Review Article

Horticultural Importance of Bryophytes——A Review  

Afroz  Alam , Vinay Sharma
Department of Bioscience and biotechnology, Banasthali University, Rajasthan, India
Author    Correspondence author
International Journal of Horticulture, 2015, Vol. 5, No. 19   doi: 10.5376/ijh.2015.05.0019
Received: 10 Sep., 2015    Accepted: 20 Oct., 2015    Published: 04 Dec., 2015
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Afroz Alam, and Vinay Sharma, 2015, Horticultural importance of bryophytes-a review, International Journal of Horticulture, Vol.5, No.19 1-9 (doi: 10.5376/ijh.2015.05.0019)


Bryophytes, the first land plants hold an indispensable place in the plant kingdom as they have similarities with algae on one hand and pteridophytes on the other. Beside this they also have unique characteristics of their own. Nature makes them an imperative component of the ecosystem. All three classes of bryophytes have been used for various purposes since time immemorial. In this review their potential uses in horticultural industry have been discussed.

Bryophyta; Horticulture; Moss; Peat; Sphagnum

Bryophytes (liverworts, hornworts and mosses) can be found in every ecosystem in the world. They are also recognized as a group of non-vascular plants. They are called atracheate as they lack xylem and phloem, the conductive tissues of vascular plants. However, bryophytes and tracheophytes are monophyletic and together called embryophytes. As the name entails, there is an embryonic phase in these organisms as compared with their closest relative of green algae. All embryophytes have an alternation between sporophyte and gametophyte generations in the life cycle. Unlike vascular plants where the sporophyte is the dominant, obvious stage, bryophytes, on the other hand, have a conspicuous gametophyte stage (in most cases) that forms the green mats and tufts we see. The leaves of bryophytes are structurally very unusual from those of tracheophytes; they are generally one cell layer thick. Bryophytes show an immense diversity in growth form and habitat. There are bryophytes that can withstand extended periods of desiccation and others that are aquatic. They are one of the pioneers in ecological succession and one can find them on virtually every substrate including soil, leaves and bark of trees, rocks, decaying wood, and even cars and other man-made materials. Owing to their miniature size and little human uses they have remained a basically ignored group of plants.
On the basis of recent studies, there are 960 genera and 24,000 species of bryophytes exist in the world (Starr et al., 2009). In spite of the huge diversity of these plants, very few economic uses have been known. Even though bryophytes are considered first land plants, their importance is relatively unfamiliar. Bryophytes are used in the pharma industry, in horticulture, for domestic purposes, and are also environmentally imperative. The miscellaneous uses and applications of this miniature plant group are being progressively more recognized around the humankind. Their prospective in the horticulture is also colossal. In this article, the various uses of bryophytes in gardening or horticulture have been examined and discussed.
Various Uses of Bryophytes in Horticulture
In horticulture, position of mosses is almost incomparable in any other existing bryophyte industry (Nelson and Carpenter, 1965). Peat mosses have played a foremost task in horticulture for centuries (Perin, 1962; Arzeni, 1963; Adderley, 1965). Even if their application as part of the background beautification has conventionally been used by Asian, they have frequently been utilized as soil additives and bedding for hothouse crops, potted decorative plants, and seedling beds (Cox and Westing, 1963; Sjors, 1980).

Figure 1 Various uses of bryophytes 


In horticultural practice bryophytes are being used as soil additives since long time. Bryophytes are predominantly accepted soil additives because of their remarkably high water holding capability and permeability to atmospheric air. Peat is one of the important soil conditioners and is frequently used for various horticultural purposes around the world. Numerous taxa of bryophytes have also been used for greenhouse plants, potted decorative plants, and in garden soil.
