Mosses are small,
non-vascular flower
Flowers, also known as blooms and blossoms, are the reproductive structures of flowering plants ( angiosperms). Typically, they are structured in four circular levels, called whorls, around the end of a stalk. These whorls include: calyx, m ...
less
plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s in the taxonomic
division Bryophyta (, ) ''
sensu stricto''. Bryophyta (''
sensu lato'',
Schimp. 1879
) may also refer to the parent group
bryophyte
Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s, which comprise
liverworts
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
, mosses, and
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s.
Mosses typically form dense green clumps or mats, often in damp or shady locations. The individual plants are usually composed of simple
leaves
A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
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.
Mosses do not have
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s and after fertilisation develop
sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
s with unbranched stalks topped with single capsules containing
spores
In biology, a spore is a unit of sexual (in fungi) or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plant ...
. They are typically tall, though some species are much larger.
''Dawsonia'', the tallest moss in the world, can grow to in height. There are approximately 12,000 species.
Mosses are commonly confused with liverworts, hornworts and
lichen
A lichen ( , ) is a hybrid colony (biology), colony of algae or cyanobacteria living symbiotically among hypha, filaments of multiple fungus species, along with yeasts and bacteria embedded in the cortex or "skin", in a mutualism (biology), m ...
s.
[Lichens of North America, Irwin M. Brodo, Sylvia Duran Sharnoff, , 2001] Although often described as
non-vascular plants, many mosses have advanced vascular systems. Like liverworts and hornworts, the
haploid
Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
gametophyte
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 se ...
generation of mosses is the dominant phase of the
life cycle. This contrasts with the pattern in all vascular plants (
seed plants and
pteridophytes), where the
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
sporophyte generation is dominant. Lichens may superficially resemble mosses, and sometimes have common names that include the word "moss" (e.g., "
reindeer moss
''Cladonia rangiferina'', also known as reindeer cup lichen, reindeer lichen (cf. Swedish language, Sw. ''renlav'') or grey reindeer lichen, is a light-coloured fruticose lichen, fruticose, Cladonia, cup lichen species in the family Cladoniaceae. ...
" or "
Iceland moss"), but they are fungal symbioses and not related to mosses.
The main commercial significance of mosses is as the main constituent of
peat
Peat is an accumulation of partially Decomposition, decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, Moorland, moors, or muskegs. ''Sphagnum'' moss, also called peat moss, is one of the most ...
(mostly the genus ''
Sphagnum
''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
''), although they are also used for decorative purposes, such as in gardens and in the
florist
Floristry is the production, commerce, and trade in flowers. It encompasses flower care and handling, floral design and arrangement, merchandising, production, display and flower delivery. Wholesale florists sell bulk flowers and related s ...
trade. Traditional uses of mosses included as insulation and for the ability to absorb liquids up to 20 times their weight. Mosses are
keystone species
A keystone species is a species that has a disproportionately large effect on its natural environment relative to its abundance. The concept was introduced in 1969 by the zoologist Robert T. Paine. Keystone species play a critical role in main ...
and benefit
habitat restoration
Ecological restoration, or ecosystem restoration, is the process of assisting the recovery of an ecosystem that has been degraded, damaged, destroyed or transformed. It is distinct from conservation in that it attempts to retroactively repair ...
and
reforestation
Reforestation is the practice of restoring previously existing forests and woodlands that have been destroyed or damaged. The prior forest destruction might have happened through deforestation, clearcutting or wildfires. Three important purpose ...
.
Physical characteristics
Description

Botanically, mosses are
non-vascular plant
Non-vascular plants are plants without a vascular system consisting of xylem and phloem. Instead, they may possess simpler tissues that have specialized functions for the internal transport of water.
Non-vascular plants include two distantly rel ...
s in the land plant division Bryophyta. They are usually small (a few centimeters tall)
herbaceous
Herbaceous plants are vascular plants that have no persistent woody stems above ground. This broad category of plants includes many perennials, and nearly all annuals and biennials.
Definitions of "herb" and "herbaceous"
The fourth edition of ...
(non-woody) plants that absorb water and nutrients mainly through their leaves and harvest
carbon dioxide
Carbon dioxide is a chemical compound with the chemical formula . It is made up of molecules that each have one carbon atom covalent bond, covalently double bonded to two oxygen atoms. It is found in a gas state at room temperature and at norma ...
and sunlight to create food by
photosynthesis
Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
.
