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44-515: Levänluhta ( Finnish: [ˈleʋænˌluhtɑ] ; lit. " alga 's floodmeadow ") or Leväluhta is a spring and prominent archaeological site in the village of Orismala in Isokyrö , Finland . It was used as an Iron Age water graveyard from the 5th to the 8th centuries, containing remains of around 100 individuals. It's one of the oldest known burial sites in Finland with preserved human bones, as

88-571: A and b . Their chloroplasts are surrounded by four and three membranes, respectively, and were probably retained from ingested green algae. Chlorarachniophytes , which belong to the phylum Cercozoa , contain a small nucleomorph , which is a relict of the algae's nucleus . Euglenids , which belong to the phylum Euglenozoa , live primarily in fresh water and have chloroplasts with only three membranes. The endosymbiotic green algae may have been acquired through myzocytosis rather than phagocytosis . (Another group with green algae endosymbionts

132-408: A and c , and phycobilins. The shape can vary; they may be of discoid, plate-like, reticulate, cup-shaped, spiral, or ribbon shaped. They have one or more pyrenoids to preserve protein and starch. The latter chlorophyll type is not known from any prokaryotes or primary chloroplasts, but genetic similarities with red algae suggest a relationship there. In the first three of these groups ( Chromista ),

176-528: A polyphyletic group since they do not include a common ancestor , and although their chlorophyll -bearing plastids seem to have a single origin (from symbiogenesis with cyanobacteria ), they were acquired in different ways. Green algae are a prominent examples of algae that have primary chloroplasts derived from endosymbiont cyanobacteria. Diatoms and brown algae are examples of algae with secondary chloroplasts derived from endosymbiotic red algae , which they acquired via phagocytosis . Algae exhibit

220-539: A form and capabilities not possessed by the symbiont species alone (they can be experimentally isolated). The photobiont possibly triggers otherwise latent genes in the mycobiont. Trentepohlia is an example of a common green alga genus worldwide that can grow on its own or be lichenised. Lichen thus share some of the habitat and often similar appearance with specialized species of algae ( aerophytes ) growing on exposed surfaces such as tree trunks and rocks and sometimes discoloring them. Coral reefs are accumulated from

264-598: A lineage that eventually led to the higher land plants. The innovation that defines these nonalgal plants is the presence of female reproductive organs with protective cell layers that protect the zygote and developing embryo. Hence, the land plants are referred to as the Embryophytes . The term algal turf is commonly used but poorly defined. Algal turfs are thick, carpet-like beds of seaweed that retain sediment and compete with foundation species like corals and kelps , and they are usually less than 15 cm tall. Such

308-483: A red dye derived from it. The Latinization, fūcus , meant primarily the cosmetic rouge. The etymology is uncertain, but a strong candidate has long been some word related to the Biblical פוך ( pūk ), 'paint' (if not that word itself), a cosmetic eye-shadow used by the ancient Egyptians and other inhabitants of the eastern Mediterranean. It could be any color: black, red, green, or blue. The study of algae

352-468: A turf may consist of one or more species, and will generally cover an area in the order of a square metre or more. Some common characteristics are listed: Many algae, particularly species of the Characeae , have served as model experimental organisms to understand the mechanisms of the water permeability of membranes, osmoregulation , turgor regulation , salt tolerance , cytoplasmic streaming , and

396-989: A wide range of reproductive strategies, from simple asexual cell division to complex forms of sexual reproduction via spores . Algae lack the various structures that characterize plants (which evolved from freshwater green algae), such as the phyllids (leaf-like structures) and rhizoids of bryophytes ( non-vascular plants ), and the roots , leaves and other xylemic / phloemic organs found in tracheophytes ( vascular plants ). Most algae are autotrophic , although some are mixotrophic , deriving energy both from photosynthesis and uptake of organic carbon either by osmotrophy , myzotrophy or phagotrophy . Some unicellular species of green algae, many golden algae , euglenids , dinoflagellates , and other algae have become heterotrophs (also called colorless or apochlorotic algae), sometimes parasitic , relying entirely on external energy sources and have limited or no photosynthetic apparatus. Some other heterotrophic organisms, such as

440-454: Is an informal term for any organisms of a large and diverse group of photosynthetic eukaryotes , which include species from multiple distinct clades . Such organisms range from unicellular microalgae such as Chlorella , Prototheca and the diatoms , to multicellular macroalgae such as the giant kelp , a large brown alga which may grow up to 50 metres (160 ft) in length. Most algae are aquatic organisms and lack many of

484-550: Is most commonly called phycology (from Greek phykos  'seaweed'); the term algology is falling out of use. One definition of algae is that they "have chlorophyll as their primary photosynthetic pigment and lack a sterile covering of cells around their reproductive cells ". On the other hand, the colorless Prototheca under Chlorophyta are all devoid of any chlorophyll. Although cyanobacteria are often referred to as "blue-green algae", most authorities exclude all prokaryotes , including cyanobacteria, from