In air layering, a method of propagating plants, use of Sphagnum is very common. Fresh plants of Sphagnum spp. are also mixed with the topsoil or put on the soil as a mulch. It helps to uphold wetness and prevent redundant growth of weeds. It is permeable to atmosphere and has flexibility, making it an ultimate growth medium. In Japan, use of mosses as ornamental plants for culture is very trendy. Landscape tray is another horticultural skill of Japan in which a number of mosses like Bartramia pomiformis, Leucobryum neilgherrense,and Polytrichum commune are used. Mosses also offer an imperative component for bonsai, where they help in the stabilization of soil and in preserving moisture. Moreover, the moss seems to be stylish and provides a green cooling feel. Moss gardens are also one of the aspects of daily life in Japan, where mosses have commonly been used in moss gardens as they bestow a calm splendor and olden gaze to gardens by covering tree trunks, rocks and stone. Buddhist temple in Kyoto, an ancient capital of Japan holds a much admired moss garden.
The great range of horticultural uses of bryophytes has been given in Figure 2.


Figure 2 Range of horticultural uses of bryophytes


Horticulture benefits from a extensive practice involving bryophytes as additives to the soil, soil cover, undersized plants, greenhouse crops, potted ornamental plants, and for seedling beds. Sphagnum (moss)is used in making totem poles to support climbers and moss-filled garlands, admired in southeastern U.S. Other pretty horticultural uses comprise floral arrangements, making flower baskets and covering flower container etc. Gardeners typically employ soggy Sphagnum for transportation of live plants. Occasional burning of Sphagnum is also used to create a smoke screen against frostiness (Glime, 2007).
Conditioning of Soil
Various moss taxa are often used to condition the soil. Mosses with coarse textured augment water-storage capacity, while smooth-textured mosses offer required air spaces. Mosses perk up the nutrient state by holding nutrients, particularly those originate by rainfall and dust, and discharging the nutrients bit by bit over a much longer phase of time than normal nutrient occurrence near the top soil. Typically mosses build up K, Mg, and Ca from precipitation, but they do not contend for P in soil. After the accumulation of these ensnared nutrients mosses release them slowly to the soil. When mosses become dried out, their cell membranes undergo damage, so when rehydrated again, it becomes permeable due to leaky situation. It normally takes about a daytime to fix this damage, and during that time, the moss can pour out its additional soluble contents, thus providing a number of these nutrients to root zone of other plant during the initial stages of precipitation. It is established that bryophytes augment the safeguard ability of the soil, predominantly in opposition to the alterations usually caused by adding up of manure. The dawdling decay of several bryophyte taxa makes them appropriate for continuing mulch. When Sphagnum is spread over the soil or mixed with top soil, it keeps dampness and averts the growth of unwanted weeds. It also dejects damping-off disease caused by fungi. Peat mosses mixed with wastes obtained after fish-processing supply a fertilizer better to sawdust and wood flakes in preserving nitrogen, however, it is also more costly (Ando, 1957; Bernier, 1992; Rao and Burns, 1990).
For growing exotic ornamental ferns, orchids (Adderley, 1965) and other bonsai, mosses are particularly excellent Taxa like Hypnum imponens, Leucobryum spp., Camptothecium arenarium, Rhytidiopsis robusta, and Thuidium delicatulum are specially used for this reason. Leucobryum spp. is used as a replacement for peat moss that encourages good root initiation on cuttings of orchids. For the enhanced and quality production of mushrooms, Sungrow, Inc., has had a contract with the Campbell (soup) corporation to grow mushrooms using a mixture of Sphagnum spp. Sphagnum appears to be indispensable in the method of air-layering. The moss is coupled firmly with sheet of plastic in the specific region of the stems to maintain wetness, encouraging the expansion of adventitious roots. Pant (1989) observed the use of such padding for grafting fruit trees in western Himalayas. He also reported more prolific bud and flower formation in Fuchsia and Begonia,if the pots have a moss layer to take apart the humus loaded top and the bed soil. Similar practice has also perform in Japan, where for Rhododendron cultivation, fragments of Hypnum plumaeforme, Leucobryum bowringii, L. neilgherrense, and rarely L. scabrum are mixed with sand or soil (Ando, 1957).
