplasts through a secondary endosymbiosis from a red alga. One branch of Myzozoa contains the apicomplexans and their closest relatives, a small clade of flagellates known as Chrompodellida where phototrophic and heterotrophic flagellates, called chromerids and colpodellids respectively, are evolutionarily intermingled. In contrast, the apicomplexans ( Apicomplexa ) are
3266-477: The classification more stable in the long term and easier to update. In this new cladistic scheme, the protists are divided into various branches informally named supergroups . Most photosynthetic eukaryotes fall under the Diaphoretickes clade, which contains the supergroups Archaeplastida (which includes plants) and TSAR (including Telonemia , Stramenopiles , Alveolata and Rhizaria ), as well as
3337-444: The common characteristic of a ventral groove in the cell used for suspension feeding , which is considered to be an ancestral trait present in the last eukaryotic common ancestor . The Excavata is composed of three clades: Discoba , Metamonada and Malawimonadida , each including 'typical excavates' that are free-living phagotrophic flagellates with the characteristic ventral groove. According to most phylogenetic analyses, this group
3408-427: 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
3479-498: The current consensus, the term 'protist' specifically excludes animals, embryophytes (land plants) —meaning that all algae fall under this category— and all fungi, although lower fungi are often studied by protistologists and mycologists alike. The names of some protists (called ambiregnal protists), because of their mixture of traits similar to both animals and plants or fungi (e.g. slime molds and flagellated algae like euglenids ), have been published under either or both of
3550-622: 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
3621-757: The dinoflagellates and their closest relatives, the perkinsids ( Perkinsozoa ), a small group (26 species) of aquatic intracellular parasites which have lost their photosynthetic ability similarly to apicomplexans. They reproduce through flagellated spores that infect dinoflagellates, molluscs and fish . In contrast, the dinoflagellates ( Dinoflagellata ) are a highly diversified (~4,500 species) group of aquatic algae that have mostly retained their chloroplasts, although many lineages have lost their own and instead either live as heterotrophs or reacquire new chloroplasts from other sources, including tertiary endosymbiosis and kleptoplasty . Most dinoflagellates are free-living and compose an important portion of phytoplankton, as well as
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#17327722846263692-447: The emergence of meiosis and sex (such as Giardia lamblia and Trichomonas vaginalis ) are now known to descend from ancestors capable of meiosis and meiotic recombination , because they have a set core of meiotic genes that are present in sexual eukaryotes. Most of these meiotic genes were likely present in the common ancestor of all eukaryotes , which was likely capable of facultative (non-obligate) sexual reproduction. This view
3763-406: 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
3834-484: 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
3905-646: 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
3976-530: The meiosis undertaken in the trypanosomes. The species diversity of protists is severely underestimated by traditional methods that differentiate species based on morphological characteristics. The number of described protist species is very low (ranging from 26,000 to 74,400 as of 2012) in comparison to the diversity of plants, animals and fungi, which are historically and biologically well-known and studied. The predicted number of species also varies greatly, ranging from 1.4×10 to 1.6×10, and in several groups
4047-401: The naked eye. The term 'protist' is defined as a paraphyletic group of all eukaryotes that are not animals , plants or fungi . Because of this definition by exclusion, protists encompass almost all of the broad spectrum of biological characteristics expected in eukaryotes. The distinction between protists and the other three eukaryotic kingdoms has been difficult to settle. Historically,
4118-856: The number of predicted species is arbitrarily doubled. Most of these predictions are highly subjective. Molecular techniques such as environmental DNA barcoding have revealed a vast diversity of undescribed protists that accounts for the majority of eukaryotic sequences or operational taxonomic units (OTUs), dwarfing those from plants, animals and fungi. As such, it is considered that protists dominate eukaryotic diversity. Stramenopiles Alveolata Rhizaria Telonemia Haptista Microhelida Cryptista Archaeplastida Provora Hemimastigophora Meteora sporadica Discoba Metamonada Ancyromonadida Malawimonadida CRuMs Amoebozoa Breviatea Apusomonadida Opisthokonta The evolutionary relationships of protists have been explained through molecular phylogenetics ,