With the exception of the ancient group
Takakiopsida
''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.
Discovery
''Takakia'' was discovered in the Himalayas and ...
, no known mosses form
mycorrhiza
A mycorrhiza (; , mycorrhiza, or mycorrhizas) is a symbiotic association between a fungus and a plant. The term mycorrhiza refers to the role of the fungus in the plant's rhizosphere, the plant root system and its surroundings. Mycorrhizae play ...
, but bryophilous fungi is widespread in moss and other bryophytes, where they live as saprotrophs, parasites, pathogens and mutualists, some of them
endophyte
An endophyte is an endosymbiont, often a bacterium or fungus, that lives within a plant for at least part of its life cycle without causing apparent disease. Endophytes are ubiquitous and have been found in all species of plants studied to date; ...
s. Mosses differ from
vascular plants in lacking water-bearing
xylem
Xylem is one of the two types of transport tissue (biology), tissue in vascular plants, the other being phloem; both of these are part of the vascular bundle. The basic function of the xylem is to transport water upward from the roots to parts o ...
tracheids or
vessels. As in
liverworts
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
and
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s, the
haploid
Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
gametophyte
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 se ...
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
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
generation is dominant. Mosses reproduce using
spore
In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s, not
seed
In botany, a seed is a plant structure containing an embryo and stored nutrients in a protective coat called a ''testa''. More generally, the term "seed" means anything that can be Sowing, sown, which may include seed and husk or tuber. Seeds ...
s, and have no flowers.

Moss gametophytes have stems which may be simple or branched and upright (acrocarp) or prostrate (pleurocarp). The early divergent classes Takakiopsida, Sphagnopsida, Andreaeopsida and Andreaeobryopsida either lack
stoma
In botany, a stoma (: stomata, from Greek language, Greek ''στόμα'', "mouth"), also called a stomate (: stomates), is a pore found in the Epidermis (botany), epidermis of leaves, stems, and other organs, that controls the rate of gas exc ...
ta or have pseudostomata that do not form pores. In the remaining classes, stomata have been lost more than 60 times. Their leaves are simple, usually only a single layer of cells with no internal air spaces, often with thicker midribs (nerves). The nerve can run beyond the edge of the leaf tip, termed excurrent. The tip of the leaf blade can be extended as a hair point, made of colourless cells. These appear white against the dark green of the leaves. The edge of the leaf can be smooth or it may have teeth. There may be a distinct type of cell defining the edge of the leaf, distinct in shape and/or colour from the other leaf cells.
Moss has threadlike
rhizoids that anchor them to their substrate, comparable to
root hairs rather than the more substantial
root
In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...
structures of
spermatophytes.
Mosses do not absorb water or nutrients from their substrate through their rhizoids. They can be distinguished from
liverworts
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
(
Marchantiophyta
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
or Hepaticae) by their multi-cellular rhizoids. Spore-bearing capsules or
sporangia
A sporangium (from Late Latin, ; : sporangia) is an enclosure in which spores are formed. It can be composed of a unicellular organism, single cell or can be multicellular organism, multicellular. Virtually all plants, fungus, fungi, and many ot ...
of mosses are borne singly on long, unbranched stems, thereby distinguishing them from the
polysporangiophytes, which include all vascular plants. The spore-producing sporophytes (i.e. the
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
multicellular generation) are short-lived and usually capable of photosynthesis, but are dependent on the gametophyte for water supply and most or all of its nutrients.
Also, in the majority of mosses, the spore-bearing capsule enlarges and matures after its stalk elongates, while in liverworts the capsule enlarges and matures before its stalk elongates.
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
plant
Plants are the eukaryotes that form the Kingdom (biology), kingdom Plantae; they are predominantly Photosynthesis, photosynthetic. This means that they obtain their energy from sunlight, using chloroplasts derived from endosymbiosis with c ...
s have two sets of
chromosome
A chromosome is a package of DNA containing part or all of the genetic material of an organism. In most chromosomes, the very long thin DNA fibers are coated with nucleosome-forming packaging proteins; in eukaryotic cells, the most import ...
s in their vegetative cells and are said to be
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Here ''sets of chromosomes'' refers to the number of maternal and paternal chromosome copies, ...