Seed Beds
Use of bryophytes as seed beds offers both benefits and troubles, because it may promote seed germination, but as a side effect also inhibits survival of seedlings. However, species of Polytrichum are used as a carpet to promote germination pioneering white spruce (Picea glauca) in Nova Scotia. Conversely, a mixed mat of Polytrichum and Cladonia is too intense for Populus seed spread. As a result, germination is failed because the lichen and moss mat absorbs water too fast to permit satisfactory soaking of seeds, beside this, recurrent dripping and desiccation of surface soil causes the hardly any thriving seedlings to lift. Actually, moss has been well thought-out as a “pest” in the case of containers growing of conifer seedlings, because it obstructs juvenile seedlings by creating a contest for minerals nutrients, and withdraws soil of water. One of the problems seems to be that, in water deficit soils, Sphagnum peat has a high attraction for water, providing deprived hydraulic conductance for seedlings and shoot water potentials are poorer than those obtained in sandy loam (Al-Kanani et al., 1992; Bernier et al., 1995).
In Picea mariana, development of roots is pretty slow. Roots are too undersized to get into the soil ahead of the moss to attain nourishment (Al-Kanani et al., 1992; Bernier et al., 1995). Conversely, in case of prairie soils, cryptogamic crusts augment the establishment of seedling. In such conditions, Sphagnum extracts persuade germination of Pinus banksiana seeds, and extracts of Polytrichum commune and Sphagnum spp. encourage growth of Larix seedlings. Extracts of the above mentioned moss taxa, on the other hand, inhibit the growth of other pine and spruce seedlings. Some of this control of germination may be due to the production of IAA by these mosses, but under usual conditions, it is uncertain if this phytohormone would influence other plants. Nevertheless, when extracts of mosses are applied, conflicting responses are observed with different plant species. Bigger moss taxa, for instance, Pleurozium schreberi supports seed germination in coniferous plants, but the seedlings hardly ever carry on to next year. This appears to be the outcome of diminutive seedling height that makes them unfeasible to compete with superior mosses for light, or they germinate in the mat too distant over the soil and are incapable to acquire adequate nutrients and water by their roots. In contrast, it was established that amplified bryophyte envelop reduces thriving seedling coming out, but amplified seedling survival. The low per/Pfr light ratio beneath the cover of bryophyte lessens successful emergence. While it has also recommended that chances of survival, perhaps improved by discharge of nutrients from bryoflora during hot and humid weather conditions. But there are fewer uncertain sensation accounts about the use of mosses as ‘seed bed’. Some reports suggest that spreading of Rhododendron ponticum in Ireland is basically caused by an augment in bryophyte cover as a consequence of excessive-grazing. In this instance, moss flora supply crucial wetness required for successful germination, and seedlings are not consumed because of the un-palatability of mosses that provide protected situates to the seeds. In Dutch chalk grasslands, more or less comparable safeguard has been experimental. It was observed that Calluna vulgaris grew better with more flowering when it took set in moss beds. However, as mentioned above, strong retardation on the germination process was observed. Bogdanov (1963) believed that bryoflora is actually a serious problem for germination in some regions and the elimination of moss cover is necessary. Later on, it was that found moss extracts have no inhibitory outcome on seeds of Calluna vulgaris, however, they found liverworts behave as inhibitors due to their several sesquiterpenoids (Matsuo et al., 1983; Miller and Trigoboff, 2001; Stark, 2002).
Moss Gardens
Japanese are very fanatical of gardens and scenic attractiveness. They use mosses to craft a sensation of calmness in their backyards and lawns. They love an orderly gaze of shades of green instead of the blend of variously coloured grasses. Moreover, moss gardens habitually connected with Buddhist places of worship, the most prominent of which is Kyoto’s Kokedera, exactly decoded as “moss temple.” One splendid example of use of mosses in garden is the Sanboin Temple, Kyoto, where two guitar-shaped and three circular patches of moss taxa signify the 1598 cherry blossom feast of Lord Hideyoshi Toyotomi (Glime and Saxena, 1990). For gardens the most preferred moss taxa are Pogonatum and Polytrichum. Other frequently used widespread species are Dicranum scoparium, Leucobryum bowringii, L. neilgherrense, Rhizogonium dozyanum, and Trachycystis microphylla, these species grow in piles or cushions, forming a tender, undulating background akin to mini hills. Besides Japan, other European countries, including United Kingdom using mosses for their gardens to form a serene ambiance. Mosses like Neckera crispa, Plagiomnium undulatum, Dicranella heteromalla, Hylocomium splendens, Polytrichum commune, P. piliferum, Dicranum scoparium, Rhizomnium punctatum, and Thamnobryum alopecurum are mostly used for this purpose. Ando (1957) once highlighted the use of Polytrichum commune in the built-up garden of Poet Laureate W. Wordsworth.
