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Lichen

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:''For other things named "lichen", see: Lichen (disambiguation).'' Image:Haeckel Lichenes.jpg thumb|230px|"Lichenes" from [[Ernst Haeckel's ''Artforms of Nature'', 1904]] image:N2 Lichen.jpg thumb|230 px|right|Crustose and foliose lichens on a wall image:Lichen_squamulose.jpg thumb|right|230px|A foliose lichen on basalt. '''Lichens''' are symbiosis symbiotic organisms made up by the association of microscopic green algae or cyanobacteria and filamentous fungi. There is evidence that lichens might involve a controlled form of parasitism of the algal cells. In laboratory settings, algae grow faster when they are alone rather than when they are part of a lichen. Lichens take the external shape of the fungal partner and hence are named based on the fungus. The fungus most commonly forms the majority of a lichen's bulk, though in filamentous and gelatinous lichens this may not always be the case. The lichen fungus is typically a member of the Ascomycota—rarely a member of the Basidiomycota. Some lichen taxonomists place lichens in their own division, the '''Mycophycophyta''', but this practice ignores the fact that the components belong to separate Scientific classification lineages. The algal cells contain chlorophyll, permitting them to live in a purely mineral environment by producing their own organic compounds (see photosynthesis). The fungus protects the alga against drying out and, in some cases, provides it with minerals obtained from the substratum. If a Cyanobacteria cyanobacterium is present, as in certain terricolous lichens, this can nitrogen fixation fix atmospheric nitrogen, complementing the activities of the green alga.

Morphology and structure
Lichens live on various surfaces, such as soil, wood, and rock. They are often the first to settle in places lacking soil, constituting the sole vegetation in some extreme environments such as those found at high mountain elevations and at high latitudes. Some survive in the tough conditions of deserts, and others on frozen soil of the arctic regions. Recent NASA research shows that lichens can even endure extended exposure to space. Some lichens have the aspect of leaves (foliose lichens); others cover the substratum like a crust (crustose lichens); others adopt shrubby forms (fruticose lichens); and there are gelatinous lichens (see lichen forms to the right -->). image:Lichen_fruiticose.jpg thumb|right|230px|''Usnea australis'', a fruticose form, growing on a tree branch Image:Lichen02.jpg thumb|right|230px|[[Map lichen (''Rhizocarpon geographicum'') on rock]] image:Hyella caespitosa hypae.jpg thumb|right|230px|The cyanobacterium ''Hyella caespitosa'' with fungal hyphae in the lichen ''Pyrenocollema halodytes'' Although the form of a lichen is determined by the DNA genetic material of the fungal partner, association with a photobiont is required for the development of that form. When grown in the laboratory in the absence of its photobiont, a lichen fungus develops as an undifferentiated mass of hyphae. If combined with its photobiont under appropriate conditions, the morphogenesis of the lichen occurs and its characteristic form emerges. (Brodo, Sharnoff & Sharnoff, 2001) There is evidence to suggest that the lichen symbiosis is parasitism parasitic rather than mutualism mutualistic. The photosynthetic partner can exist in nature independently of the fungal partner, but not vice versa. Furthermore, photobiont cells are routinely destroyed in the course of nutrient exchange. The association is able to continue because photobiont cells reproduce faster than they are destroyed. (Ibid.) When seen under magnification, a section through a typical foliose lichen Thallus_(tissue) thallus reveals four layers of interlaced filaments (fungus). The upper layer is formed by densely agglutinated fungal hyphae building a protective outer layer called the cortex. Cyanobacteria may be held in small eruptions of or under the surface, called cephalopodium cephalopodia. Beneath the upper cortex is an algal layer composed of algal cells embedded in rather densely interwoven fungal hyphae. Each cell or group of cells of the photobiont is usually individually wrapped by hyphae, and in some cases penetrated by an haustorium. Beneath the algal layer is a layer of loosely interwoven fungal hyphae without algal cells. This layer is called the medulla. Beneath the medulla, the bottom surface resembles the upper surface and is called the lower cortex, consisting of densely packed fungal hyphae. The lower cortex often bears structures, such as rhizines or a tomentum, serving to attach the thallus to the substratum on which it grows. Lichens also sometimes contain structures made from fungal metabolites, for example crustose lichens sometimes have a polysaccharide layer in the cortex. Although each lichen thallus generally appears homogeneous, it may consist of several different species of fungus and photobiont living together.

