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Archaeplastida

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51-408: The Archaeplastida (or kingdom Plantae sensu lato "in a broad sense"; pronounced / ɑːr k ɪ ˈ p l æ s t ɪ d ə / ) 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

102-452: A and phycobiliproteins , like most cyanobacteria, and accumulate starch outside the chloroplasts. The green algae and land plants – together known as Viridiplantae (Latin for "green plants") or Chloroplastida – are pigmented with chlorophylls a and b , but lack phycobiliproteins, and starch is accumulated inside the chloroplasts. The glaucophytes have typical cyanobacterial pigments, but their plastids (called cyanelles) differ in having

153-485: A peptidoglycan outer layer. Archaeplastida should not be confused with the older and obsolete name Archiplastideae, which refers to cyanobacteria and other groups of bacteria. The consensus in 2005, when the group consisting of the glaucophytes and red and green algae and land plants was named 'Archaeplastida', was that it was a clade , i.e. was monophyletic . Many studies published since then have provided evidence in agreement. Other studies, though, have suggested that

204-457: A polyphyletic grouping are not inherited from a common ancestor, but evolved independently. Monophyletic groups are typically characterised by shared derived characteristics ( synapomorphies ), which distinguish organisms in the clade from other organisms. An equivalent term is holophyly . The word "mono-phyly" means "one-tribe" in Greek. These definitions have taken some time to be accepted. When

255-545: A protective response. The first such plant receptors were identified in rice and in Arabidopsis thaliana . Plants have some of the largest genomes of all organisms. The largest plant genome (in terms of gene number) is that of wheat ( Triticum aestivum ), predicted to encode ≈94,000 genes and thus almost 5 times as many as the human genome . The first plant genome sequenced was that of Arabidopsis thaliana which encodes about 25,500 genes. In terms of sheer DNA sequence,

306-420: A range of physical and biotic stresses which cause DNA damage , but they can tolerate and repair much of this damage. Plants reproduce to generate offspring, whether sexually , involving gametes , or asexually , involving ordinary growth. Many plants use both mechanisms. When reproducing sexually, plants have complex lifecycles involving alternation of generations . One generation, the sporophyte , which

357-471: A single origin. This evidence is disputed. Based on the evidence to date, it is not possible to confirm or refute alternative evolutionary scenarios to a single primary endosymbiosis . Photosynthetic organisms with plastids of different origin (such as brown algae ) do not belong to the Archaeplastida. The archaeplastidans fall into two main evolutionary lines. The red algae are pigmented with chlorophyll

408-404: Is diploid (with 2 sets of chromosomes ), gives rise to the next generation, the gametophyte , which is haploid (with one set of chromosomes). Some plants also reproduce asexually via spores . In some non-flowering plants such as mosses, the sexual gametophyte forms most of the visible plant. In seed plants (gymnosperms and flowering plants), the sporophyte forms most of the visible plant, and

459-518: Is Plastida, defined as the clade sharing "plastids of primary (direct prokaryote) origin [as] in Magnolia virginiana Linnaeus 1753". Although many studies have suggested the Archaeplastida form a monophyletic group, a 2009 paper argues that they are in fact paraphyletic . The enrichment of novel red algal genes in a recent study demonstrates a strong signal for Plantae (Archaeplastida) monophyly and an equally strong signal of gene sharing history between

510-547: Is a similar process. Structures such as runners enable plants to grow to cover an area, forming a clone . Many plants grow food storage structures such as tubers or bulbs which may each develop into a new plant. Some non-flowering plants, such as many liverworts, mosses and some clubmosses, along with a few flowering plants, grow small clumps of cells called gemmae which can detach and grow. Plants use pattern-recognition receptors to recognize pathogens such as bacteria that cause plant diseases. This recognition triggers

561-496: Is known as botany , a branch of biology . All living things were traditionally placed into one of two groups, plants and animals . This classification dates from Aristotle (384–322 BC), who distinguished different levels of beings in his biology , based on whether living things had a "sensitive soul" or like plants only a "vegetative soul". Theophrastus , Aristotle's student, continued his work in plant taxonomy and classification. Much later, Linnaeus (1707–1778) created

