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Moss
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"Mosses" redirects here. For the mountain pass, see Col des Mosses.
For other uses, see Moss (disambiguation).
Moss
Temporal range: Carboniferous[1] – present
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Clumps of moss on the ground and base of trees in the Allegheny National Forest, Pennsylvania, USA.
Scientific classification
Kingdom: Plantae
Division: Bryophyta
Schimp. sensu stricto
Classes[2]
Takakiopsida
Sphagnopsida
Andreaeopsida
Andreaeobryopsida
Oedipodiopsida
Polytrichopsida
Tetraphidopsida
Bryopsida
Synonyms
Musci L.
Muscineae Bisch.
Mosses are small flowerless plants that typically grow in dense green clumps or mats, often in damp or shady locations. The individual plants are usually composed of simple leaves that are generally only one cell thick, attached to a stem that may be branched or unbranched and has only a limited role in conducting water and nutrients. Although some species have conducting tissues, these are generally poorly developed and structurally different from similar tissue found in vascular plants.[3] Mosses do not have seeds and after fertilisation develop sporophytes with unbranched stalks topped with single capsules containing spores. They are typically 0.2–10 cm (0.1–3.9 in) tall, though some species are much larger. Dawsonia, the tallest moss in the world, can grow to 50 cm (20 in) in height.
Mosses are commonly confused with lichens, hornworts, and liverworts.[4] Lichens may superficially look like mosses, and have common names that include the word "moss" (e.g., "reindeer moss" or "iceland moss"), but are not related to mosses.[4]:3 Mosses used to be grouped together with the hornworts and liverworts as "non-vascular" plants in the former division "bryophytes", all of them having the haploid gametophyte generation as the dominant phase of the life cycle. This contrasts with the pattern in all vascular plants (seed plants and pteridophytes), where the diploid sporophyte generation is dominant.
Mosses are now classified on their own as the division Bryophyta. There are approximately 12,000 species.[2]
The main commercial significance of mosses is as the main constituent of peat (mostly the genus Sphagnum), although they are also used for decorative purposes, such as in gardens and in the florist trade. Traditional uses of mosses included as insulation and for the ability to absorb liquids up to 20 times their weight.
Contents
1 Physical characteristics
1.1 Description
1.2 Life cycle
1.3 Dwarf males
1.4 DNA repair
2 Classification
3 Geological history
4 Ecology
4.1 Habitat
4.2 Moss-cyanobacteria relationship
5 Cultivation
5.1 Inhibiting moss growth
5.2 Mossery
6 Uses
6.1 Traditional
6.2 Commercial
7 See also
8 References
9 External links
Physical characteristics
Description
Chloroplasts and accumulated starch granules in Bryum capillare
Botanically, mosses are non-vascular plants in the land plant division Bryophyta. They are small (a few centimeters tall) herbaceous (non-woody) plants that absorb water and nutrients mainly through their leaves and harvest carbon dioxide and sunlight to create food by photosynthesis.[5][6] They differ from vascular plants in lacking water-bearing xylem tracheids or vessels. As in liverworts and hornworts, the haploid gametophyte generation is the dominant phase of the life cycle. This contrasts with the pattern in all vascular plants (seed plants and pteridophytes), where the diploid sporophyte generation is dominant. Mosses reproduce using spores, not seeds, and have no flowers.
Moss gametophytes have stems which may be simple or branched and upright or prostrate. Their leaves are simple, usually only a single layer of cells with no internal air spaces, often with thicker midribs. They do not have proper roots, but have threadlike rhizoids that anchor them to their substrate. Mosses do not absorb water or nutrients from their substrate through their rhizoids. They can be distinguished from liverworts (Marchantiophyta or Hepaticae) by their multi-cellular rhizoids. Spore-bearing capsules or sporangia of mosses are borne singly on long, unbranched stems, thereby distinguishing them from the polysporangiophytes, which include all vascular plants. The spore-bearing sporophytes (i.e. the diploid multicellular generation) are short-lived and dependent on the gametophyte for water supply and nutrition. Also, in most mosses, the spore-bearing capsule enlarges and matures after its stalk elongates, while in liverworts the capsule enlarges and matures before its stalk elongates.[6] Other differences are not universal for all mosses and all liverworts, but the presence of clearly differentiated stem with simple-shaped, ribbed leaves, without deeply lobed or segmented leaves and not arranged in three ranks, all point to the plant being a moss.
Life cycle
Vascular plants have two sets of chromosomes in their vegetative cells and are said to be diploid, i.e. each chromosome has a partner that contains the same, or similar, genetic information. By contrast, mosses and other bryophytes have only a single set of chromosomes and so are haploid (i.e. each chromosome exists in a unique copy within the cell). There is a period in the moss life cycle when they do have a double set of paired chromosomes, but this happens only during the sporophyte stage.
Life cycle of a typical moss (Polytrichum commune)
The moss life-cycle starts with a haploid spore that germinates to produce a protonema (pl. protonemata), which is either a mass of thread-like filaments or thalloid (flat and thallus-like). Massed moss protonemata typically look like a thin green felt, and may grow on damp soil, tree bark, rocks, concrete, or almost any other reasonably stable surface. This is a transitory stage in the life of a moss, but from the protonema grows the gametophore ("gamete-bearer") that is structurally differentiated into stems and leaves. A single mat of protonemata may develop several gametophore shoots, resulting in a clump of moss.
From the tips of the gametophore stems or branches develop the sex organs of the mosses. The female organs are known as archegonia (sing. archegonium) and are protected by a group of modified leaves known as the perichaetum (plural, perichaeta). The archegonia are small flask-shaped clumps of cells with an open neck (venter) down which the male sperm swim. The male organs are known as antheridia (sing. antheridium) and are enclosed by modified leaves called the perigonium (pl. perigonia). The surrounding leaves in some mosses form a splash cup, allowing the sperm contained in the cup to be splashed to neighboring stalks by falling water droplets.
