Moss

Last updated

Moss
Temporal range: Carboniferous [1] present
Tionesta-ac-moss2.jpg
Clumps of moss on the ground and base of trees in the Allegheny National Forest, Pennsylvania, United States
Scientific classification Red Pencil Icon.png
Kingdom: Plantae
Clade: Embryophytes
Division: Bryophyta
Schimp. sensu stricto
Classes [2]
Synonyms
  • Musci L.
  • Muscineae Bisch.

Mosses, the taxonomic division Bryophyta ( /brˈɒfɪtə/ , [3] /ˈbr.ftə/ ) or the bryophytes ( /ˈbrəfts/ ), [4] are small, non-vascular flowerless plants that typically form 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. [5] 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.

Contents

Mosses are commonly confused with hornworts, liverworts and lichens. [6] Mosses were formerly grouped with the hornworts and liverworts as "non-vascular" plants in the 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. Lichens may superficially resemble mosses, and sometimes have common names that include the word "moss" (e.g., "reindeer moss" or "Iceland moss"), but they are not related to mosses. [6] :3 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.

Physical characteristics

Description

Chloroplasts and accumulated starch granules in Bryum capillare Bryum capillare leaf cells.jpg
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. [7] [8] 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.[ citation needed ] 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. [8] Other differences are not universal for all mosses and all liverworts, but the presence of a clearly differentiated stem with simple-shaped, non-vascular leaves that are 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) Lifecycle moss svg diagram.svg
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. [9]

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. [10] [11]

A patch of moss showing both gametophytes (the low, leaf-like forms) and sporophytes (the tall, stalk-like forms) Moss Gametophytes Sporophytes.JPG
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 [12] and that this process is mediated by moss-emitted scents. Male and female fire moss, for example emit different and complex volatile organic scents. [13] 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. [13] 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. [14]

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. [15] 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. [15] [16] [17] [18] 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. [17] [18] The nature of such a substance is unknown, but the phytohormone auxin may be involved [15]

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. [19] [20]

Dwarf males occur in several unrelated lineages [20] [21] and is showing to be more common than previously thought. [20] For example, it is estimated that between one quarter and half of all dioicous pleurocarps have dwarf males. [20]

DNA repair

The moss Physcomitrella 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. [22] 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. [23] 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. [24]

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. The old Bryophyte division is now split into three divisions: Bryophyta (mosses), Marchantiophyta (liverworts) and Anthocerotophyta (hornworts). 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] [25]
"Muscinae" from Ernst Haeckel's Kunstformen der Natur, 1904 Haeckel Muscinae.jpg
"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

Bristly Haircap moss, a winter native of the Yorkshire Dales moorland RedMoss.jpg
Bristly Haircap moss, 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. [26] It has further been claimed that tube-like fossils from the Silurian are the macerated remains of moss calyptræ. [27] Mosses also appear to evolve 2–3 times slower than ferns, gymnosperms and angiosperms. [28]

Recent research shows 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. [29] [30]

Ecology

Habitat

Since moss gametophytes are autotrophic they require enough sunlight to perform photosynthesis. [31] Shade tolerance varies by species, just as it does with higher plants. In most areas, mosses grow chiefly in moist, shaded areas, such as wooded areas and at the edges of streams, but they can grow anywhere in cool, humid, 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 over broadleaf trees, oaks over alders, or vice versa. [8] 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. [8] 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. [31]

It is generally believed that in the Northern Hemisphere, the north side of trees and rocks will generally have more luxuriant moss growth on average than other sides. [32] The reason is assumed to be because sunshine on the south side causes a dry environment. The reverse would be true in the Southern Hemisphere. However, naturalists feel that mosses grow on the damper side of trees and rocks. [7] 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, humid, cloudy climates, all sides of tree trunks and rocks may be equally moist enough for moss growth. Each species of moss requires certain amounts of moisture and sunlight and thus will grow on certain sections of the same tree or rock.

Some mosses grow underwater, or completely waterlogged. Many prefer well-drained locations. There are mosses that preferentially grow on rocks and tree trunks of various chemistries. [33]

Relationship with cyanobacteria

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 colonize moss and receive 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. [34]

Cultivation

A moss lawn in a garden in Japan 500px photo (189849595).jpeg
A moss lawn in a garden in Japan
The moss garden at the Bloedel Reserve, Bainbridge Island, Washington State. Moss Garden, Bloedel Reserve.jpg
The moss garden at the Bloedel Reserve, Bainbridge Island, Washington State.