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612-649: 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. The cells of

663-608: The Antarctic flora , consisting of algae, mosses, liverworts, lichens, and just two flowering plants, have adapted to the prevailing conditions on that southern continent. Plants are often the dominant physical and structural component of the habitats where they occur. Many of the Earth's biomes are named for the type of vegetation because plants are the dominant organisms in those biomes, such as grassland , savanna , and tropical rainforest . Monophyly In biological cladistics for

714-679: The Cretaceous so rapid that Darwin called it an " abominable mystery ". Conifers diversified from the Late Triassic onwards, and became a dominant part of floras in the Jurassic . In 2019, a phylogeny based on genomes and transcriptomes from 1,153 plant species was proposed. The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced. Both

765-457: The carpels or ovaries , which develop into fruits that contain seeds . Fruits may be dispersed whole, or they may split open and the seeds dispersed individually. Plants reproduce asexually by growing any of a wide variety of structures capable of growing into new plants. At the simplest, plants such as mosses or liverworts may be broken into pieces, each of which may regrow into whole plants. The propagation of flowering plants by cuttings

816-412: The eukaryotes which took into account morphology, biochemistry, and phylogenetics, and which had "some stability in the near term." They rejected the use of formal taxonomic ranks in favour of a hierarchical arrangement where the clade names do not signify rank. Thus, the phylum name 'Glaucophyta' and the class name 'Rhodophyceae' appear at the same level in their classification. The divisions proposed for

867-1003: The "chlorophyte algae" and the "streptophyte algae" are treated as paraphyletic (vertical bars beside phylogenetic tree diagram) in this analysis, as the land plants arose from within those groups. The classification of Bryophyta is supported both by Puttick et al. 2018, and by phylogenies involving the hornwort genomes that have also since been sequenced. Rhodophyta [REDACTED] Glaucophyta [REDACTED] Chlorophyta [REDACTED] Prasinococcales   Mesostigmatophyceae Chlorokybophyceae Spirotaenia [REDACTED] Klebsormidiales [REDACTED] Chara [REDACTED] Coleochaetales [REDACTED] Hornworts [REDACTED] Liverworts [REDACTED] Mosses [REDACTED] Lycophytes [REDACTED] [REDACTED] Gymnosperms [REDACTED] Angiosperms [REDACTED] Plant cells have distinctive features that other eukaryotic cells (such as those of animals) lack. These include

918-551: The Archaeplastida are shown below in both tabular and diagrammatic form . Archaeplastida : Below is a consensus reconstruction of the relationships of Archaeplastida with its nearest neighbours, mainly based on molecular data. Hemimastigophora [REDACTED] Provora Haptista [REDACTED] Telonemia Rhizaria [REDACTED] Stramenopiles [REDACTED] Alveolata [REDACTED] Cryptista [REDACTED] Microheliella maris Archaeplastida [REDACTED] There has been disagreement near

969-523: The Archaeplastida root, e.g. whether Cryptista emerged within the Archaeplastida. In 2014 a thorough review was published on these inconsistencies. The position of Telonemia and Picozoa are not clear. Also Hacrobia (Haptista + Cryptista) may be completely associated with the SAR clade. The SAR are often seen as eukaryote-eukaryote hybrids, contributing to the confusion in the genetic analyses. A sister of Gloeomargarita lithophora has been engulfed by an ancestor of

1020-407: The Archaeplastida typically lack centrioles and have mitochondria with flat cristae . They usually have a cell wall that contains cellulose , and food is stored in the form of starch . However, these characteristics are also shared with other eukaryotes. The main evidence that the Archaeplastida form a monophyletic group comes from genetic studies, which indicate their plastids probably had

1071-469: The Archaeplastida, leading to the plastids which are living in permanent endosymbiosis in most of the descendant lineages. Because both Gloeomargarita and related cyanobacteria, in addition to the most primitive archaeplastids, all live in freshwater, it seems the Archaeplastida originated in freshwater, and only colonized the oceans in the late Proterozoic. In 2019, a phylogeny of the Archaeplastida based on genomes and transcriptomes from 1,153 plant species