4189-449: 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
4260-441: The photosynthetic Ochrophyta or Heterokontophyta (>23,000 species), which contain chloroplasts originated from a red alga . Among these are many lineages of algae that encompass a wide range of structures and morphologies. The three most diverse ochrophyte classes are: the diatoms , unicellular or colonial organisms encased in silica cell walls ( frustules ) that exhibit widely different shapes and ornamentations, responsible for
4331-524: The phyla Cryptista and Haptista . The animals and fungi fall into the Amorphea supergroup, which contains the phylum Amoebozoa and several other protist lineages. Various groups of eukaryotes with primitive cell architecture are collectively known as the Excavata . Excavata is a group that encompasses diverse protists, mostly flagellates, ranging from aerobic and anaerobic predators to phototrophs and heterotrophs. The common name 'excavate' refers to
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#17327722846264402-536: The presence of cortical alveoli , cytoplasmic sacs underlying the cell membrane of unknown physiological function. Among them are three of the most well-known groups of protists: apicomplexans, dinoflagellates and ciliates. The ciliates ( Ciliophora ) are a highly diverse (>8,000 species) and probably the most thoroughly studied group of protists. They are mostly free-living microbes characterized by large cells covered in rows of cilia and containing two kinds of nuclei, micronucleus and macronucleus (e.g., Paramecium ,
4473-417: 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
4544-472: The remaining eukaryotes. Protists generally reproduce asexually under favorable environmental conditions, but tend to reproduce sexually under stressful conditions, such as starvation or heat shock. Oxidative stress , which leads to DNA damage , also appears to be an important factor in the induction of sex in protists. Eukaryotes emerged in evolution more than 1.5 billion years ago. The earliest eukaryotes were protists. Although sexual reproduction
4615-498: 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
4686-422: The same principles of physiology and biochemistry described for those cells within the "higher" eukaryotes (animals, fungi or plants): they are aerobic organisms that consume oxygen to produce energy through mitochondria , and those with chloroplasts perform carbon fixation through photosynthesis in chloroplasts . However, many have evolved a variety of unique physiological adaptations that do not appear in
4757-487: The sister clade to Ochrophyta are the predominantly osmotrophic and filamentous Pseudofungi (>1,200 species), which include three distinct lineages: the parasitic oomycetes or water moulds (e.g., Phytophthora infestans , the agent behind the Irish Potato Famine ), which encompass most of the pseudofungi species; the less diverse non-parasitic hyphochytrids that maintain a fungus-like lifestyle; and
4828-717: The study of environmental DNA and is still in the process of being fully described. They are present in all ecosystems as important components of the biogeochemical cycles and trophic webs . They exist abundantly and ubiquitously in a variety of forms that evolved multiple times independently, such as free-living algae , amoebae and slime moulds , or as important parasites . Together, they compose an amount of biomass that doubles that of animals. They exhibit varied types of nutrition (such as phototrophy , phagotrophy or osmotrophy ), sometimes combining them (in mixotrophy ). They present unique adaptations not present in multicellular animals, fungi or land plants. The study of protists
4899-503: 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
4970-578: Was further supported by a 2011 study on amoebae . Amoebae have been regarded as asexual organisms , but the study describes evidence that most amoeboid lineages are ancestrally sexual, and that the majority of asexual groups likely arose recently and independently. Even in the early 20th century, some researchers interpreted phenomena related to chromidia ( chromatin granules free in the cytoplasm ) in amoebae as sexual reproduction. Some commonly found protist pathogens such as Toxoplasma gondii are capable of infecting and undergoing asexual reproduction in
5041-477: Was the last supergroup to be described, because it lacks any defining characteristic and was discovered exclusively through molecular phylogenetics . Three major clades are included, namely the phyla Cercozoa , Endomyxa and Retaria . Retaria contains the most familiar rhizarians: forams and radiolarians , two groups of large free-living marine amoebae with pseudopodia supported by microtubules , many of which are macroscopic. The radiolarians (Radiolaria) are
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