Mosses can be either dioicous (compare dioecious in seed plants) or monoicous (compare monoecious). In dioicous mosses, male and female sex organs are borne on different gametophyte plants. In monoicous (also called autoicous) mosses, both are borne on the same plant. In the presence of water, sperm from the antheridia swim to the archegonia and fertilisation occurs, leading to the production of a diploid sporophyte. The sperm of mosses is biflagellate, i.e. they have two flagellae that aid in propulsion. Since the sperm must swim to the archegonium, fertilisation cannot occur without water. Some species (for example Mnium hornum or several species of Polytrichum) keep their antheridia in so called 'splash cups', bowl-like structures on the shoot tips that propel the sperm several decimeters when water droplets hit it, increasing the fertilization distance.[7]
After fertilisation, the immature sporophyte pushes its way out of the archegonial venter. It takes about a quarter to half a year for the sporophyte to mature. The sporophyte body comprises a long stalk, called a seta, and a capsule capped by a cap called the operculum. The capsule and operculum are in turn sheathed by a haploid calyptra which is the remains of the archegonial venter. The calyptra usually falls off when the capsule is mature. Within the capsule, spore-producing cells undergo meiosis to form haploid spores, upon which the cycle can start again. The mouth of the capsule is usually ringed by a set of teeth called peristome. This may be absent in some mosses.
Most mosses rely on the wind to disperse the spores. In the genus Sphagnum the spores are projected about 10–20 cm (4–8 in) off the ground by compressed air contained in the capsules; the spores are accelerated to about 36,000 times the earth's gravitational acceleration g.[8][9]
A patch of moss showing both gametophytes (the low, leaf-like forms) and sporophytes (the tall, stalk-like forms)
It has recently been found that microarthropods, such as springtails and mites, can effect moss fertilization[10] and that this process is mediated by moss-emitted scents. Male and female fire moss, for example emit different and complex volatile organic scents.[11] Female plants emit more compounds than male plants. Springtails were found to choose female plants preferentially, and one study found that springtails enhance moss fertilization, suggesting a scent-mediated relationship analogous to the plant-pollinator relationship found in many seed plants.[11] The stinkmoss species Splachnum sphaericum develops insect pollination further by attracting flies to its sporangia with a strong smell of carrion, and providing a strong visual cue in the form of red-coloured swollen collars beneath each spore capsule. Flies attracted to the moss carry its spores to fresh herbivore dung, which is the favoured habitat of the species of this genus.[12]
In many mosses, e.g. Ulota phyllantha, green vegetative structures called gemmae are produced on leaves or branches, which can break off and form new plants without the need to go through the cycle of fertilization. This is a means of asexual reproduction, and the genetically identical units can lead to the formation of clonal populations.
Dwarf males
Moss dwarf males (also known as nannandry or phyllodioicy) originate from wind-dispersed male spores that settle and germinate on the female shoot where their growth is restricted to a few millimeters. In some species, dwarfness is genetically determined, in that all male spores become dwarf.[13] More often, however, it is environmentally determined in that male spores that land on a female become dwarf, while those that land elsewhere develop into large, female-sized males.[13][14][15][16] In the latter case, dwarf males that are transplanted from females to another substrate develop into large shoots, suggesting that the females emit a substance which inhibits the growth of germinating males and possibly also quickens their onset of sexual maturation.[15][16] The nature of such a substance is unknown, but the phytohormone auxin may be involved[13]
Having the males growing as dwarfs on the female is expected to increase the fertilization efficiency by minimizing the distance between male and female reproductive organs. Accordingly, it has been observed that fertilization frequency is positively associated with the presence of dwarf males in several phyllodioicous species.[17][18]
Dwarf males occur in several unrelated lineages[18][19] and is showing to be more common than previously thought.[18] For example, it is estimated that between one quarter and half of all dioicous pleurocarps have dwarf males.[18]
DNA repair
The moss Phycomitrella patens has been used as a model organism to study how plants repair damage to their DNA, especially the repair mechanism known as homologous recombination. If the plant cannot repair DNA damage, e.g. double-strand breaks, in their somatic cells, the cells can lose normal functions or die. If this occurs during meiosis (part of sexual reproduction), they could become infertile. The genome of P. patens has been sequenced, which has allowed several genes involved in DNA repair to be identified.[20] P. patens mutants that are defective in key steps of homologous recombination have been used to work out how the repair mechanism functions in plants. For example, a study of P. patens mutants defective in RpRAD51, a gene that encodes a protein at the core of the recombinational repair reaction, indicated that homologous recombination is essential for repairing DNA double-strand breaks in this plant.[21] Similarly, studies of mutants defective in Ppmre11 or Pprad50 (that encode key proteins of the MRN complex, the principal sensor of DNA double-strand breaks) showed that these genes are necessary for repair of DNA damage as well as for normal growth and development.[22]
Classification
Traditionally, mosses were grouped with the liverworts and hornworts in the division Bryophyta (bryophytes, or Bryophyta sensu lato), within which the mosses made up the class Musci. However, this definition of Bryophyta was paraphyletic, as the mosses appear to be more closely related to vascular plants than to liverworts. Bryophytes are now split among three divisions: Bryophyta, Marchantiophyta and Anthocerotophyta. The current circumscription of Bryophyta includes only the mosses.
The mosses, now division Bryophyta (Bryophyta sensu stricto), are divided into eight classes:
division Bryophyta
class Takakiopsida
class Sphagnopsida
class Andreaeopsida
class Andreaeobryopsida
class Oedipodiopsida
class Polytrichopsida
class Tetraphidopsida
class Bryopsida
liverworts
hornworts
vascular plants
Bryophyta
Takakiopsida
Sphagnopsida
Andreaeopsida
Andreaeobryopsida
Neomusci
Oedipodiopsida
Cenomusci
Polytrichopsida
Altamusci
Tetraphidopsida
Bryopsida
The current phylogeny and composition of the Bryophyta.[2][23]
"Muscinae" from Ernst Haeckel's Kunstformen der Natur, 1904
Six of the eight classes contain only one or two genera each. Polytrichopsida includes 23 genera, and Bryopsida includes the majority of moss diversity with over 95% of moss species belonging to this class.
The Sphagnopsida, the peat-mosses, comprise the two living genera Ambuchanania and Sphagnum, as well as fossil taxa. However, the genus Sphagnum is a diverse, widespread, and economically important one. These large mosses form extensive acidic bogs in peat swamps. The leaves of Sphagnum have large dead cells alternating with living photosynthetic cells. The dead cells help to store water. Aside from this character, the unique branching, thallose (flat and expanded) protonema, and explosively rupturing sporangium place it apart from other mosses.