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1122-472: The algae. By the definition used in this article, plants form the clade Viridiplantae (green plants), which consists of the green algae and the embryophytes or land plants ( hornworts , liverworts , mosses , lycophytes , ferns , conifers and other gymnosperms , and flowering plants ). A definition based on genomes includes the Viridiplantae, along with the red algae and the glaucophytes , in

1173-801: The amount of cytoplasm stays the same. Most plants are multicellular . Plant cells differentiate into multiple cell types, forming tissues such as the vascular tissue with specialized xylem and phloem of leaf veins and stems , and organs with different physiological functions such as roots to absorb water and minerals, stems for support and to transport water and synthesized molecules, leaves for photosynthesis, and flowers for reproduction. Plants photosynthesize , manufacturing food molecules ( sugars ) using energy obtained from light . Plant cells contain chlorophylls inside their chloroplasts, which are green pigments that are used to capture light energy. The end-to-end chemical equation for photosynthesis is: This causes plants to release oxygen into

1224-431: The atmosphere. Green plants provide a substantial proportion of the world's molecular oxygen, alongside the contributions from photosynthetic algae and cyanobacteria. Plants that have secondarily adopted a parasitic lifestyle may lose the genes involved in photosynthesis and the production of chlorophyll. Growth is determined by the interaction of a plant's genome with its physical and biotic environment. Factors of

1275-429: The bacterium, and the other to the eukaryote that captured it. Over time, many genes from the chloroplast have been transferred to the nucleus of the host cell through endosymbiotic gene transfer (EGT). It is estimated that 6–20% of the archaeplastidan genome consist of genes transferred from the endosymbiont. The presence of such genes in the nuclei of eukaryotes without chloroplasts suggests this transfer happened early in

1326-461: The basis of the modern system of scientific classification , but retained the animal and plant kingdoms , naming the plant kingdom the Vegetabilia. When the name Plantae or plant is applied to a specific group of organisms or taxa , it usually refers to one of four concepts. From least to most inclusive, these four groupings are: There are about 382,000 accepted species of plants, of which

1377-728: The clade Archaeplastida . There are about 380,000 known species of plants, of which the majority, some 260,000, produce seeds . They range in size from single cells to the tallest trees . Green plants provide a substantial proportion of the world's molecular oxygen; the sugars they create supply the energy for most of Earth's ecosystems and other organisms , including animals, either eat plants directly or rely on organisms which do so. Grain , fruit , and vegetables are basic human foods and have been domesticated for millennia. People use plants for many purposes , such as building materials , ornaments, writing materials , and, in great variety, for medicines . The scientific study of plants

1428-404: The cladistics school of thought became mainstream in the 1960s, several alternative definitions were in use. Indeed, taxonomists sometimes used terms without defining them, leading to confusion in the early literature, a confusion which persists. The first diagram shows a phylogenetic tree with two monophyletic groups. The several groups and subgroups are particularly situated as branches of

1479-689: The classification of organisms , monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of taxa which meets these criteria: Monophyly is contrasted with paraphyly and polyphyly as shown in the second diagram. A paraphyletic grouping meets 1. but not 2., thus consisting of the descendants of a common ancestor, excepting one or more monophyletic subgroups. A polyphyletic grouping meets neither criterion, and instead serves to characterize convergent relationships of biological features rather than genetic relationships – for example, night-active primates, fruit trees, or aquatic insects. As such, these characteristic features of

1530-473: The degree of their cell organization, from isolated cells to filaments to colonies to multi-celled organisms. The earliest were unicellular, and many groups remain so today. Multicellularity evolved separately in several groups, including red algae, ulvophyte green algae , and in the green algae that gave rise to stoneworts and land plants. Because the ancestral archaeplastidan is hypothesized to have acquired its chloroplasts directly by engulfing cyanobacteria,

1581-483: The development of forests in swampy environments dominated by clubmosses and horsetails, including some as large as trees, and the appearance of early gymnosperms , the first seed plants . The Permo-Triassic extinction event radically changed the structures of communities. This may have set the scene for the evolution of flowering plants in the Triassic (~ 200  million years ago ), with an adaptive radiation in