, i.e. each chromosome has a partner that contains the same, or similar, genetic information. By contrast, mosses and other
bryophyte
Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s have only a single set of chromosomes and so are
haploid
Ploidy () is the number of complete sets of chromosomes in a cell (biology), cell, and hence the number of possible alleles for Autosome, autosomal and Pseudoautosomal region, pseudoautosomal genes. Here ''sets of chromosomes'' refers to the num ...
(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
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
stage.

The moss life-cycle starts with a haploid
spore
In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
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
Gametophores are prominent structures in seedless plants on which the reproductive organs are borne. The word gametophore and ‘-phore’ (Greek Φορά, "to be carried"). In mosses, liverworts and ferns (Archegoniata), the gametophores suppor ...
("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.
Gametophore tip growth is disrupted by fungal
chitin
Chitin (carbon, C8hydrogen, H13oxygen, O5nitrogen, N)n ( ) is a long-chain polymer of N-Acetylglucosamine, ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is the second most abundant polysaccharide in nature (behind only cell ...
.
Galotto ''et al.'', 2020 applied
chitooctaose and found that tips detected and responded to this chitin derivative by changing
gene expression
Gene expression is the process (including its Regulation of gene expression, regulation) by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, proteins or non-coding RNA, ...
.
They concluded that this defense response was probably
conserved from the
most recent common ancestor
A most recent common ancestor (MRCA), also known as a last common ancestor (LCA), is the most recent individual from which all organisms of a set are inferred to have descended. The most recent common ancestor of a higher taxon is generally assu ...
of
bryophyte
Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s and
tracheophytes.
Orr ''et al.'', 2020 found that the
microtubule
Microtubules are polymers of tubulin that form part of the cytoskeleton and provide structure and shape to eukaryotic cells. Microtubules can be as long as 50 micrometres, as wide as 23 to 27 nanometer, nm and have an inner diameter bet ...
s of growing tip cells were structurally similar to
F-actin
Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ...
and served a similar purpose.
Mosses can be either
dioicous (compare
dioecious
Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
in seed plants) or
monoicous (compare
monoecious
Monoecy (; adj. monoecious ) is a sexual system in seed plants where separate male and female cones or flowers are present on the same plant. It is a monomorphic sexual system comparable with gynomonoecy, andromonoecy and trimonoecy, and contras ...
). 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
Fertilisation or fertilization (see spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give rise to a zygote and initiate its development into a new individual organism or of ...
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.
After fertilisation, the immature sporophyte pushes its way out of the archegonial venter. It takes several months for the
sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
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
Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
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
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
''
Sphagnum
''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
'' the
spores are projected about 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''.

It has recently been found that microarthropods, such as
springtails
Springtails (class Collembola) form the largest of the three lineages of modern Hexapoda, hexapods that are no longer considered insects. Although the three lineages are sometimes grouped together in a class called Entognatha because they have in ...
and
mites
Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...
, can effect moss fertilization
and that this process is mediated by moss-emitted scents. Male and female
fire moss, for example, emit different and complex volatile organic scents.
Female plants emit more compounds than male plants.
Springtails
Springtails (class Collembola) form the largest of the three lineages of modern Hexapoda, hexapods that are no longer considered insects. Although the three lineages are sometimes grouped together in a class called Entognatha because they have in ...
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.
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.
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
Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the f ...
, 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
spore
In biology, a spore is a unit of sexual reproduction, sexual (in fungi) or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores fo ...
s 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.
More often, 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.
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.
The nature of such a substance is unknown, but the phytohormone
auxin
Auxins (plural of auxin ) are a class of plant hormones (or plant-growth regulators) with some morphogen-like characteristics. Auxins play a cardinal role in coordination of many growth and behavioral processes in plant life cycles and are essent ...
may be involved
Having the males growing as dwarfs on the female is expected to increase the
fertilization
Fertilisation or fertilization (see American and British English spelling differences#-ise, -ize (-isation, -ization), spelling differences), also known as generative fertilisation, syngamy and impregnation, is the fusion of gametes to give ...
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.
Dwarf males occur in several unrelated
lineages and may be more common than previously thought.
For example, it is estimated that between one quarter and half of all
dioicous pleurocarps have dwarf males.