Moss Industry
Physically Sphagnum is highly permeable and porous and it has been found to soak up metals, and thus, peat moss is used as an efficient absorption and filtering means for the treatment of throwing away water and waste matter of industrial units with the acidic and noxious release containing heavy metals (Cd, Cu, Ag, Hg, Fe, Sb and Pb), and untreated organic materials for instance detergents, oils, dyes and microorganisms. For developing nations utilization of Sphagnum in this task emerges cost-effective. It entails contact of peat with throw away water with subsequent exposure to air for drying the peat and lastly burning the plants and recovery of the metals. Sphagnum can also be used as an efficient adsorption material for oil trickles. Sphagnum is also used for the production of activated carbon, an important adaptive material used in many chemical industries. Several diverse premium products are now commercially being formed to serve purchaser requirements in the field of bioremediation and pollution management. These are used for the bio-treatment of oil spills in swamps, mangroves, and marshland. Usual absorbents work gradually when applied to a spill out and may afterward liberate the oil, defying controlled standard. Products of bryophytic origin can take up liquid up to 12 times of their weight, and need a lesser storage space than conventionally used resources. Shipping and management of bryophyte-derived products are also easier, and disposal costs are lower than conventional products. Some of these bryophyte derivative materials does not absorb water, i.e. ‘hydrophobic’ and can absorb oil (oleophilic), makes them particularly useful for oil spill outs on water stream. In addition, benzene, diesel fuels, toluene, gasoline, ethyl ether, ethyl benzene, oil base ink, kerosene, jet fuels, butanol, methanol, chloroform, tetrachloroethane, xylenes, carbon  tetracholoride, paraffin oils, styrene, motor oils, oil based paint, cutting oils, corn oils, acetone, vinyl acetate etc., can be absorbed by the bryophyte-derived products (Chen  et al., 1992; Chen and Chang, 2001a,b).
Peat moss is an outstanding stuff for delivery of fresh vegetables, plants and flowers. Sphagnum is also in use for hydrophonics crop growing, and for commercially important bulbs and roots storage. Sphagnum has remarkable insulation property, hence also used in the buildup of insulator expanse for dwelling. In France moss industries construct carpets of moss in a range of sizes. They are uncomplicated to fix along the desired area. The saleable production of peat moss has been running for over 150 years in the United States of America. In Asia, Sri Lanka manufactures an extensive array of environment-friendly products like cocopeat (coir fiber pith ), coir pots, hanging wire baskets, basket liners, moss sticks etc., by the use of different species  of bryophytes (Glime, 2007).
Air Layering
Nature has always been a teacher to horticulturists, and invariably provides imperative instructions. in natural conditions, bryophytes especially mosses endow with appropriate media for practice of air layering of higher plants like Calluna  spp. and yet various tropical trees, hence obviously they are used more or less fully for air layering as a propagation method of selected plants. The moss is enveloped all over the area where roots are to be promoted, frequently held in place with fabric mesh, wire, or gloomy plastic. Several moss taxa offers a unremitting supply of moisture and promotes the expansion of adventitious roots while checking the growth of fungi. Once the plant attains appropriate maturity with formation of sufficient roots, the stalk can be cut underneath that point and the explant can be used to grow into a new plant (Scandrett and Gimingham, 1991; Macdonald et al., 1995). In western Himalayan regions of India, Pant (1989) reports comparable utilization of mosses for grafting many fruit trees.