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1632-419: The event is known as a primary endosymbiosis (as reflected in the name chosen for the group 'Archaeplastida' i.e. 'ancient plastid'). In 2013 it was discovered that one species of green algae, Cymbomonas tetramitiformis in the order Pyramimonadales , is a mixotroph and able to support itself through both phagotrophy and phototrophy . It is not yet known if this is a primitive trait and therefore defines

1683-465: The evolution of the group. Plant#Current definitions of Plantae See text Plants are the eukaryotes that form the kingdom Plantae ; they are predominantly photosynthetic . This means that they obtain their energy from sunlight , using chloroplasts derived from endosymbiosis with cyanobacteria to produce sugars from carbon dioxide and water, using the green pigment chlorophyll . Exceptions are parasitic plants that have lost

1734-413: The fact that a monophyletic group includes organisms (e.g., genera, species) consisting of all the descendants of a unique common ancestor. Conversely, the term polyphyly , or polyphyletic , builds on the ancient Greek prefix πολύς ( polús ), meaning "many, a lot of", and refers to the fact that a polyphyletic group includes organisms arising from multiple ancestral sources. By comparison,

1785-409: The gametophyte is very small. Flowering plants reproduce sexually using flowers, which contain male and female parts: these may be within the same ( hermaphrodite ) flower, on different flowers on the same plant , or on different plants . The stamens create pollen , which produces male gametes that enter the ovule to fertilize the egg cell of the female gametophyte. Fertilization takes place within

1836-450: The genes for chlorophyll and photosynthesis, and obtain their energy from other plants or fungi. Most plants are multicellular , except for some green algae. Historically, as in Aristotle's biology , the plant kingdom encompassed all living things that were not animals , and included algae and fungi . Definitions have narrowed since then; current definitions exclude the fungi and some of

1887-575: The great majority, some 283,000, produce seeds . The table below shows some species count estimates of different green plant (Viridiplantae) divisions . About 85–90% of all plants are flowering plants. Several projects are currently attempting to collect records on all plant species in online databases, e.g. the World Flora Online . Plants range in scale from single-celled organisms such as desmids (from 10  micrometres   (μm) across) and picozoa (less than 3 μm across), to

1938-401: The group is paraphyletic . To date, the situation appears unresolved, but a strong signal for Plantae (Archaeplastida) monophyly has been demonstrated in a recent study (with an enrichment of red algal genes). The assumption made here is that Archaeplastida is a valid clade. Various names have been given to the group. Some authors have simply referred to the group as plants or Plantae. However,

1989-990: The hornwort genomes that have also since been sequenced. Recent work on non-photosynthetic algae places Rhodelphidia as sister to Rhodophyta or to Glaucophyta and Viridiplantae; and Picozoa sister to that pair of groups. Picozoa [REDACTED] Rhodelphidia Rhodophyta [REDACTED] Glaucophyta [REDACTED] Chlorophyta [REDACTED] Prasinococcales   Mesostigmatophyceae Chlorokybophyceae Spirotaenia [REDACTED] Klebsormidiales [REDACTED] Chara [REDACTED] Coleochaetales Zygnematophyceae [REDACTED] Hornworts [REDACTED] Liverworts [REDACTED] Mosses [REDACTED] Lycophytes [REDACTED] Ferns [REDACTED] Gymnosperms [REDACTED] Angiosperms [REDACTED] All archaeplastidans have plastids (chloroplasts) that carry out photosynthesis and are believed to be derived from endosymbiotic cyanobacteria. In glaucophytes, perhaps

2040-517: The land 1,200  million years ago , but it was not until the Ordovician , around 450  million years ago , that the first land plants appeared, with a level of organisation like that of bryophytes. However, fossils of organisms with a flattened thallus in Precambrian rocks suggest that multicellular freshwater eukaryotes existed over 1000 mya. Primitive land plants began to diversify in

2091-412: The large water-filled central vacuole , chloroplasts , and the strong flexible cell wall , which is outside the cell membrane . Chloroplasts are derived from what was once a symbiosis of a non-photosynthetic cell and photosynthetic cyanobacteria . The cell wall, made mostly of cellulose , allows plant cells to swell up with water without bursting. The vacuole allows the cell to change in size while