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. [35] 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, humid, cloudy Pacific Northwest, moss is sometimes allowed to grow naturally as a moss lawn, one that needs little or no mowing, fertilizing or watering. In this case, grass is considered to be the weed. [36] 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. [31] 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. [37]

Green roofs and walls

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. [38] Mosses are also used on green walls.

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.

Aquascaping

Aquascaping uses many aquatic mosses. They do best at low nutrient, light, and heat levels, and propagate fairly readily. They help maintain a water chemistry suitable for aquarium fish. [39] They grow more slowly than many aquarium plants, and are fairly hardy. [40]

Inhibiting moss growth

Moss can be a troublesome weed in containerized nursery operations and greenhouses. [41] 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:

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. [42]

Uses

Wall covered in moss Moss on a wall @ Kanjirappally.jpg
Wall covered in moss

Traditional

Preindustrial societies made use of the mosses growing in their areas.

Laplanders, North American tribes, and other circumpolar people used mosses for bedding. [7] [31] 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. [31] Circumpolar and alpine peoples have used mosses for insulation in boots and mittens. Ötzi the Iceman had moss-packed boots. [31]

The capacity of dried mosses 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. [31] Tribes of the Pacific Northwest in the United States and Canada used mosses to clean salmon prior to drying it, and packed wet moss into pit ovens for steaming camas bulbs. Food storage baskets and boiling baskets were also packed with mosses. [31]

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.

Some Sphagnum mosses can absorb up to 20 times their own weight in water. [43] 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. [43] It is also claimed to have antibacterial properties. [44] Native Americans were one of the peoples to use Sphagnum for diapers and napkins, which is still done in Canada. [45]

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. [46] In Mexico, Moss is used as a Christmas decoration.

Moss bioreactor cultivating the moss Physcomitrella patens Bioreaktor quer2.jpg
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 [47] and the safe production of complex biopharmaceuticals in the moss bioreactor, developed by Ralf Reski and his co-workers. [48]

London installed several structures called "City Trees": moss-filled walls, each of which is claimed to have "the air-cleaning capability of 275 regular trees" by consuming nitrogen oxides and other types of air pollution and producing oxygen. [49]

See also

Related Research Articles

Gametophyte Haploid stage in the life cycle of plants and algae

A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.

Alternation of generations Reproductive cycle of plants and algae

Alternation of generations is the type of life cycle that occurs in those plants and algae in the Archaeplastida and the Heterokontophyta that have distinct haploid sexual and diploid asexual stages. In these groups, a multicellular haploid gametophyte with n chromosomes alternates with a multicellular diploid sporophyte with 2n chromosomes, made up of n pairs. A mature sporophyte produces haploid spores by meiosis, a process which reduces the number of chromosomes to half, from 2n to n.

Bryophyte Terrestrial plants that lack vascular tissue

Bryophytes are an informal group consisting of three divisions of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. They are characteristically limited in size and prefer moist habitats although they can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce via spores. Bryophytes are usually considered to be a paraphyletic group and not a monophyletic group, although some studies have produced contrary results. Regardless of their status, the name is convenient and remains in use as an informal collective term. The term "bryophyte" comes from Greek βρύον, bryon "tree-moss, oyster-green" and φυτόν, phyton "plant".

Embryophyte Subclade of green plants, also known as land plants

The Embryophyta or land plants are the most familiar group of green plants that form vegetation on earth. Embryophyta is a clade within the Phragmoplastophyta, a larger clade that also includes several green algae groups, and within this large clade the embryophytes are sister to the Zygnematophyceae/Mesotaeniaceae and consist of the bryophytes plus the polysporangiophytes. Living embryophytes therefore include hornworts, liverworts, mosses, ferns, lycophytes, gymnosperms and flowering plants. The Embryophyta are informally called land plants because they live primarily in terrestrial habitats, while the related green algae are primarily aquatic. Embryophytes are complex multicellular eukaryotes with specialized reproductive organs. The name derives from their innovative characteristic of nurturing the young embryo sporophyte during the early stages of its multicellular development within the tissues of the parent gametophyte. With very few exceptions, embryophytes obtain their energy by photosynthesis, that is by using the energy of sunlight to synthesize their food from carbon dioxide and water.

Marchantiophyta Botanical division of non-vascular land plants that have a gametophyte-dominant life cycle and lack stomata

The Marchantiophyta are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of genetic information.