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528-544: Is quite different from those of reds and browns, because they have distinct nodes, separated by internode 'stems'; whorls of branches reminiscent of the horsetails occur at the nodes. Conceptacles are another polyphyletic trait; they appear in the coralline algae and the Hildenbrandiales , as well as the browns. Most of the simpler algae are unicellular flagellates or amoeboids , but colonial and nonmotile forms have developed independently among several of

572-528: Is the Latin word for 'seaweed' and retains that meaning in English. The etymology is obscure. Although some speculate that it is related to Latin algēre , 'be cold', no reason is known to associate seaweed with temperature. A more likely source is alliga , 'binding, entwining'. The Ancient Greek word for 'seaweed' was φῦκος ( phŷkos ), which could mean either the seaweed (probably red algae) or

616-423: Is the dinoflagellate genus Lepidodinium , which has replaced its original endosymbiont of red algal origin with one of green algal origin. A nucleomorph is present, and the host genome still have several red algal genes acquired through endosymbiotic gene transfer. Also the euglenid and chlorarachniophyte genome contain genes of apparent red algal ancestry) These groups have chloroplasts containing chlorophylls

660-476: Is thus protected from predators; the sponge is provided with oxygen and sugars which can account for 50 to 80% of sponge growth in some species. Non-vascular plant Non-vascular plants include two distantly related groups: These groups are sometimes called "lower plants", referring to their status as the earliest plant groups to evolve, but the usage is imprecise since both groups are polyphyletic and may be used to include vascular cryptogams , such as

704-751: The Calymmian period , early in Boring Billion , but it is difficult to track the key events because of so much time gap. Primary symbiogenesis gave rise to three divisions of archaeplastids , namely the Viridiplantae ( green algae and later plants ), Rhodophyta ( red algae ) and Glaucophyta ("grey algae"), whose plastids further spread into other protist lineages through eukaryote-eukaryote predation , engulfments and subsequent endosymbioses (secondary and tertiary symbiogenesis). This process of serial cell "capture" and "enslavement" explains

748-614: The Infusoria (microscopic organisms). Unlike macroalgae , which were clearly viewed as plants, microalgae were frequently considered animals because they are often motile. Even the nonmotile (coccoid) microalgae were sometimes merely seen as stages of the lifecycle of plants, macroalgae, or animals. Although used as a taxonomic category in some pre-Darwinian classifications, e.g., Linnaeus (1753), de Jussieu (1789), Lamouroux (1813), Harvey (1836), Horaninow (1843), Agassiz (1859), Wilson & Cassin (1864), in further classifications,

792-612: The International Association for Lichenology to be "an association of a fungus and a photosynthetic symbiont resulting in a stable vegetative body having a specific structure". The fungi, or mycobionts, are mainly from the Ascomycota with a few from the Basidiomycota . In nature, they do not occur separate from lichens. It is unknown when they began to associate. One or more mycobiont associates with

836-589: The Late Cambrian / Early Ordovician period, from sessile shallow freshwater charophyte algae much like Chara , which likely got stranded ashore when riverine / lacustrine water levels dropped during dry seasons . These charophyte algae probably already developed filamentous thalli and holdfasts that superficially resembled plant stems and roots , and probably had an isomorphic alternation of generations . They perhaps evolved some 850 mya and might even be as early as 1  Gya during

880-986: The Vindhya basin have been dated to 1.6 to 1.7 billion years ago. Because of the wide range of algae types, they have increasingly different industrial and traditional applications in human society. Traditional seaweed farming practices have existed for thousands of years and have strong traditions in East Asia food cultures. More modern algaculture applications extend the food traditions for other applications, including cattle feed, using algae for bioremediation or pollution control, transforming sunlight into algae fuels or other chemicals used in industrial processes, and in medical and scientific applications. A 2020 review found that these applications of algae could play an important role in carbon sequestration to mitigate climate change while providing lucrative value-added products for global economies. The singular alga

924-601: The apicomplexans are also parasites derived from ancestors that possessed plastids, but are not included in any group traditionally seen as algae. Algae are polyphyletic thus their origin cannot be traced back to single hypothetical common ancestor . It is thought that they came into existence when photosynthetic coccoid cyanobacteria got phagocytized by a unicellular heterotrophic eukaryote (a protist ), giving rise to double-membranous primary plastids . Such symbiogenic events (primary symbiogenesis) are believed to have occurred more than 1.5 billion years ago during