Andreaeopsida and Andreaeobryopsida are distinguished by the biseriate (two rows of cells) rhizoids, multiseriate (many rows of cells) protonema, and sporangium that splits along longitudinal lines. Most mosses have capsules that open at the top.
Polytrichopsida have leaves with sets of parallel lamellae, flaps of chloroplast-containing cells that look like the fins on a heat sink. These carry out photosynthesis and may help to conserve moisture by partially enclosing the gas exchange surfaces. The Polytrichopsida differ from other mosses in other details of their development and anatomy too, and can also become larger than most other mosses, with e.g. Polytrichum commune forming cushions up to 40 cm (16 in) high. The tallest land moss, a member of the Polytrichidae is probably Dawsonia superba, a native to New Zealand and other parts of Australasia.
Geological history
Red moss capsules, a winter native of the Yorkshire Dales moorland
The fossil record of moss is sparse, due to their soft-walled and fragile nature. Unambiguous moss fossils have been recovered from as early as the Permian of Antarctica and Russia, and a case is put forwards for Carboniferous mosses.[24] It has further been claimed that tube-like fossils from the Silurian are the macerated remains of moss calyptræ.[25] Mosses also appear to evolve 2–3 times slower than ferns, gymnosperms and angiosperms.[26]
Recent research show that ancient moss could explain why the Ordovician ice ages occurred. When the ancestors of today's moss started to spread on land 470 million years ago, they absorbed CO2 from the atmosphere and extracted minerals by secreting organic acids that dissolved the rocks they were growing on. These chemically altered rocks in turn reacted with the atmospheric CO2 and formed new carbonate rocks in the ocean through the weathering of calcium and magnesium ions from silicate rocks. The weathered rocks also released a lot of phosphorus and iron which ended up in the oceans, where it caused massive algal blooms, resulting in organic carbon burial, extracting more carbon dioxide from the atmosphere. Small organisms feeding on the nutrients created large areas without oxygen, which caused a mass extinction of marine species, while the levels of CO2 dropped all over the world, allowing the formation of ice caps on the poles.[27][28]
Ecology
Habitat
Dense moss colonies in a cool coastal forest
Moss with sporophytes on brick
A closeup of moss on a rock
Young sporophytes of the common moss Tortula muralis (wall screw-moss)
Retaining wall covered in moss
A small clump of moss.
The moss garden at the Bloedel Reserve, Bainbridge Island, Washington State.
Since moss gametophytes are autotrophic they require enough sunlight to perform photosynthesis.[29] Shade tolerance varies by species, just as it does with higher plants. In most areas, mosses grow chiefly in areas of dampness and shade, such as wooded areas and at the edges of streams; but they can grow anywhere in cool damp cloudy climates, and some species are adapted to sunny, seasonally dry areas like alpine rocks or stabilized sand dunes.
Choice of substrate varies by species as well. Moss species can be classed as growing on: rocks, exposed mineral soil, disturbed soils, acid soil, calcareous soil, cliff seeps and waterfall spray areas, streamsides, shaded humusy soil, downed logs, burnt stumps, tree trunk bases, upper tree trunks, and tree branches or in bogs. Moss species growing on or under trees are often specific about the species of trees they grow on, such as preferring conifers to broadleaf trees, oaks to alders, or vice versa.[6] While mosses often grow on trees as epiphytes, they are never parasitic on the tree.
Mosses are also found in cracks between paving stones in damp city streets, and on roofs. Some species adapted to disturbed, sunny areas are well adapted to urban conditions and are commonly found in cities. Examples would be Rhytidiadelphus squarrosus, a garden weed in Vancouver and Seattle areas; Bryum argenteum, the cosmopolitan sidewalk moss, and Ceratodon purpureus, red roof moss, another cosmopolitan species. A few species are wholly aquatic, such as Fontinalis antipyretica, common water moss; and others such as Sphagnum inhabit bogs, marshes and very slow-moving waterways.[6] Such aquatic or semi-aquatic mosses can greatly exceed the normal range of lengths seen in terrestrial mosses. Individual plants 20–30 cm (8–12 in) or more long are common in Sphagnum species for example.
Wherever they occur, mosses require liquid water for at least part of the year to complete fertilisation. Many mosses can survive desiccation, sometimes for months, returning to life within a few hours of rehydration.[29]
It is generally believed that in northern latitudes, the north side of trees and rocks will generally have more luxuriant moss growth on average than other sides.[30] This is assumed to be because the sun on the south side creates a dry environment. South of the equator the reverse would be true. However, naturalists feel that mosses grow on the damper side of trees and rocks.[5] In some cases, such as sunny climates in temperate northern latitudes, this will be the shaded north side of the tree or rock. On steep slopes it may be the uphill side. For mosses that grow on tree branches, this is generally the upper side of the branch on horizontally growing sections or near the crotch. In cool damp cloudy climates, all sides of tree trunks and rocks may be equally damp enough for mosses. And different species of mosses have different moisture and sun requirements so will grow on different sections of the same tree or rock.
Moss-cyanobacteria relationship
In boreal forests, some species of moss play an important role in providing nitrogen for the ecosystem due to their relationship with nitrogen-fixing cyanobacteria. Cyanobacteria colonizes moss and receives shelter in return for providing fixed nitrogen. Moss releases the fixed nitrogen, along with other nutrients, into the soil "upon disturbances like drying-rewetting and fire events," making it available throughout the ecosystem.[31]
Cultivation
Moss is often considered a weed in grass lawns, but is deliberately encouraged to grow under aesthetic principles exemplified by Japanese gardening. In old temple gardens, moss can carpet a forest scene. Moss is thought to add a sense of calm, age, and stillness to a garden scene. Moss is also used in bonsai to cover the soil and enhance the impression of age.[32] Rules of cultivation are not widely established. Moss collections are quite often begun using samples transplanted from the wild in a water-retaining bag. However, specific species of moss can be extremely difficult to maintain away from their natural sites with their unique requirements of combinations of light, humidity, substrate chemistry, shelter from wind, etc.