Sporophyte Diploid multicellular stage in the life cycle of a plant or alga

A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga. It develops from the zygote produced when a haploid egg cell is fertilized by a haploid sperm and each sporophyte cell therefore has a double set of chromosomes, one set from each parent. All land plants, and most multicellular algae, have life cycles in which a multicellular diploid sporophyte phase alternates with a multicellular haploid gametophyte phase. In the seed plants, the largest groups of which are the gymnosperms and flowering plants (angiosperms), the sporophyte phase is more prominent than the gametophyte, and is the familiar green plant with its roots, stem, leaves and cones or flowers. In flowering plants the gametophytes are very reduced in size, and are represented by the germinated pollen and the embryo sac.

Hornwort One of the three Divisions of bryophytic plants

Hornworts are a group of bryophytes constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, the flattened, green plant body of a hornwort is the gametophyte plant.

Non-vascular plant

Non-vascular plants are plants without the vascular system consisting of xylem and phloem. Although non-vascular plants lack these particular tissues, many possess simpler tissues that have specialized functions for the internal transport of water.

<i>Fissidens adianthoides</i>

Fissidens adianthoides, the maidenhair pocketmoss, is a moss in the family of Fissidentaceaea. It was first collected by Hedwig in 1801.

<i>Sphagnum</i>

Sphagnum is a genus of approximately 380 accepted species of mosses, commonly known as "peat moss" though they are different as peat moss has a more acidic pH level. Accumulations of Sphagnum can store water, since both living and dead plants can hold large quantities of water inside their cells; plants may hold 16 to 26 times as much water as their dry weight, depending on the species. The empty cells help retain water in drier conditions.

<i>Takakia</i>

Takakia is a genus of two species of mosses known from western North America and central and eastern Asia. The genus is placed as a separate family, order and class among the mosses. It has had a history of uncertain placement, but the discovery of sporophytes clearly of the moss-type firmly supports placement with the mosses.

Monoicous plants are those species that bear both sperm and eggs on the same gametophyte. Dioicous plants are those that have gametophytes that produce only sperm or eggs but never both. The terms are used largely but not exclusively in the context of bryophytes. Both monoicous and dioicous gametophytes produce gametes in gametangia by mitosis rather than meiosis, so that sperm and eggs are genetically identical with their parent gametophyte. The states of being monoicous or dioicous are called monoicy and dioicy respectively.

<i>Buxbaumia</i>

Buxbaumia is a genus of twelve species of moss (Bryophyta). It was first named in 1742 by Albrecht von Haller and later brought into modern botanical nomenclature in 1801 by Johann Hedwig to commemorate Johann Christian Buxbaum, a German physician and botanist who discovered the moss in 1712 at the mouth of the Volga River. The moss is microscopic for most of its existence, and plants are noticeable only after they begin to produce their reproductive structures. The asymmetrical spore capsule has a distinctive shape and structure, some features of which appear to be transitional from those in primitive mosses to most modern mosses.

<i>Physcomitrella patens</i>

Physcomitrium patens, the spreading earthmoss, is a moss (bryophyte) used as a model organism for studies on plant evolution, development, and physiology.

<i>Hypnodendron comosum</i>

Hypnodendron comosum, commonly known as palm moss or palm tree moss, is a ground moss which can be divided into two varieties: Hypnodendron comosum var. comosum and Hypnodendron comosum var. sieberi. Both Hypnodendron varieties most commonly grow in damp locations in the temperate and tropical rainforests of New South Wales, Victoria, and Tasmania in southern Australia and in New Zealand.

<i>Pogonatum urnigerum</i>

Pogonatum urnigerum is a species of moss in the family Polytrichaceae, commonly called urn haircap. The name comes from "urna" meaning "urn" and "gerere" meaning "to bear" which is believed to be a reference made towards the plant's wide-mouthed capsule. It can be found on gravelly banks or similar habitats and can be identified by the blue tinge to the overall green colour. The stem of this moss is wine red and it has rhizoids that keep the moss anchored to substrates. It is an acrocarpous moss that grows vertically with an archegonium borne at the top of each fertilized female gametophyte shoot which develops an erect sporophyte.

<i>Climacium dendroides</i>

Climacium dendroides, also known as tree climacium moss, belongs in the order Hypnales and family Climaciaceae, in class Bryopsida and subclass Bryidae. It is identified as a "tree moss" due to its distinctive morphological features, and has four species identified across the Northern Hemisphere. The species name "dendroides" describes the tree-like morphology of the plant, and its genus name came from the structure of the perforations of peristome teeth. This plant was identified by Weber and Mohr in 1804. They often have stems that are around 2-10 cm tall and growing in the form of patches, looking like small palm-trees. They have yellow-green branches at the tip of stems. The leaves are around 2.5-3 mm long, with rounder stem leaves and pointier branch leaves. Their sporophytes are only abundant in late winter and early spring, and appears as a red-brown shoot with long stalk and cylindrical capsules.