DNA repair
The moss ''
Physcomitrium patens'' has been used as a
model organism
A model organism is a non-human species that is extensively studied to understand particular biological phenomena, with the expectation that discoveries made in the model organism will provide insight into the workings of other organisms. Mo ...
to study how plants
repair
The technical meaning of maintenance involves functional checks, servicing, repairing or replacing of necessary devices, equipment, machinery, building infrastructure and supporting utilities in industrial, business, and residential installat ...
damage to their DNA, especially the repair mechanism known as
homologous recombination
Homologous recombination is a type of genetic recombination in which genetic information is exchanged between two similar or identical molecules of double-stranded or single-stranded nucleic acids (usually DNA as in Cell (biology), cellular organi ...
. If the plant cannot repair DNA damage, e.g.,
double-strand breaks
DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. A weakened capacity for DNA repair is a risk factor for the development of cancer. DNA is constantly modified ...
, in their
somatic cell
In cellular biology, a somatic cell (), or vegetal cell, is any biological cell forming the body of a multicellular organism other than a gamete, germ cell, gametocyte or undifferentiated stem cell. Somatic cells compose the body of an organism ...
s, the cells can lose normal functions or die. If this occurs during
meiosis
Meiosis () is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, the sperm or egg cells. It involves two rounds of division that ultimately result in four cells, each with only one c ...
(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.
''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 ''Rp''RAD51, 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.
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.
Classification
More recently, mosses have been grouped with the
liverworts
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
and
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s in the division Bryophyta (
bryophyte
Bryophytes () are a group of embryophyte, land plants (embryophytes), sometimes treated as a taxonomic Division (taxonomy), division referred to as Bryophyta ''Sensu#Common qualifiers, sensu lato'', that contains three groups of non-vascular pla ...
s, or Bryophyta ''sensu lato'').
The bryophyte division itself contains three (former) divisions: Bryophyta (mosses), Marchantiophyta
Liverworts are a group of non-vascular plant, non-vascular embryophyte, land plants forming the division Marchantiophyta (). They may also be referred to as hepatics. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in wh ...
(liverworts) and Anthocerotophyta
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
(hornworts); it has been proposed that these latter divisions are de-ranked to the classes Bryopsida, Marchantiopsida, and Anthocerotopsida, respectively. The mosses and liverworts are now considered to belong to a clade called Setaphyta.
The mosses, (Bryophyta sensu stricto), are divided into eight classes:
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
''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
'', as well as fossil taxa. ''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 high. The tallest land moss, a member of the Polytrichidae is probably '' Dawsonia superba'', a native to New Zealand
New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
and other parts of Australasia
Australasia is a subregion of Oceania, comprising Australia, New Zealand (overlapping with Polynesia), and sometimes including New Guinea and surrounding islands (overlapping with Melanesia). The term is used in a number of different context ...
.
Geological history
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
The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
of Antarctica and Russia, and a case has been made for Carboniferous
The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
mosses. It has further been claimed that tube-like fossils from the Silurian
The Silurian ( ) is a geologic period and system spanning 23.5 million years from the end of the Ordovician Period, at million years ago ( Mya), to the beginning of the Devonian Period, Mya. The Silurian is the third and shortest period of t ...
are the macerated remains of moss calyptræ. Mosses also appear to evolve 2–3 times slower than ferns, gymnosperm
The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...
s and angiosperm
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (). The term angiosperm is derived from the Greek words (; 'container, vessel') and (; 'seed'), meaning that the seeds are enclosed within a fruit ...
s.
Recent research shows that ancient moss could explain why the Ordovician
The Ordovician ( ) is a geologic period and System (geology), system, the second of six periods of the Paleozoic Era (geology), Era, and the second of twelve periods of the Phanerozoic Eon (geology), Eon. The Ordovician spans 41.6 million years f ...
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 significant amounts 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.
Ecology
Habitat
File:Mossopolis.jpg, Dense moss colonies in a cool coastal forest
File:MossForest.jpg, A cool high altitude/latitude moss forest; the forest floor is covered in moss, beneath conifers
File:Route 1 , Between Vik and Kirkjubæjarklaustur - panoramio (7).jpg, Moss colonizes a basalt
Basalt (; ) is an aphanite, aphanitic (fine-grained) extrusive igneous rock formed from the rapid cooling of low-viscosity lava rich in magnesium and iron (mafic lava) exposed at or very near the planetary surface, surface of a terrestrial ...
flow, in Iceland
Iceland is a Nordic countries, Nordic island country between the Atlantic Ocean, North Atlantic and Arctic Oceans, on the Mid-Atlantic Ridge between North America and Europe. It is culturally and politically linked with Europe and is the regi ...