Growing moss from spores is even less controlled. Moss spores fall in a constant rain on exposed surfaces; those surfaces which are hospitable to a certain species of moss will typically be colonised by that moss within a few years of exposure to wind and rain. Materials which are porous and moisture retentive, such as brick, wood, and certain coarse concrete mixtures, are hospitable to moss. Surfaces can also be prepared with acidic substances, including buttermilk, yogurt, urine, and gently puréed mixtures of moss samples, water and ericaceous compost.
In the cool cloudy damp Pacific Northwest, moss is sometimes allowed to grow naturally as a lawn substitute, one that needs little or no mowing, fertilizing or watering. In this case, grass is considered to be the weed.[33] Landscapers in the Seattle area sometimes collect boulders and downed logs growing mosses for installation in gardens and landscapes. Woodland gardens in many parts of the world can include a carpet of natural mosses.[29] The Bloedel Reserve on Bainbridge Island, Washington State, is famous for its moss garden. The moss garden was created by removing shrubby underbrush and herbaceous groundcovers, thinning trees, and allowing mosses to fill in naturally.[34]
Mosses are sometimes used in green roofs. Advantages of mosses over higher plants in green roofs include reduced weight loads, increased water absorption, no fertilizer requirements, and high drought tolerance. Since mosses do not have true roots, they require less planting medium than higher plants with extensive root systems. With proper species selection for the local climate, mosses in green roofs require no irrigation once established and are low maintenance.[35]
Inhibiting moss growth
Moss can be a troublesome weed in containerized nursery operations and greenhouses.[36] Vigorous moss growth can inhibit seedling emergence and penetration of water and fertilizer to the plant roots.
Moss growth can be inhibited by a number of methods:
Decreasing availability of water through drainage.
Increasing direct sunlight.
Increasing number and resources available for competitive plants like grasses.
Increasing the soil pH with the application of lime.
Heavy traffic or manually disturbing the moss bed with a rake
Application of chemicals such as ferrous sulfate (e.g. in lawns) or bleach (e.g. on solid surfaces).
In containerized nursery operations, coarse mineral materials such as sand, gravel, and rock chips are used as a fast-draining top dressing in plant containers to discourage moss growth.
The application of products containing ferrous sulfate or ferrous ammonium sulfate will kill moss; these ingredients are typically in commercial moss control products and fertilizers. Sulfur and Iron are essential nutrients for some competing plants like grasses. Killing moss will not prevent regrowth unless conditions favorable to their growth are changed.[37]
Mossery
A passing fad for moss-collecting in the late 19th century led to the establishment of mosseries in many British and American gardens. The mossery is typically constructed out of slatted wood, with a flat roof, open to the north side (maintaining shade). Samples of moss were installed in the cracks between wood slats. The whole mossery would then be regularly moistened to maintain growth.
Uses
Traditional
Preindustrial societies made use of the mosses growing in their areas.
Laplanders, North American tribes, and other circumpolar people used mosses for bedding.[5][29] Mosses have also been used as insulation both for dwellings and in clothing. Traditionally, dried moss was used in some Nordic countries and Russia as an insulator between logs in log cabins, and tribes of the northeastern United States and southeastern Canada used moss to fill chinks in wooden longhouses.[29] Circumpolar and alpine peoples have used mosses for insulation in boots and mittens. Ötzi the Iceman had moss-packed boots.[29]
The capacity dried mosses have to absorb fluids, has made their use practical in both medical and culinary uses. North American tribal people used mosses for diapers, wound dressing, and menstrual fluid absorption.[29] Tribes of the Pacific Northwest in the United States and Canada used mosses to clean salmon prior to drying, and packed wet moss into pit ovens for steaming camas bulbs. Food storage baskets and boiling baskets were also packed with mosses.[29]
Commercial
There is a substantial market in mosses gathered from the wild. The uses for intact moss are principally in the florist trade and for home decoration. Decaying moss in the genus Sphagnum is also the major component of peat, which is "mined" for use as a fuel, as a horticultural soil additive, and in smoking malt in the production of Scotch whisky.
Sphagnum moss, generally the species S. cristatum and S. subnitens, is harvested while still growing and is dried out to be used in nurseries and horticulture as a plant growing medium.
The practice of harvesting peat moss should not be confused with the harvesting of moss peat. Peat moss can be harvested on a sustainable basis and managed so that regrowth is allowed, whereas the harvesting of moss peat is generally considered to cause significant environmental damage as the peat is stripped with little or no chance of recovery.[citation needed]
Some Sphagnum mosses can absorb up to 20 times their own weight in water.[38] In World War I, Sphagnum mosses were used as first-aid dressings on soldiers' wounds, as these mosses said to absorb liquids three times faster than cotton, retain liquids better, better distribute liquids uniformly throughout themselves, and are cooler, softer, and be less irritating.[38] It is also claimed to have antibacterial properties.[39] Native Americans were one of the peoples to use Sphagnum for diapers and napkins, which is still done in Canada.[40]
In rural UK, Fontinalis antipyretica was traditionally used to extinguish fires as it could be found in substantial quantities in slow-moving rivers and the moss retained large volumes of water which helped extinguish the flames. This historical use is reflected in its specific Latin/Greek name, the approximate meaning of which is "against fire".
In Finland, peat mosses have been used to make bread during famines.[41] In Mexico, Moss is used as a Christmas decoration.
Moss bioreactor cultivating the moss Physcomitrella patens
Physcomitrella patens is increasingly used in biotechnology. Prominent examples are the identification of moss genes with implications for crop improvement or human health[42] and the safe production of complex biopharmaceuticals in the moss bioreactor, developed by Ralf Reski and his co-workers.[43]
See also
Mosses portal
Embryophyte
Lichen
References
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^ Jump up to: a b c d Hedenäs, Lars; Bisang, Irene (2011). "The overlooked dwarf males in mosses—unique among green land plants". Perspectives in Plant Ecology, Evolution and Systematics. 13 (2): 121–135. doi:10.1016/j.ppees.2011.03.001.
Jump up ^ Ramsay, Helen P.; Berrie, G. K. (1982). "Sex determination in bryophytes". Journal of the Hattori Botanical Laboratory. 52: 255–274.