<i>Polytrichum strictum</i>

Polytrichum strictum, commonly known as bog haircap moss or strict haircap, is an evergreen and perennial species of moss native to Sphagnum bogs and other moist habitats in temperate climates. It has a circumboreal distribution, and is also found in South America and Antarctica.

<i>Buxbaumia viridis</i>

Buxbuamia viridis, also known as the green shield-moss, is a rare bryophyte found sporadically throughout the northern hemisphere. The gametophyte of this moss is not macroscopically visible; the large, distinct sporophyte of B. viridis is the only identifying structure of this moss. This moss can be found singularly or in small groups on decaying wood, mostly in humid, sub-alpine to alpine Picea abies, Abies alba, or mixed tree forests. This moss is rare and conservation efforts are being made in most countries B. viridis is found in.

<i>Tortula muralis</i>

Tortula muralis, commonly known as wall screw-moss, is a species of moss in the family Pottiaceae. T. muralis is found throughout the world.

References

  1. Hubers, M.; Kerp, H. (2012). "Oldest known mosses discovered in Mississippian (late Visean) strata of Germany". Geology. 40 (8): 755–758. Bibcode:2012Geo....40..755H. doi:10.1130/G33122.1.
  2. 1 2 3 Goffinet, Bernard; William R. Buck (2004). Systematics of the Bryophyta (Mosses): From molecules to a revised classification. Monographs in Systematic Botany. Molecular Systematics of Bryophytes. 98. Missouri Botanical Garden Press. pp. 205–239. ISBN   978-1-930723-38-2.
  3. "Bryophyta". Merriam-Webster Dictionary .
  4. "bryophyte" . Oxford English Dictionary (Online ed.). Oxford University Press. (Subscription or participating institution membership required.)
  5. Ligrone, R.; Duckett, J.G.; Renzaglia, K.S. (2000). "Conducting tissues and phyletic relationships of bryophytes". Philos Trans R Soc Lond B Biol Sci. 355 (1398): 795–813. doi:10.1098/rstb.2000.0616. PMC   1692789 . PMID   10905610.
  6. 1 2 Lichens of North America, Irwin M. Brodo, Ms. Sylvia Duran Sharnoff, ISBN   978-0300082494, 2001
  7. 1 2 3 Mathews, Daniel (1994). Cascade-Olympic Natural History . Portland, Oregon: Audubon Society of Portland/Raven Editions. ISBN   978-0-9620782-0-0.
  8. 1 2 3 4 Pojar and MacKinnon (1994). Plants of the Pacific Northwest Coast. Vancouver, British Columbia: Lone Pine Publishing. ISBN   978-1-55105-040-9.
  9. van der Velde, M.; During, H. J.; van de Zande, L.; Bijlsma, R. (2001). "The reproductive biology of Polytrichum formosum: clonal structure and paternity revealed by microsatellites". Molecular Ecology. 10 (10): 2423–2434. doi:10.1046/j.0962-1083.2001.01385.x. PMID   11742546. S2CID   19716812.
  10. Johan L. van Leeuwen (July 23, 2010). "Launched at 36,000g". Science. 329 (5990): 395–6. doi:10.1126/science.1193047. PMID   20651138.
  11. Dwight K. Whitaker & Joan Edwards (July 23, 2010). "Sphagnum Moss Disperses Spores with Vortex Rings". Science. 329 (5990): 406. Bibcode:2010Sci...329..406W. doi:10.1126/science.1190179. PMID   20651145. S2CID   206526774.
  12. Cronberg, N.; Natcheva, R.; Hedlund, K. (2006). "Microarthropods Mediate Sperm Transfer in Mosses". Science. 313 (5791): 1255. doi:10.1126/science.1128707. PMID   16946062. S2CID   11555211.
  13. 1 2 Rosenstiel, T. N.; Shortlidge, E. E.; Melnychenko, A. N.; Pankow, J. F.; Eppley, S. M. (2012). "Sex-specific volatile compounds influence microarthropod-mediated fertilization of moss". Nature. 489 (7416): 431–433. Bibcode:2012Natur.489..431R. doi:10.1038/nature11330. PMID   22810584. S2CID   4419337.
  14. Vaizey, J. R. (1890). "On the Morphology of the Sporophyte of Splachnum luteum". Annals of Botany. 1: 1–8. doi:10.1093/oxfordjournals.aob.a090623.
  15. 1 2 3 Une, Kouji (1985). "Sexual dimorphism in the Japanese species of Macromitrium Brid.(Musci: Orthotrichaceae)". The Journal of the Hattori Botanical Laboratory Devoted to Bryology and Lichenology. 59: 487–513.