File:Moss (Iceland) 03.jpg, Moss growing along seep
A seep or flush is a moist or wet place where water, usually groundwater, reaches the Earth's surface from an underground aquifer.
Description
Seeps are usually not of sufficient volume to be flowing beyond their immediate above-ground location. ...
s and springs in newly deposited basaltic
Basalt (; ) is an aphanitic (fine-grained) extrusive igneous rock formed from the rapid cooling of low-viscosity lava rich in magnesium and iron ( mafic lava) exposed at or very near the surface of a rocky planet or moon. More than 90% ...
rock, Iceland.
File:Steinerne Rinne bei Obererasbach im Altmühltal.jpg, Moss growing along the stream from a karst spring
A karst spring or karstic spring is a Spring (hydrology), spring (exsurgence, outflow of groundwater) that is part of a karst hydrological system.
Description
Because of their often conical or inverted bowl shape, karst springs are also known i ...
; travertine deposits from the stream water and the moss overgrows it, forming this ridge, with the stream on top.
File:Winter moss.jpg, Moss with sporophytes on brick
File:Mech plonnik mlode sporofity.jpg, Young sporophyte
A sporophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the biological life cycle, life cycles of plants and algae. It is a diploid multicellular organism which produces asexual Spo ...
s of the common moss ''Tortula muralis'' (wall screw-moss)
File:Taiwan 2009 JinGuaShi Historic Gold Mine Moss Covered Retaining Wall FRD 8940.jpg, Retaining wall
Retaining walls are relatively rigid walls used for supporting soil laterally so that it can be retained at different levels on the two sides. Retaining walls are structures designed to restrain soil to a slope that it would not naturally keep to ...
covered in moss
File:Michiganmosspatch.jpg, A small clump of moss beneath a conifer (a shady, usually dry place)
File:MossOnConcreteWall.jpg, Moss on a concrete wall
File:Moss (Bryophyta) on the forest floor in Broken Bow, Oklahoma.jpg, Moss (Bryophyta) on the forest floor in Broken Bow, Oklahoma
Mosses live in almost every terrestrial habitat type on Earth. Though mosses are particularly abundant in certain habitats such as peatlands, where Sphagnum mosses are the dominant organism, and in moist boreal, temperate, and montane tropical forests, mosses grow in many other habitats, including habitats with conditions too extreme for vascular plants to survive. Desiccation tolerant mosses are important in arid and semi-arid ecosystems, where they help form biocrusts that mediate extremes of soil temperature, regulate soil moisture, and regulate the release and uptake of carbon. Mosses can live on substrates heated by geothermal activity to temperatures exceeding 50 degrees Celsius, on walls and pavement in urban areas, and in Antarctica. Moss diversity is generally not associated with latitude; boreal and temperate moss diversity is similar to tropical moss diversity. Moss diversity hotspots are found in the northern Andes mountains, Mexico, the Himalayan mountains, Madagascar, Japan, the highlands of eastern Africa, Southeast Asia, central Europe, Scandinavia, and British Columbia.
Moss gametophyte
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 se ...
s are autotroph
An autotroph is an organism that can convert Abiotic component, abiotic sources of energy into energy stored in organic compounds, which can be used by Heterotroph, other organisms. Autotrophs produce complex organic compounds (such as carbohy ...
ic and require sunlight
Sunlight is the portion of the electromagnetic radiation which is emitted by the Sun (i.e. solar radiation) and received by the Earth, in particular the visible spectrum, visible light perceptible to the human eye as well as invisible infrare ...
to perform photosynthesis
Photosynthesis ( ) is a system of biological processes by which photosynthetic organisms, such as most plants, algae, and cyanobacteria, convert light energy, typically from sunlight, into the chemical energy necessary to fuel their metabo ...
. In most areas, mosses grow chiefly in moist, shaded areas, such as wooded areas and at the edges of streams, but shade tolerance varies by species.