Jump up ^ Rensing SA, Lang D, Zimmer AD, Terry A, Salamov A, Shapiro H, Nishiyama T, Perroud PF, Lindquist EA, Kamisugi Y, Tanahashi T, Sakakibara K, Fujita T, Oishi K, Shin-I T, Kuroki Y, Toyoda A, Suzuki Y, Hashimoto S, Yamaguchi K, Sugano S, Kohara Y, Fujiyama A, Anterola A, Aoki S, Ashton N, Barbazuk WB, Barker E, Bennetzen JL, Blankenship R, Cho SH, Dutcher SK, Estelle M, Fawcett JA, Gundlach H, Hanada K, Heyl A, Hicks KA, Hughes J, Lohr M, Mayer K, Melkozernov A, Murata T, Nelson DR, Pils B, Prigge M, Reiss B, Renner T, Rombauts S, Rushton PJ, Sanderfoot A, Schween G, Shiu SH, Stueber K, Theodoulou FL, Tu H, Van de Peer Y, Verrier PJ, Waters E, Wood A, Yang L, Cove D, Cuming AC, Hasebe M, Lucas S, Mishler BD, Reski R, Grigoriev IV, Quatrano RS, Boore JL (January 2008). "The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants" (PDF). Science. 319 (5859): 64–9. doi:10.1126/science.1150646. PMID 18079367.
Jump up ^ Markmann-Mulisch U, Wendeler E, Zobell O, Schween G, Steinbiss HH, Reiss B (October 2007). "Differential requirements for RAD51 in Physcomitrella patens and Arabidopsis thaliana development and DNA damage repair". Plant Cell. 19 (10): 3080–9. doi:10.1105/tpc.107.054049. PMC 2174717. PMID 17921313.
Jump up ^ Kamisugi Y, Schaefer DG, Kozak J, Charlot F, Vrielynck N, Holá M, Angelis KJ, Cuming AC, Nogué F (April 2012). "MRE11 and RAD50, but not NBS1, are essential for gene targeting in the moss Physcomitrella patens". Nucleic Acids Res. 40 (8): 3496–510. doi:10.1093/nar/gkr1272. PMC 3333855. PMID 22210882.
Jump up ^ Buck, William R. & Bernard Goffinet. (2000). "Morphology and classification of mosses", pages 71-123 in A. Jonathan Shaw & Bernard Goffinet (Eds.), Bryophyte Biology. (Cambridge: Cambridge University Press). ISBN 0-521-66097-1.
Jump up ^ Thomas, B.A. (1972). "A probable moss from the Lower Carboniferous of the Forest of Dean, Gloucestershire". Annals of Botany. 36 (1): 155–161. ISSN 1095-8290. JSTOR 42752024.
Jump up ^ Kodner, R. B.; Graham, L. E. (2001). "High-temperature, acid-hydrolyzed remains of Polytrichum (Musci, Polytrichaceae) resemble enigmatic Silurian-Devonian tubular microfossils". American Journal of Botany. 88 (3): 462–466. doi:10.2307/2657111. JSTOR 2657111. PMID 11250824. Archived from the original on 2009-03-03.
Jump up ^ Stenøien, H. K. (2008). "Slow molecular evolution in 18S rDNA, rbcL and nad5 genes of mosses compared with higher plants". Journal of Evolutionary Biology. 21: 566–571. doi:10.1111/j.1420-9101.2007.01479.x. PMID 18205784. Archived from the original on 2013-06-03. Retrieved 2013-09-11.
Jump up ^ "First land plants plunged Earth into ice age". Newscientist.com. Archived from the original on 2013-09-23. Retrieved 2013-09-11.
Jump up ^ "First Plants Caused Ice Ages, New Research Reveals". Sciencedaily.com. 2012-02-01. Archived from the original on 2013-10-03. Retrieved 2013-09-11.
^ Jump up to: a b c d e f g h Kimmerer, Robin Wall (2003). Gathering Moss. Corvallis, Oregon: Oregon State University Press. ISBN 0-87071-499-6.
Jump up ^ Porley, Ron; Hodgetts, Nick (2005). Mosses & Liverworts. London: Collins. pp. 80–81. ISBN 0-00-220212-3.
Jump up ^ Rousk, Kathrin; Jones, Davey L.; DeLuca, Thomas H. (2013-01-01). "Moss-cyanobacteria associations as biogenic sources of nitrogen in boreal forest ecosystems". Frontiers in Microbiology. 4. doi:10.3389/fmicb.2013.00150. ISSN 1664-302X. PMC 3683619. PMID 23785359.
Jump up ^ Chan, Peter (1993). Bonsai Masterclass. New York City: Sterling Publishing Co. ISBN 0-8069-6763-3.
Jump up ^ Smith, Sally W. (1998). Sunset Western Garden Problem Solver. Menlo Park, California: Sunset Books. ISBN 0-376-06132-4.
Jump up ^ "The Bloedel Reserve". Archived from the original on 16 April 2011. Retrieved 24 April 2011.
Jump up ^ "RoofTopGarden". Archived from the original on 24 April 2011. Retrieved 22 May 2011.
Jump up ^ Haglund, William A.; Russell and Holland (Summer 1981). "Moss Control in Container-Grown Conifer Seedlings" (PDF). Tree Planter's Notes(USFS). 32 (3): 27–29. Archived (PDF) from the original on 19 July 2011. Retrieved 24 April 2011.
Jump up ^ Steve Whitcher; Master Gardener (1996). "Moss Control in Lawns". Gardening in Western Washington. Washington State University. Archived from the original on 2007-02-05. Retrieved 2007-02-10.
^ Jump up to: a b The Plant Underworld, Sphagnum and Water, Australian Botanic Garden Archived 2014-02-17 at the Wayback Machine.
Jump up ^ Stalheim, T.; Ballance, S.; Christensen, B. E.; Granum, P. E. (2009-03-01). "Sphagnan – a pectin-like polymer isolated from Sphagnum moss can inhibit the growth of some typical food spoilage and food poisoning bacteria by lowering the pH". Journal of Applied Microbiology. 106 (3): 967–976. doi:10.1111/j.1365-2672.2008.04057.x. ISSN 1365-2672.
Jump up ^ Hotson, J. W. (1921). "Sphagnum Used as Surgical Dressing in Germany during the World War (Concluded)". The Bryologist. 24 (6): 89–96. doi:10.1639/0007-2745(1921)24[89:suasdi]2.0.co;2. JSTOR 3237483.