Reproduction
Image:Lichen reproduction.jpg thumb|230px|Thalli and apothecia on a foliose lichen Lichens most frequently reproduce asexually, either by vegetative reproduction or through the dispersal of diaspores containing algal and fungal cells. '''Soredia''' (sing. soredium) are small groups of algal cells surrounded by fungal filaments that form in cavities called soralia, which open when the lichen dries or surrounding tissues die and release the soredia to be dispersed by wind. Another form of diaspore are isidia, elongated outgrowths from the thallus that break off for dispersal. Fruticose lichens in particular can easily fragment. Due to the relative lack of differentiation in the thallus, the line between diaspore formation and vegetative reproduction is often blurred. Many lichens break up into fragments when they dry, dispersing themselves to resume growth when moisture returns. Lichens also reproduce sexually in a manner typical of fungi, forming fungal and algal "propagules" that, following germination, must meet with a compatible partner before a functional lichen can form. This is generally not a common means of reproduction for most lichens, though it is more common in basidiomycetous lichens since they appear to lack structures specifically designed for asexual reproduction. Spores are produced in spore-producing bodies, the three most common spore body types are the '''apothecia''', '''perithecia''' and the '''pycnidia'''.

Ecology
Image:CladonioPinetum.jpg right|thumb|230px|Pine forest with lichen ground-cover Because lichens are morphologically small relative to most terrestrial plants, yet require access to sunlight in order to grow, most forms are attached to either large boulders, other inert surfaces, or woody plants in somewhat to completely open or exposed situations. However, where adequate moisture exists, lichens develop on surfaces (particularly those of slow-growing trees) in forests as part of an epiphyte community. Stability (that is, longevity) of a surface is a commonality of most lichen habitats. Lichens are relatively slow-growing, and do not always compete well against even similarly low-growing moss mosses or liverwort liverworts where conditions of adequate moisture will favor the latter. Lichens are a part of the food available for many animals, such as reindeer, living in arctic regions. The larva larvae of a surprising number of Lepidoptera species feed exclusively on lichens. These include Common Footman and Marbled Beauty. However, Lichen is very low in protein and high in carbohydrates, making it unsuitable for some animals. Lichen is also used by the Northern Flying squirrel for nesting and a water source during winter. Although lichens typically grow in harsh environments in nature, many lichens are sensitive to man-made pollution pollutants. Hence, they have potential as pollution indicator organisms. When growing on mineral surfaces, some lichens slowly degrade their substrate by secreting acids that dissolve the minerals, contributing to the process of weathering by which rocks are gradually turned into soil. While this contribution to weathering is usually benign, it can cause problems for man-made stone structures. For example, there is an ongoing lichen growth problem on Mount Rushmore National Memorial that requires the employment of mountain-climbing conservators to clean the monument. Many lichens produce secondary compounds, including pigments that reduce harmful amounts of sunlight and powerful toxins the reduce herbivory or kill bacteria. These compounds are very useful for lichen identification, and have (or had) economic importance as dyes or primitive antibiotics. Extracts from many Usnea http://www.tcbmed.com/Newsletters/Volume4-Issue4-Usnea.html //www.tcbmed.com/Newsletters/Volume4-Issue4-Usnea.html species were used to treat wounds in Russia in the mid twentieth century. Orcein and other lichen dyes have largely been replaced by synthetic versions [http://waynesword.palomar.edu/ecoph5.htm]. The European Space Agency has discovered that lichens can survive unprotected in space. In an experiment led by Leopoldo Sancho from the Complutense University of Madrid, two species of lichen – Rhizocarpon geographicum and Xanthoria elegans – were sealed in a capsule and launched on a Russian Soyuz rocket on 31 May 2005. The Lichens were exposed to temperatures ranging from -20°C to 20°C, the vacuum of space and radiation for 15 days. Afterward, they were found to be in the same state as at the start of the experiment.

Growth Form
Lichens are informally classified by growth form into: * crustose (paint-like, flat), e.g., ''Caloplaca flavescens'' * filamentose (hair-like), e.g., ''Ephebe lanata'' * foliose (leafy), e.g., ''Hypogymnia physodes'' * fruticose (branched), e.g., ''Cladina evensii'', ''C. subtenuis'', and ''Usnea australis'' * leprose (powdery), e.g., ''Lepraria incana'' * squamulose (consisting of small scale-like structures, lacking a lower cortex), e.g., ''Normandina pulchella'' * gelatinous lichens, in which the cyanobacteria produce a polysaccharide that absorbs and retains water.

References
Brodo, I.M., S.D. Sharnoff, and S. Sharnoff, 2001. Lichens of North America. Yale University Press, New Haven. http://www.newscientistspace.com/article/dn8297 Hardy lichen shown to survive in space http://www.lichen.com

External links
{{Commons|Category:Lichenes}}
- University of Sydney lichen biology
- Crustose species of lichen order Caliciales in Norway
- ESA article on lichen survivability in low earth orbit Category:Lichens Category:Symbiosis Category:Mycology Category: cryptogams {{Link FA|nl}} {{Link FA|de}} ar:شيبيات ca:Liquen da:Lav (symbiotisk organisme) de:Flechte et:Samblikud el:Λειχήνα es:Liquen eo:Likeno fr:Lichen gl:Lique it:Licheni he:חזזית la:Lichen lt:Kerpės mk:Лишаи nl:Korstmos ja:地衣類 no:Lav pl:Porosty pt:Líquen ru:Лишайники fi:Jäkälä sv:Lav wa:Liken zh:地衣

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