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2142-507: The largest trees ( megaflora ) such as the conifer Sequoia sempervirens (up to 120 metres (380 ft) tall) and the angiosperm Eucalyptus regnans (up to 100 m (325 ft) tall). The naming of plants is governed by the International Code of Nomenclature for algae, fungi, and plants and the International Code of Nomenclature for Cultivated Plants . The ancestors of land plants evolved in water. An algal scum formed on

2193-418: The last common ancestor of Archaeplastida, which could explain how it obtained its chloroplasts, or if it is a trait regained by horizontal gene transfer . Since then more species of mixotrophic green algae, such as Pyramimonas tychotreta and Mantoniella antarctica , has been found. Evidence for primary endosymbiosis includes the presence of a double membrane around the chloroplasts; one membrane belonged to

2244-656: The late Silurian , around 420  million years ago . Bryophytes, club mosses, and ferns then appear in the fossil record. Early plant anatomy is preserved in cellular detail in an early Devonian fossil assemblage from the Rhynie chert . These early plants were preserved by being petrified in chert formed in silica-rich volcanic hot springs. By the end of the Devonian, most of the basic features of plants today were present, including roots, leaves and secondary wood in trees such as Archaeopteris . The Carboniferous period saw

2295-428: The most primitive members of the group, the chloroplast is called a cyanelle and shares several features with cyanobacteria, including a peptidoglycan cell wall, that are not retained in other members of the group. The resemblance of cyanelles to cyanobacteria supports the endosymbiotic theory . The cells of most archaeplastidans have walls, commonly but not always made of cellulose. The Archaeplastida vary widely in

2346-423: The name Plantae is ambiguous, since it has also been applied to less inclusive clades , such as Viridiplantae and embryophytes . To distinguish, the larger group is sometimes known as Plantae sensu lato ("plants in the broad sense"). To avoid ambiguity, other names have been proposed. Primoplantae, which appeared in 2004, seems to be the first new name suggested for this group. Another name applied to this node

2397-501: The physical or abiotic environment include temperature , water , light, carbon dioxide , and nutrients in the soil. Biotic factors that affect plant growth include crowding, grazing, beneficial symbiotic bacteria and fungi, and attacks by insects or plant diseases . Frost and dehydration can damage or kill plants. Some plants have antifreeze proteins , heat-shock proteins and sugars in their cytoplasm that enable them to tolerate these stresses . Plants are continuously exposed to

2448-401: The red/green algae and other lineages. This study provides insight on how rich mesophilic red algal gene data are crucial for testing controversial issues in eukaryote evolution and for understanding the complex patterns of gene inheritance in protists. The name Archaeplastida was proposed in 2005 by a large international group of authors (Adl et al. ), who aimed to produce a classification for

2499-465: The smallest published genome is that of the carnivorous bladderwort ( Utricularia gibba) at 82 Mb (although it still encodes 28,500 genes) while the largest, from the Norway spruce ( Picea abies ), extends over 19.6 Gb (encoding about 28,300 genes). Plants are distributed almost worldwide. While they inhabit several biomes which can be divided into a multitude of ecoregions , only the hardy plants of

2550-493: The tree to indicate ordered lineal relationships between all the organisms shown. Further, any group may (or may not) be considered a taxon by modern systematics , depending upon the selection of its members in relation to their common ancestor(s); see second and third diagrams. The term monophyly , or monophyletic , derives from the two Ancient Greek words μόνος ( mónos ), meaning "alone, only, unique", and φῦλον ( phûlon ), meaning "genus, species", and refers to

2601-489: Was proposed. The placing of algal groups is supported by phylogenies based on genomes from the Mesostigmatophyceae and Chlorokybophyceae that have since been sequenced. Both the "chlorophyte algae" and the "streptophyte algae" are treated as paraphyletic (vertical bars beside phylogenetic tree diagram) in this analysis. The classification of Bryophyta is supported both by Puttick et al. 2018, and by phylogenies involving

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