Jump up ^ Engman, Max; D. G. Kirby (1989). Finland: people, nation, state. C. Hurst & Co. p. 45. ISBN 0-253-32067-4.
Jump up ^ Ralf Reski and Wolfgang Frank (2005): Moss (Physcomitrella patens) functional genomics – Gene discovery and tool development with implications for crop plants and human health. Briefings in Functional Genomics and Proteomics 4, 48-57.
Jump up ^ Decker, E. L.; Reski, R. (2007). "Moss bioreactors producing improved biopharmaceuticals". Current Opinion in Biotechnology. 18 (5): 393–398. doi:10.1016/j.copbio.2007.07.012. PMID 17869503.
External links
Wikimedia Commons has media related to Bryophyta.
Wikispecies has information related to Bryophyta
The dictionary definition of moss at Wiktionary
Information, diagrams and photos
Moss grower's handbook- 2.39MB, PDF file
The British Bryological Society
Picture Gallery of Mosses
World of Mosses - Watercolour paintings of moss by Robert Muma
show
v
t
e
Classification of Archaeplastida / Plantae sensu lato
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Rhodophyta
(red algae)
Cyanidiophyceae
Porphyridiophyceae
Compsopogonophyceae
Stylonematophyceae
Rhodellophyceae
Bangiophyceae
Florideophyceae
Glaucocystophyta
(glaucophytes)
Glaucocystophyceae
Glaucocystis
Cyanophora
Gloeochaete
Viridiplantae
(green algae,
& land plants)
M+C+S
Mesostigmatophyceae
Chlorokybophyceae
Spirotaenia
C+S
Chlorophyta
(prasinophytes)
Mamiellophyceae
Nephrophyceae
Palmophyllales
Prasinophyceae
Pseudoscourfieldiales
Pyramimonadophyceae
Scourfieldiales
Pedinophyceae
Chlorodendrophyceae
UTC clade
Ulvophyceae
Trebouxiophyceae
Chlorophyceae
Streptophyta
(charophytes,
& land plants)
Charophytes
(basal)
Streptofilum
Klebsormidiophyceae
Phragmo-
plastophyta
Charophytes
(non-basal)
Charophyceae
Coleochaetophyceae
Zygnematophyceae
Mesotaeniaceae
Embryophyta
(land plants)
Bryophytes
(non-vascular)
Marchantiophyta
Anthocerotophyta
Bryophyta "Moss"
†Horneophytopsida
Tracheophyta
(vascular)
Lycopodiophyta
(microphylls)
†Zosterophyllopsida
†Sawdoniales
Isoetopsida
Lycopodiopsida
Euphyllophyta
(megaphylls)
Moniliformopses
(ferns)
†Cladoxylopsida
†Stauropteridales
†Zygopteridales
Equisetopsida
Psilotopsida
Marattiopsida
Polypodiopsida
Spermatophyta
(seed plants)
†Seed ferns
Gymnosperms
Gnetopsida
Pinopsida
Cycadopsida
Ginkgoopsida
Angiosperms or flowering plants
Amborellales
Nymphaeales
Austrobaileyales
Magnoliids
Monocots
Eudicots
Other
†Trimerophytopsida
†Progymnosperm
Other
†Rhyniopsida
† = extinct. See also the list of plant orders.
show
v
t
e
Classification of Bryophyta
Domain
Archaea
Bacteria
Eukaryota
(Supergroup
Plant
Hacrobia
Heterokont
Alveolata
Rhizaria
Excavata
Amoebozoa
Opisthokonta
Animal
Fungi)
Sphagnopsida
†Protosphagnales
Ambuchananiales
Sphagnales
Takakiopsida
Takakiales
Andreaeopsida
Andreaeales
Andreaeobryopsida
Andreaeobryales
Tetraphidopsida
Tetraphidales
Oedipodiopsida
Oedipodiales
Polytrichopsida
Polytrichales
Bryopsida
Buxbaumiidae
Buxbaumiales
Diphysciidae
Diphysciales
Funariidae
Gigaspermales
Encalyptales
Funariales
Timmiidae
Timmiales
Dicranidae
Archidiales
Scouleriales
Grimmiales
Bryoxiphiales
Pottiales
Dicranales
Bryidae
Bryanae
Bartramiales
Hedwigiales
Splachnales
Bryales
Orthotrichales
Orthodontiales
Rhizogoniales
Aulacomniales
Hypnanae
Hypnodendrales
Ptychomniales
Hookeriales
Hypnales

Taxon identifiers
Wikidata: Q25347
Wikispecies: Bryophyta
EoL: 3768
EPPO: 1BRYP
Fossilworks: 55135
GBIF: 35
ITIS: 14189
NCBI: 3208
NZOR: 8c5e5c2b-067d-4b78-ae2b-65a94ce407e6
WoRMS: 147480
Authority control
GND: 4166892-3
LCCN: sh85087489
NDL: 00570784
Retrieved from "https://en.wikipedia.org/w/index.php?title=Moss&oldid=866375799"
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text:
National Basketball Association on television
From Wikipedia, the free encyclopedia
Jump to navigation Jump to search
The National Basketball Association is shown on national television on broadcast channel ABC, cable networks ESPN and TNT, as well as the NBA TV network. The NBA is also shown on multiple regional sports networks.
Contents
[hide]
1 History
2 Regular season
3 NBA playoffs
4 NBA Finals
5 Single games
5.1 NBA on Christmas Day
5.2 NBA All-Star Game
6 Regional broadcasters
7 See also
8 References
9 External links
History[edit]
As one of the major sports leagues in North America, the National Basketball Association has a long history of partnership with television networks in the US. The League signed a contract with DuMont in its 8th season (1953–54), marking the first year the NBA had a national television broadcaster. Similar to NFL, the lack of television stations leads to NBC taking over the rights beginning the very next season until April 7, 1962 - NBC's first tenure with the NBA. After the deal expired, Sports Network Incorporated (later known as the Hughes Television Network) signed up for two-year coverage in the 1962–63, 1963–64 season. ABC gained the NBA in 1964, the network aired its first NBA game on January 3, 1965, but lost the broadcast rights to CBS after the 1972–73 season with the initial tenure ending on May 10, 1973. As the national broadcaster of the NBA, CBS aired NBA games from the 1973-74 until the 1989–90 season, during which the early 1980s is notoriously known as the tape delay playoff era.