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968-435: The apicomplexans , are also derived from cells whose ancestors possessed chlorophyllic plastids, but are not traditionally considered as algae. Algae have photosynthetic machinery ultimately derived from cyanobacteria that produce oxygen as a byproduct of splitting water molecules , unlike other organisms that conduct anoxygenic photosynthesis such as purple and green sulfur bacteria . Fossilized filamentous algae from

1012-545: The calcareous exoskeletons of marine invertebrates of the order Scleractinia (stony corals ). These animals metabolize sugar and oxygen to obtain energy for their cell-building processes, including secretion of the exoskeleton, with water and carbon dioxide as byproducts. Dinoflagellates (algal protists) are often endosymbionts in the cells of the coral-forming marine invertebrates, where they accelerate host-cell metabolism by generating sugar and oxygen immediately available through photosynthesis using incident light and

1056-497: The ferns and fern allies that reproduce using spores. Non-vascular plants are often among the first species to move into new and inhospitable territories, along with prokaryotes and protists , and thus function as pioneer species . Mosses and leafy liverworts have structures called phyllids that resemble leaves , but only consist of single sheets of cells with no internal air spaces, no cuticle or stomata , and no xylem or phloem. Consequently, phyllids are unable to control

1100-568: The waterlogged environment had contributed to an exceptionally good preservation of the remains. Ancient DNA of 13 individuals from Levänluhta has been further studied, revealing that most of them had some Siberian-derived genetic ancestry like all Fenno-Ugric peoples, with a slightly closer resemblance to Sami than Finnish people , but the difference was not considerable. Algae Algae ( UK : / ˈ æ l ɡ iː / AL -ghee , US : / ˈ æ l dʒ iː / AL -jee ; sg. : alga / ˈ æ l ɡ ə / AL -gə )

1144-720: The "algae" are seen as an artificial, polyphyletic group. Throughout the 20th century, most classifications treated the following groups as divisions or classes of algae: cyanophytes , rhodophytes , chrysophytes , xanthophytes , bacillariophytes , phaeophytes , pyrrhophytes ( cryptophytes and dinophytes ), euglenophytes , and chlorophytes . Later, many new groups were discovered (e.g., Bolidophyceae ), and others were splintered from older groups: charophytes and glaucophytes (from chlorophytes), many heterokontophytes (e.g., synurophytes from chrysophytes, or eustigmatophytes from xanthophytes), haptophytes (from chrysophytes), and chlorarachniophytes (from xanthophytes). With

1188-675: The abandonment of plant-animal dichotomous classification, most groups of algae (sometimes all) were included in Protista , later also abandoned in favour of Eukaryota . However, as a legacy of the older plant life scheme, some groups that were also treated as protozoans in the past still have duplicated classifications (see ambiregnal protists ). Some parasitic algae (e.g., the green algae Prototheca and Helicosporidium , parasites of metazoans, or Cephaleuros , parasites of plants) were originally classified as fungi , sporozoans , or protistans of incertae sedis , while others (e.g.,

1232-451: The carbon dioxide produced by the host. Reef-building stony corals ( hermatypic corals ) require endosymbiotic algae from the genus Symbiodinium to be in a healthy condition. The loss of Symbiodinium from the host is known as coral bleaching , a condition which leads to the deterioration of a reef. Endosymbiontic green algae live close to the surface of some sponges, for example, breadcrumb sponges ( Halichondria panicea ). The alga

1276-420: The chloroplast has four membranes, retaining a nucleomorph in cryptomonads , and they likely share a common pigmented ancestor, although other evidence casts doubt on whether the heterokonts , Haptophyta , and cryptomonads are in fact more closely related to each other than to other groups. The typical dinoflagellate chloroplast has three membranes, but considerable diversity exists in chloroplasts within

1320-624: The composition of the three major groups of algae. Their lineage relationships are shown in the figure in the upper right. Many of these groups contain some members that are no longer photosynthetic. Some retain plastids, but not chloroplasts, while others have lost plastids entirely. Phylogeny based on plastid not nucleocytoplasmic genealogy: Cyanobacteria Glaucophytes Rhodophytes Stramenopiles Cryptophytes Haptophytes Euglenophytes Chlorarachniophytes Chlorophytes Charophytes Land plants (Embryophyta) These groups have green chloroplasts containing chlorophylls

1364-436: The definition of algae. The algae contain chloroplasts that are similar in structure to cyanobacteria. Chloroplasts contain circular DNA like that in cyanobacteria and are interpreted as representing reduced endosymbiotic cyanobacteria . However, the exact origin of the chloroplasts is different among separate lineages of algae, reflecting their acquisition during different endosymbiotic events. The table below describes

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1408-499: The distinct cell and tissue types, such as stomata , xylem and phloem that are found in land plants . The largest and most complex marine algae are called seaweeds . In contrast, the most complex freshwater forms are the Charophyta , a division of green algae which includes, for example, Spirogyra and stoneworts . Algae that are carried passively by water are plankton , specifically phytoplankton . Algae constitute