Different moss species grow on different substrates 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 humus
In classical soil science, humus is the dark organic matter in soil that is formed by the decomposition of plant and animal matter. It is a kind of soil organic matter. It is rich in nutrients and retains moisture in the soil. Humus is the Lati ...
y soil, downed logs, burnt stumps, tree trunk bases, upper tree trunks, and tree branches or in bog
A bog or bogland is a wetland that accumulates peat as a deposit of dead plant materials often mosses, typically sphagnum moss. It is one of the four main types of wetlands. Other names for bogs include mire, mosses, quagmire, and musk ...
s. Moss species growing on or under trees are often specific about the species of trees they grow on, such as preferring conifers
Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
over broadleaf trees, oaks over alder
Alders are trees of the genus ''Alnus'' in the birch family Betulaceae. The genus includes about 35 species of monoecious trees and shrubs, a few reaching a large size, distributed throughout the north temperate zone with a few species ex ...
s, or vice versa. While mosses often grow on trees as epiphytes
An epiphyte is a plant or plant-like organism that grows on the surface of another plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphyt ...
, 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. Such aquatic or semi-aquatic mosses can greatly exceed the normal range of lengths seen in terrestrial mosses. Individual plants or more long are common in ''Sphagnum'' species for example. But even aquatic species of moss and other bryophytes needs their mature capsules to be exposed to air by seta elongation or seasonal lowering of water level to be able to reproduce.
Wherever they occur, mosses require liquid water for at least part of the year to complete fertilisation. Many mosses can survive desiccation
Desiccation is the state of extreme dryness, or the process of extreme drying. A desiccant is a hygroscopic (attracts and holds water) substance that induces or sustains such a state in its local vicinity in a moderately sealed container. The ...
, sometimes for months, returning to life within a few hours of rehydration. Mosses in arid habitats, such as the moss '' Syntrichia caninervis'', have adaptations for collecting non-rainfall sources of moisture like dew
Dew is water in the form of droplets that appears on thin, exposed objects in the morning or evening due to condensation.
As the exposed surface cools by thermal radiation, radiating its heat, atmospheric moisture condenses at a rate grea ...
and fog, capturing condensation from the air.
It is generally believed that in the Northern Hemisphere
The Northern Hemisphere is the half of Earth that is north of the equator. For other planets in the Solar System, north is defined by humans as being in the same celestial sphere, celestial hemisphere relative to the invariable plane of the Solar ...
, the north side of trees and rocks will generally have more luxuriant moss growth on average than other sides. 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. Some naturalists feel that mosses grow on the damper side of trees and rocks. In some cases, such as sunny climates in temperate
In geography, the temperate climates of Earth occur in the middle latitudes (approximately 23.5° to 66.5° N/S of the Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ran ...
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.
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
Nitrogen fixation is a chemical process by which molecular dinitrogen () is converted into ammonia (). It occurs both biologically and abiological nitrogen fixation, abiologically in chemical industry, chemical industries. Biological nitrogen ...
cyanobacteria
Cyanobacteria ( ) are a group of autotrophic gram-negative bacteria that can obtain biological energy via oxygenic photosynthesis. The name "cyanobacteria" () refers to their bluish green (cyan) color, which forms the basis of cyanobacteri ...
. 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.
Cultivation
Moss is often considered a weed
A weed is a plant considered undesirable in a particular situation, growing where it conflicts with human preferences, needs, or goals.Harlan, J. R., & deWet, J. M. (1965). Some thoughts about weeds. ''Economic botany'', ''19''(1), 16-24. Pla ...
in grass lawns, but is deliberately encouraged to grow under aesthetic principles exemplified by Japanese garden
are traditional gardens whose designs are accompanied by Japanese aesthetics and philosophical ideas, avoid artificial ornamentation, and highlight the natural landscape. Plants and worn, aged materials are generally used by Japanese garden desig ...
ing. 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
Bonsai (; , ) is the Japanese art of Horticulture, growing and shaping miniature trees in containers, with a long documented history of influences and native Japanese development over a thousand years, and with unique aesthetics, cultural hist ...
to cover the soil and enhance the impression of age. Rules of cultivation are not widely established. Moss collections are quite often begun using samples transplanted from the wild in a water-retaining bag. Some 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
A brick is a type of construction material used to build walls, pavements and other elements in masonry construction. Properly, the term ''brick'' denotes a unit primarily composed of clay. But is now also used informally to denote building un ...
, wood
Wood is a structural tissue/material found as xylem in the stems and roots of trees and other woody plants. It is an organic materiala natural composite of cellulosic fibers that are strong in tension and embedded in a matrix of lignin t ...