NBC then succeeded the broadcast rights from 1990 to 2002.[1] During NBC's partnership with the NBA in the 1990s, the league rose to unprecedented popularity, with ratings surpassing the days of Magic Johnson and Larry Bird in the mid 1980s. Upon expiration of the contract in 2002, the league signed an agreement with ABC, which began airing games in the 2002-03 season. NBC had made a four-year $1.3 billion ($330 million/year) bid in the spring of 2002 to renew its NBA rights, but the league instead went to ESPN and ABC with a six-year deal worth $2.4 billion ($400 million/year), a total of $4.6 billion ($766 million/year) when adding the cable deal with Turner Sports.[2] Partially due to the retirement of Michael Jordan, the league suffered ratings decline after ESPN and ABC took over the rights. NBA extended its national TV package on June 27, 2007 worth eight-year $7.4 billion ($930 million/year) through 2015–16 season,[2] during which the league had its new resurgence leading by renewed Celtics–Lakers rivalry and LeBron James. On October 6, 2014, NBA announced a nine-year $24 billion ($2.7 billion/year) extension with ESPN, ABC and Turner Sports beginning with the 2016–17 season and running through the 2024–25 season[3] - the second most expensive media rights in the world after NFL and on a par with Premier League in annual rights fee from 2016–17 to 2018–19 season.[4]
NBA entered the cable territory in 1979 when USA Network signed a three-year $1.5 million deal and extended for two years until the 1983-84 season, ESPN also had a brief affair with NBA from 1982 to 1984. Turner Sports obtained rights to air NBA games beginning with the 1984-85 season (replacing ESPN and USA Network as national cable partners) under a four-year deal, in which TBS shared the NBA television package along with CBS. In the summer of 1987, Turner Broadcasting System signed a new joint broadcast contract between TBS and TNT to split broadcast NBA games starting from the 1988-89 season. TNT held rights to broadcast the NBA draft, most NBA regular season and playoff games, while TBS only aired single games or doubleheaders once a week. The 2001-02 season would mark the final year of regular NBA coverage on TBS, Turner Sports signed a new NBA television contract in which TNT would assume rights to the company's NBA package while TBS would discontinue game coverage altogether. In recent years however, TBS has served as an overflow feed during the playoffs while also simulcasting the 2015, 2016, and 2017 NBA All-Star Game. Subsequently ESPN regained the NBA in 2002-03 season and took over TBS's half of cable television rights.
Regular season[edit]
Average rating per regular season game on broadcast networks
(1995–present)
Season
Network
Rating
2017–18 ABC
2.2
2016–17 ABC
1.9
2015–16 ABC
2.3
2014–15 ABC
2.2
2013–14 ABC
2.3
2012–13 ABC
2.9
2011–12 (lockout shortened) ABC
3.3
2010–11 ABC
3.0
2009–10 ABC
2.3
2008–09 ABC
2.3
2007–08 ABC
2.2
2006–07 ABC
2.0
2005–06 ABC
2.2
2004–05 ABC
2.3
2003–04 ABC
2.4
2002–03 ABC
2.6
2001–02 NBC
2.9
2000–01 NBC
3.0
1999–2000 NBC
3.3
1998–99 (lockout shortened) NBC
4.3
1997–98 NBC
4.6
1996–97 NBC
4.7
1995–96 NBC
5.0
Average viewership per regular season game in millions by networks
(2007–present)
Season
ABC
TNT
ESPN
NBA TV
2017–18
3.82M
1.74M
1.63M
0.31M
2016–17
3.27M
1.54M
1.57M
0.31M
2015–16
3.93M
1.68M
1.65M
0.35M
2014–15
3.59M
1.67M
1.50M
0.29M
2013–14
3.58M
1.90M
1.68M
0.32M
2012–13
4.70M
2.00M
1.77M
0.34M
2011–12 (lockout shortened)
5.42M
2.50M
1.86M
0.34M
2010–11
5.11M
2.40M
1.99M
0.25M
2009–10
3.69M
1.72M
1.56M
---
2008–09
3.68M
1.71M
1.68M
---
2007–08
3.18M
1.47M
1.47M
---
NBA playoffs[edit]
Main article: NBA playoffs
Ever since NBC lost the rights, the Conference Finals have been on cable networks almost exclusively with TNT and ESPN alternating coverage each year.
Most watched NBA games on cable networks
Year
Network
Game
Rating
Viewership
2016 TNT Oklahoma City Thunder vs Golden State Warriors WCF Game 7
8.9
16.00M
2012 ESPN Boston Celtics vs Miami Heat ECF Game 7
7.7
13.35M
2013 TNT Indiana Pacers vs Miami Heat ECF Game 7
7.1
11.57M
2011 TNT Miami Heat vs Chicago Bulls ECF Game 1
6.2
11.11M
2012 ESPN Miami Heat vs Boston Celtics ECF Game 6
6.8
11.07M
2012 ESPN Miami Heat vs Boston Celtics ECF Game 4
6.8
11.07M
2011 TNT Chicago Bulls vs Miami Heat ECF Game 3
6.4
10.89M
2016 TNT Golden State Warriors vs Oklahoma City Thunder WCF Game 6
6.2
10.81M
2011 TNT Miami Heat vs Chicago Bulls ECF Game 5
6.4
10.41M
2012 ESPN Boston Celtics vs Miami Heat ECF Game 5
6.3
10.25M
NBA Finals[edit]
Main articles: NBA Finals television ratings and List of NBA Finals broadcasters
Most watched NBA Final games on ABC
Year
Game
Rating/Share
Viewership
2016 Cleveland Cavaliers vs Golden State Warriors Game 7
15.8/29
31.02M
2010 Boston Celtics vs Los Angeles Lakers Game 7
15.6/27
28.20M
2013 San Antonio Spurs vs Miami Heat Game 7
15.3/26
26.32M
2017 Cleveland Cavaliers vs Golden State Warriors Game 5
13.5/25
24.47M
2011 Dallas Mavericks vs Miami Heat Game 6
13.3/23
23.88M
2015 Golden State Warriors vs Cleveland Cavaliers Game 6
13.4/24
23.25M
2004 Los Angeles Lakers vs Detroit Pistons Game 5
13.8/23
21.84M
2015 Cleveland Cavaliers vs Golden State Warriors Game 5
11.8/21
20.86M
2016 Golden State Warriors vs Cleveland Cavaliers Game 6
11.8/22
20.70M
2013 San Antonio Spurs vs Miami Heat Game 6
12.3/21
20.64M
Single games[edit]
NBA on Christmas Day[edit]
Main article: NBA on Christmas Day
Games on Christmas Day have drawn some of the biggest regular season audience. Since 2001, the most watched Christmas games were:
2004 Miami Heat vs Los Angeles Lakers on ABC averaged a 7.3 rating and 13.18 million viewers.
2010 Miami Heat vs Los Angeles Lakers on ABC averaged a 6.4 rating and 13.11 million viewers.
2015 Cleveland Cavaliers vs Golden State Warriors on ABC averaged a 5.7 rating and 11.12 million viewers.
Average rating/viewership per game for NBA on Christmas Day
(2001–present)
Year
Network
Games
Rating
Viewership
2017 ABC, ESPN, TNT
5
2.6
5.10M
2016 ABC, ESPN
5
2.3
4.56M
2015 ABC, ESPN
5
3.0
5.55M
2014 ABC, ESPN, TNT
5
2.8
5.22M
2013 ABC, ESPN
5
2.5
4.46M
2012 ABC, ESPN
5
3.1
5.50M
2011 (lockout shortened, opening day) ABC, ESPN, TNT
5
3.5
6.31M
2010 ABC, ESPN
5
3.2
6.00M
2009 ABC, ESPN
5
2.4
4.17M
2008 ABC, ESPN, TNT
5
2.5
4.43M
2007 ABC, ESPN
3
2.6
4.29M
2006 ABC
1
3.5
5.47M
2005 ABC
2
4.4
7.12M
2004 ABC, ESPN
2
5.2
8.92M
2003 ABC, ESPN
3
3.0
4.96M
2002 ABC, ESPN
3
2.8
4.52M
2001 NBC
2
3.2
4.99M
NBA All-Star Game[edit]
Main article: List of NBA All-Star Game broadcasters
The NBA All-Star Game was on broadcast networks until 2002, TNT began airing the All-Star Game in 2003 which featured the last appearance of Michael Jordan in the event, TBS started simulcasting the game since 2015.
NBA All-Star Game TV ratings
(1990–present)
Year
Network
Results
Rating/Share
Viewership
2017 TNT, TBS West 192, East 182
4.2/7
7.75M
2016 TNT, TBS West 196, East 173
4.3/7
7.61M
2015 TNT, TBS West 163, East 158
4.3/7
7.18M
2014 TNT East 163, West 155
4.3/7
7.51M
2013 TNT West 143, East 138
4.6/8
8.02M
2012 TNT West 152, East 149
4.4/7
7.07M
2011 TNT West 148, East 143
5.2/9
9.09M
2010 TNT East 141, West 139
3.8/6
6.85M
2009 TNT West 146, East 119
4.5/7
7.62M
2008 TNT East 134, West 128
3.8/6
6.33M
2007 TNT West 153, East 132
4.2/7
6.84M
2006 TNT East 122, West 120
4.3/8
7.07M
2005 TNT East 125, West 115
4.9/8
8.08M
2004 TNT West 136, East 132
5.1/10
8.19M
2003 TNT West 155, East 145
6.6/12
10.83M
2002 NBC West 135, East 120
8.2/15
13.10M
2001 NBC East 111, West 110
5.1/8
7.76M
2000 NBC East 137, West 126
6.9/12
10.52M
1999
Cancelled due to owners' Lockout
1998 NBC East 135, West 114
10.6/17
16.93M
1997 NBC East 132, West 120
11.2/19
16.90M
1996 NBC East 129, West 118
11.7/20
17.46M
1995 NBC West 139, East 112
10.7/17
15.78M
1994 NBC East 127, West 118
9.1/14
13.67M
1993 NBC West 135, East 132
14.3/22
22.91M
1992 NBC West 153, East 113
12.8/26
18.83M
1991 NBC East 116, West 113
7.8/21
10.61M
1990 CBS East 130, West 113
9.5/13
13.20M
Regional broadcasters[edit]
Main article: List of current National Basketball Association broadcasters
NBA games not televised by its national partners are instead broadcast by local broadcast stations and regional sports network, televising their respective local team within their respective region. Regionally broadcast games are subject to blackouts, and those games from outside of a viewer's designated market are blacked out to protect the local team.
Most NBA regional broadcasters are members of national chains:
Regional network
Entering the 2017–18 NBA season
Teams(s)
Altitude Sports Denver
AT&T SportsNet Houston, Utah
Fox Sports Networks
(incl. Yes Network) Atlanta, Brooklyn, Charlotte, Cleveland, Detroit, Dallas, Indiana, LA Clippers, Memphis, Miami, Milwaukee, Minnesota, New Orleans, Oklahoma City, Orlando, Phoenix, San Antonio
NBC Sports Regional Networks Boston, Chicago, Golden State, Philadelphia, Portland, Sacramento, Washington
MSG New York
Spectrum Sports LA Lakers
Sportsnet and TSN
(shared rights) Toronto
See also[edit]
NBA TV
NBA TV Canada
NBA League Pass
List of current National Basketball Association broadcasters
Major League Baseball on television
Major League Soccer on television
National Football League on television
National Hockey League on television
References[edit]
Jump up ^ "NBC Celebrates 12 Years of NBA on NBC". NBC Sports History Page.
^ Jump up to: a b John Lombardo & John Ourand (October 13, 2014). "Fast break: NBA media rights". SportsBusiness Daily. Retrieved June 17, 2016.
Jump up ^ Paulsen (October 6, 2014). "NBA Announces 9-Year Extension With ESPN, Turner, Through 2025". Sports Media Watch. Retrieved June 17, 2016.
Jump up ^ BBC Sports (February 10, 2015). "Premier League TV rights: Sky and BT pay £5.1bn for live games". BBC Sport. Retrieved June 17, 2016.
External links[edit]
NBA on Christmas Day numbers game
NBA All-Star Game TV rating and viewership
Most watched NBA games on cable networks
Most watched NBA games since 1998
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