Pot Culture
In potted plant culture, mosses are known to promote better growth. Pant (1989) highlighted few examples where mosses are utilized to take apart the humus-rich topsoil from the bottom soil, that promotes more profusely bud and flower formation in case of Fuchsia and Begonia in pots. Family Ericaceae, especially, benefit from the vital acid production of Sphagnum and other mosses, like Hypnum plumaeforme, Leucobryum bowringii, L. neilgherrense, and L. scabrum. The fragments of these mosses are used, mixed with soil or sand, to cultivate Rhododendron shrubs (Ando, 1957).
Sphagnum is vital for culturing juvenile seedlings in forestry. Heiskanen and Rikala (2000) established that use of Sphagnum (peat) is superior compared to peat with perlite or fine sand, the former resulting in more stronger seedlings as a consequence of the higher water retention of the medium. Besides this, Sphagnum is also supportive and effective in the inhibition of plant pathogens (Miller, 1981). However, in case of seedlings of coniferous plants, use of mosses is not recommended because they can compete for nutrients, choke young seedlings, and repel water (Haglund et al., 1981)
Mushrooms & Other Fungi
For cultivation of mushroom (Agaricus bisporus), Sphagnum peat is the substrate of preference as sheathing/casing medium (Beyer, 1997). Companies like Sungrow and Campbell had a multi-million-dollar contract for the improvement in mushroom culturing using a Sphagnum mix (Miller, 1981). However, in those places where there is no peat, certain substitutes are necessary for mushroom culture. The demand for peat replacements led to analysis other substrata and evaluate, but peat gave the maximum yields compared to 8 other resources (Eicker and Greuning, 1989).
In an effort to make additional enhancement in mushroom accomplishment, Beyer (1997) required ways to diminish the consequence of built up substances on overdue crops of mushroom. Unexpectedly, he noted that the adding up of Hypnum peat to the manure enhanced afterward break yield, but the adding up of Sphagnum did not. One of the apprehensions is that the peat get contaminated with nematodes and may hold Pseudomonas tolaasii, the source of bacterial blotch, both of which are known to cause severe ailments to the mushrooms (Nikandrow et al., 1982). Martin and Bailey (1983) successful in using peat as a fermentation medium in which adapted fungi could be grown. The rate of their success was more in case of Agaricus campestris than Morchella esculenta (Smith, 1983) according to them the growth inhibitors might be exist in peat. Utilizing hydrolysates of sulfuric acid with autoclaved peat released a liquor that, when added-on with nutrients, would improve the growth and content of crude protein of these two palatable fungi.
A blend of Sphagnum with fish waste assures to be a proper substrate for culturing the fungus Scytalidium acidophilum (acid-tolerant fungus), this fungus is supposed to be a capable source of microbial protein (Martin and Chintalapati, 1990). However, not all fungal cultures appear to promoted from these mixtures. In one profitable process, the yield of mushrooms enhanced when the peat was absent in the medium for cultivation (Smith, 1983).
Container Gardens
Mosses are frequently used in container gardens with bonsai, because they are very helpful for the stabilization of the soil and retainment of wetness for the superficial roots of bonsai. Dryness of the mosses is an indicator of water need for bonsai. Though, they are not forever friendly to the bonsai. It has been studied that the uninterrupted dampness of the mosses can slow up root growth and encourages abrupt fungal assaults. Therefore, periodical removal of mosses is essential to diminish damage caused by fungus (Bland, 1971; Ishikawa, 1974).
Stylish horticulturists choose specific moss taxa to provide exacting tasks in the container landscapes. Moss taxa like Atrichum, Climacium, Dicranum, Polytrichum, and Rhodobryum which are large, upright are used to replicate forests. Bryum argenteum has a silvery, compact gaze that can appears as grasslands, and Leucobryum usually has the character of a mount. Racomitrium canescens provides a frosted look which is well suited to obtain the appearance of snow-capped peaks. Physcomitrium, frequently assists to improve the gaze of flower pots in greenhouse. Leucobryum spp. due to its miniature size appear like grass, hence suitable for landscaping. Whenever, intermediate sized mosses are needed mosses like Barbula unguiculata, Funaria hygrometrica, and Weissia controversa can fulfill the purpose. A number of moss taxa are also utilized for making false bonsai, for instance, Campylopus spp., Dendropogonella rufescens, Hypnum spp., and Thuidium spp. are commonly used in such practice in Mexico.  Similarly, in North America (the Pacific Northwest), Leptobryum pyriforme, commonly known as ‘Kyoto moss’, is sold for making bonsai trays (Glime, 2007).