, and certain coarse concrete mixtures, are hospitable to moss. Surfaces can also be prepared with acidic substances, including buttermilk
Buttermilk is a fermented dairy drink. Traditionally, it was the liquid left behind after churning butter out of cultured cream. As most modern butter in Western countries is not made with cultured cream but uncultured sweet cream, most mode ...
, yogurt
Yogurt (; , from , ; also spelled yoghurt, yogourt or yoghourt) is a food produced by bacterial Fermentation (food), fermentation of milk. Fermentation of sugars in the milk by these bacteria produces lactic acid, which acts on milk protein to ...
, urine
Urine is a liquid by-product of metabolism in humans and many other animals. In placental mammals, urine flows from the Kidney (vertebrates), kidneys through the ureters to the urinary bladder and exits the urethra through the penile meatus (mal ...
, and gently puréed mixtures of moss samples, water and ericaceous compost.
In the cool, humid, cloudy Pacific Northwest
The Pacific Northwest (PNW; ) is a geographic region in Western North America bounded by its coastal waters of the Pacific Ocean to the west and, loosely, by the Rocky Mountains to the east. Though no official boundary exists, the most common ...
, 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. 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. 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.
Green roofs and walls
Mosses are sometimes used in green roof
A green roof or living roof is a roof of a building that is partially or completely covered with vegetation and a growing medium, planted over a waterproofing membrane. It may also include additional layers such as a root barrier and drainage ...
s. 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. Mosses are also used on green wall
A green wall is a vertical built structure intentionally covered by vegetation. Green walls include a vertically applied growth medium such as soil, substitute substrate, or hydroculture felt; as well as an integrated hydration and fertigation ...
s.
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
Aquascaping is the craft of arranging aquatic plants, as well as rocks, stones, cavework, or driftwood, in an aesthetically pleasing manner within an aquarium—in effect, gardening under water. Aquascape designs include a number of distinct styl ...
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. They grow more slowly than many aquarium plants, and are fairly hardy.
Growth inhibition
Moss can be a troublesome weed in containerized nursery operations and greenhouses. 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
Water is an inorganic compound with the chemical formula . It is a transparent, tasteless, odorless, and Color of water, nearly colorless chemical substance. It is the main constituent of Earth's hydrosphere and the fluids of all known liv ...
through drainage
Drainage is the natural or artificial removal of a surface's water and sub-surface water from an area with excess water. The internal drainage of most agricultural soils can prevent severe waterlogging (anaerobic conditions that harm root gro ...
.
* Increasing direct sunlight.
* Increasing number and resources available for competitive plants like grass
Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family (biology), family of monocotyledonous flowering plants commonly known as grasses. It includes the cereal grasses, bamboos, the grasses of natural grassland and spe ...
es.
* Increasing the soil pH
Soil pH is a measure of the acidity or basicity (alkalinity) of a soil. Soil pH is a key characteristic that can be used to make informative analysis both qualitative and quantitatively regarding soil characteristics. pH is defined as the neg ...
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 fertilizer
A fertilizer or fertiliser is any material of natural or synthetic origin that is applied to soil or to plant tissues to supply plant nutrients. Fertilizers may be distinct from liming materials or other non-nutrient soil amendments. Man ...
s. Sulfur
Sulfur ( American spelling and the preferred IUPAC name) or sulphur ( Commonwealth spelling) is a chemical element; it has symbol S and atomic number 16. It is abundant, multivalent and nonmetallic. Under normal conditions, sulfur atoms ...
and iron
Iron is a chemical element; it has symbol Fe () and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, forming much of Earth's o ...
are essential nutrients for some competing plants like grasses. Killing moss will not prevent regrowth unless conditions favorable to their growth are changed.
Uses
Traditional
Preindustrial societies made use of the mosses growing in their areas.
Sámi people, North American tribes, and other circumpolar peoples
Circumpolar peoples and Arctic peoples are umbrella terms for the various indigenous peoples of the Arctic region.
Approximately four million people are resident in the Arctic, among which 10 percent are indigenous peoples belonging to a vast nu ...
used mosses for bedding. 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 cabin
A log cabin is a small log house, especially a minimally finished or less architecturally sophisticated structure. Log cabins have an ancient history in Europe, and in America are often associated with first-generation home building by settl ...
s, and tribes of the northeastern United States and southeastern Canada used moss to fill chinks in wooden longhouses. Circumpolar and alpine peoples have used mosses for insulation in boots and mittens. Ötzi the Iceman had moss-packed boots.