1452-425: The diversity of photosynthetic eukaryotes. Recent genomic and phylogenomic approaches have significantly clarified plastid genome evolution , the horizontal movement of endosymbiont genes to the "host" nuclear genome , and plastid spread throughout the eukaryotic tree of life . Fossils of isolated spores suggest land plants may have been around as long as 475  million years ago (mya) during

1496-432: The first to divide macroscopic algae into four divisions based on their pigmentation. This is the first use of a biochemical criterion in plant systematics. Harvey's four divisions are: red algae (Rhodospermae), brown algae (Melanospermae), green algae (Chlorospermae), and Diatomaceae. At this time, microscopic algae were discovered and reported by a different group of workers (e.g., O. F. Müller and Ehrenberg ) studying

1540-607: The functioning of peatlands. This provides essential goods and services to humans such as global carbon sinks, water purification systems, fresh water reserves as well as biodiversity and peat resources. This is achieved through nutrient acquisition from dominant plants under nutrient-stressed conditions. Non-vascular plants can also play important roles in other biomes such as deserts, tundra and alpine regions. They have been shown to contribute to soil stabilization, nitrogen fixation, carbon assimilation etc. These are all crucial components in an ecosystem in which non-vascular plants play

1584-483: The gametophyte generation is dominant. In these plants, the sporophytes grow from and are dependent on gametophytes for supply of water and mineral nutrients and photosynthate, the products of photosynthesis . Non-vascular plants play crucial roles in their environments. They often dominate certain biomes such as mires, bogs and lichen tundra where these plants perform primary ecosystem functions. Additionally, in bogs mosses host microbial communities which help support

1628-538: The genera Volvox and Corallina , and a species of Acetabularia (as Madrepora ), among the animals. In 1768, Samuel Gottlieb Gmelin (1744–1774) published the Historia Fucorum , the first work dedicated to marine algae and the first book on marine biology to use the then new binomial nomenclature of Linnaeus. It included elaborate illustrations of seaweed and marine algae on folded leaves. W. H. Harvey (1811–1866) and Lamouroux (1813) were

1672-446: The generation of action potentials . Plant hormones are found not only in higher plants, but in algae, too. Some species of algae form symbiotic relationships with other organisms. In these symbioses, the algae supply photosynthates (organic substances) to the host organism providing protection to the algal cells. The host organism derives some or all of its energy requirements from the algae. Examples are: Lichens are defined by

1716-538: The green algae Phyllosiphon and Rhodochytrium , parasites of plants, or the red algae Pterocladiophila and Gelidiocolax mammillatus , parasites of other red algae, or the dinoflagellates Oodinium , parasites of fish) had their relationship with algae conjectured early. In other cases, some groups were originally characterized as parasitic algae (e.g., Chlorochytrium ), but later were seen as endophytic algae. Some filamentous bacteria (e.g., Beggiatoa ) were originally seen as algae. Furthermore, groups like

1760-571: The group, and a number of endosymbiotic events apparently occurred. The Apicomplexa , a group of closely related parasites, also have plastids called apicoplasts , which are not photosynthetic, but appear to have a common origin with dinoflagellate chloroplasts. Linnaeus , in Species Plantarum (1753), the starting point for modern botanical nomenclature , recognized 14 genera of algae, of which only four are currently considered among algae. In Systema Naturae , Linnaeus described

1804-453: The groups. Some of the more common organizational levels, more than one of which may occur in the lifecycle of a species, are In three lines, even higher levels of organization have been reached, with full tissue differentiation. These are the brown algae, —some of which may reach 50 m in length ( kelps ) —the red algae, and the green algae. The most complex forms are found among the charophyte algae (see Charales and Charophyta ), in

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1848-417: The late phase of the Boring Billion . A range of algal morphologies is exhibited, and convergence of features in unrelated groups is common. The only groups to exhibit three-dimensional multicellular thalli are the reds and browns , and some chlorophytes . Apical growth is constrained to subsets of these groups: the florideophyte reds, various browns, and the charophytes. The form of charophytes

1892-427: The rate of water loss from their tissues and are said to be poikilohydric . Some liverworts, such as Marchantia , have a cuticle, and the sporophytes of mosses have both cuticles and stomata, which were important in the evolution of land plants . All land plants have a life cycle with an alternation of generations between a diploid sporophyte and a haploid gametophyte , but in all non-vascular land plants,

1936-405: The same phycobiont species, from the green algae, except that alternatively, the mycobiont may associate with a species of cyanobacteria (hence "photobiont" is the more accurate term). A photobiont may be associated with many different mycobionts or may live independently; accordingly, lichens are named and classified as fungal species. The association is termed a morphogenesis because the lichen has

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