  16. Blackstock, T. H. (1987). "The male gametophores of Leucobryum glaucum (Hedw.) Ångstr. and L. juniperoideum (Brid.) C. Muell. in two Welsh woodlands". Journal of Bryology. 14 (3): 535–541. doi:10.1179/jbr.1987.14.3.535.
  17. 1 2 Loveland, Hugh Frank (1956). Sexual dimorphism in the moss genus Dicranum Hedw. (Dissertation). University of Michigan.
  18. 1 2 Wallace, M. H. (1970). Developmental morphology and sexual dimorphism in Homalothecium megaptilum (Sull.) Robins. (Dissertation). Washington State University.
  19. Sagmo Solli, I. M.; Söderström, Lars; Bakken, Solveig; Flatberg, Kjell Ivar; Pedersen, Bård (1998). "Studies of fertility of Dicranum majus in two populations with contrasted sporophyte production". Journal of Bryology. 22 (1): 3–8. doi:10.1179/jbr.2000.22.1.3. S2CID   85349694.
  20. 1 2 3 4 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.
  21. Ramsay, Helen P.; Berrie, G. K. (1982). "Sex determination in bryophytes". Journal of the Hattori Botanical Laboratory. 52: 255–274.
  22. Rensing, S.A.; Lang, D; Zimmer, A.D.; Terry, A.; Salamov, A; Shapiro, H.; Nishiyama, T.; et al. (January 2008). "The Physcomitrella genome reveals evolutionary insights into the conquest of land by plants" (PDF). Science. 319 (5859): 64–9. Bibcode:2008Sci...319...64R. doi:10.1126/science.1150646. hdl:11858/00-001M-0000-0012-3787-A. PMID   18079367. S2CID   11115152.
  23. 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.
  24. 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.
  25. 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.
  26. Thomas, B.A. (1972). "A probable moss from the Lower Carboniferous of the Forest of Dean, Gloucestershire". Annals of Botany. 36 (1): 155–161. doi:10.1093/oxfordjournals.aob.a084568. ISSN   1095-8290. JSTOR   42752024.
  27. 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.
  28. 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 (2): 566–571. doi: 10.1111/j.1420-9101.2007.01479.x . PMID   18205784.
  29. "First land plants plunged Earth into ice age". Newscientist.com. Archived from the original on 2013-09-23. Retrieved 2013-09-11.
  30. "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.
  31. 1 2 3 4 5 6 7 8 Kimmerer, Robin Wall (2003). Gathering Moss. Corvallis, Oregon: Oregon State University Press. ISBN   978-0-87071-499-3.
  32. Porley, Ron; Hodgetts, Nick (2005). Mosses & Liverworts. London: Collins. pp. 80–81. ISBN   978-0-00-220212-1.
  33. Fletcher, Michael (2006). Moss Growers Handbook (3rd ed.). Reading Berkshire: SevenTy Press. ISBN   0-9517176-0-X.
  34. 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: 150. doi:10.3389/fmicb.2013.00150. ISSN   1664-302X. PMC   3683619 . PMID   23785359.
  35. Chan, Peter (1993). Bonsai Masterclass. New York City: Sterling Publishing Co. ISBN   978-0-8069-6763-9.
  36. Smith, Sally W. (1998). Sunset Western Garden Problem Solver . Menlo Park, California: Sunset Books. ISBN   978-0-376-06132-4.
  37. "The Bloedel Reserve". Archived from the original on 16 April 2011. Retrieved 24 April 2011.
  38. "RoofTopGarden". Archived from the original on 24 April 2011. Retrieved 22 May 2011.
  39. "A Guide to Keeping and Growing Aquatic Moss". Aquascaping Love. 12 April 2016.
  40. "Mosses". www.aquasabi.com.
  41. 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.
  42. 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.
  43. 1 2 The Plant Underworld, Sphagnum and Water, Australian Botanic Garden Archived 2014-02-17 at the Wayback Machine
  44. 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. PMID   19187129. S2CID   1545021.
  45. 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.
  46. Engman, Max; D. G. Kirby (1989). Finland: people, nation, state. C. Hurst & Co. p. 45. ISBN   0-253-32067-4.
  47. 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.
  48. 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.
  49. Landon, Alex (2020-01-07). "City Trees: London Has New Artificial Trees That Eat Pollution". Secret London. Retrieved 2020-02-09.