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. 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.
Recent research investigating the Neanderthals remains recovered from El Sidrón have provided evidence that their diet would have consisted primarily of pine nuts, moss and mushrooms. This is contrasted by evidence from other European locations, which point to a more carnivorous diet.
In Finland
Finland, officially the Republic of Finland, is a Nordic country in Northern Europe. It borders Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bothnia to the west and the Gulf of Finland to the south, ...
, peat mosses have been used to make bread during famines.
Commercial
There is a substantial market in mosses gathered from the wild. The uses for intact moss are principally in the florist
Floristry is the production, commerce, and trade in flowers. It encompasses flower care and handling, floral design and arrangement, merchandising, production, display and flower delivery. Wholesale florists sell bulk flowers and related s ...
trade and for home decoration. Decaying moss in the genus ''Sphagnum'' is also the major component of peat
Peat is an accumulation of partially Decomposition, decayed vegetation or organic matter. It is unique to natural areas called peatlands, bogs, mires, Moorland, moors, or muskegs. ''Sphagnum'' moss, also called peat moss, is one of the most ...
, which is "mined" for use as a fuel
A fuel is any material that can be made to react with other substances so that it releases energy as thermal energy or to be used for work (physics), work. The concept was originally applied solely to those materials capable of releasing chem ...
, as a horticultural soil additive, and in smoking malt
Malt is any cereal grain that has been made to germinate by soaking in water and then stopped from germinating further by drying with hot air, a process known as "malting".
Malted grain is used to make beer, whisky, malted milk, malt vinegar, ...
in the production of Scotch whisky.
''Sphagnum
''Sphagnum'' is a genus of approximately 380 accepted species of mosses, commonly known as sphagnum moss, also bog moss and quacker moss (although that term is also sometimes used for peat). Accumulations of ''Sphagnum'' can store water, since ...
'' 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. In World War I
World War I or the First World War (28 July 1914 – 11 November 1918), also known as the Great War, was a World war, global conflict between two coalitions: the Allies of World War I, Allies (or Entente) and the Central Powers. Fighting to ...
, ''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.[ It is also claimed to have antibacterial properties. Native Americans were one of the peoples to use ''Sphagnum'' for diapers and menstrual pads, which is still done in ]Canada
Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
.
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
Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
/Greek
Greek may refer to:
Anything of, from, or related to Greece, a country in Southern Europe:
*Greeks, an ethnic group
*Greek language, a branch of the Indo-European language family
**Proto-Greek language, the assumed last common ancestor of all kno ...
name, which means "against fire".
In Mexico
Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...
, moss is used as a Christmas
Christmas is an annual festival commemorating Nativity of Jesus, the birth of Jesus Christ, observed primarily on December 25 as a Religion, religious and Culture, cultural celebration among billions of people Observance of Christmas by coun ...
decoration.
'' Physcomitrium patens'' is increasingly used in biotechnology
Biotechnology is a multidisciplinary field that involves the integration of natural sciences and Engineering Science, engineering sciences in order to achieve the application of organisms and parts thereof for products and services. Specialists ...
. Prominent examples are the identification of moss gene
In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...
s
with implications for crop
A crop is a plant that can be grown and harvested extensively for profit or subsistence. In other words, a crop is a plant or plant product that is grown for a specific purpose such as food, Fiber, fibre, or fuel.
When plants of the same spe ...
improvement or human health
Health has a variety of definitions, which have been used for different purposes over time. In general, it refers to physical and emotional well-being, especially that associated with normal functioning of the human body, absent of disease, p ...
and the safe production of complex biopharmaceuticals in the moss bioreactor
A bioreactor is any manufactured device or system that supports a biologically active environment. In one case, a bioreactor is a vessel in which a chemical reaction, chemical process is carried out which involves organisms or biochemistry, biochem ...
, developed by Ralf Reski and his co-workers.
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.
References
Further reading
* Kimmerer, Robin Wall (2003). ''Gathering Moss: A Natural and Cultural History of Mosses''. Oregon State University Press. .
External links
Information, diagrams and photos
Moss grower's handbook (2.3 9MB PDF file)
The British Bryological Society
World of Mosses – Watercolour paintings of moss by Robert Muma
{{Authority control
Extant Carboniferous first appearances
Mississippian first appearances