Minuscule plated landscapes employ mosses to give suitable feel and color with modest hazard of spoil due to drying (Kawamoto, 1980; Oishi, 1981). In an interesting example, Gerritson (1928) positioned 16 moss taxa ('platter of mosses') in different stages of development to present a shifting landscape for his hospitalized friend and said, "Each day the mosses had changed appearance; so each day added a new joy”.
For constructing these minuscule sceneries, Schenk (1997) advocates the customary potting mix of manure, including peat moss, tree bark (ground-up), or decomposed sawdust. He concerns that vermiculite, sand, or perlite can be used, but continuous dampness is essential because they tend to have bigger spaces and dried out rapidly close to the surface, leaving the moss without any moisture source.
All these practices are although apparently harmless, but some of the moss taxa can act as allergens hence caution must be taken. For instance, Sphagnum peat as a medium for tray gardens, dish gardens and other forms of bonsai and is documented for its capacity to anchorage the sporotrichosis causing fungal strains (Dong et al., 1995).
Hanging Baskets
The use of mosses in the creation of hanging flower baskets is also common (Smith, 1996). In the USA, moss strips (Hypnum, Polytrichum and Sphagnum)of few centimeters wide are regularly used to construct hanging flower baskets. More often than not a cable framework is used to provide hold up, these frameworks of wires are laden with moss taxa that enhances the beauty also. As a result, not only they appear as an eye-catching good-looking basket, but also they lessen the requirement for recurrent watering (Lohr and Pearson-Mims, 2001).
Similar to the container gardens, the terrarium is a drier plant version of the aquarium. A terrarium is frequently set like an enclosed minuscule garden. Owing to its minute size, bryophytes are regularly used to give the appearance of hills. However, bryophytes are not straightforward to grow in such circumstances. If the container is completely open, mosses quickly dried up and become brittle. If it is completely closed, fungi can grow without difficulty. Hence it is recommended to leave the apex partly open to allow much required circulation of air.
Selection of moss taxa depends on the required moisture level and effect within the terrarium. Polytrichum spp. can endure in a fairly arid terrarium but will simply be sheltered with fungus when it is too humid. Similarly, Leucobryum spp.and Ceratodon purpureus prefer good air circulation, which is helpful to check the growth of the mold, biggest threat to the terrarium. Schenk (1997) states, "I must tell the whole truth by identifying the great enemy of terrarium gardening with native woodlanders, for there is one: mold." He warned about the short life of most terraria due to the problem caused by mold. Hence, moss taxa like, Funaria hygrometrica can be used in extra moist conditions, but it still wants proper air circulation. For proper air space and circulation the size of the container is an important parameter because small sized containers invariably contaminated with fungal growth. Charcoal may be further mixed with the substrate to soak up undue acidity and gases produced by gradual decomposition.
Moss taxa that are collected sodden normally do not require extra water and may even have to be dried up by leaving the terrarium open wide for a few hours. After that, modest watering is desirable within the nearly sealed container. Slightly dry mosses can be moistened with a few tablespoons of water, while completely dry mosses may need up to 1/4 cup of water (Schenk, 1997).  Upholding for the initial few days after planting is necessary to shun an instant mold assault. In the condition of intense dew on the container walls, container must be open to attain dryness. The life span of a terrarium can be increased by the use of fluorescent lights that shun the etiolated growth that occurs in diffuse light.
Peat mosses have been extensively used in various horticultural practices as mentioned above. The structure of bryophytes makes them well appropriate for bed linen, topiary foundation, hanging baskets and garlands. Their capacity to absorb moisture makes them the ultimate material for shipping of plants. They are used extensively in culture medium for the growth of exotic orchid. Sphagnum and other mosses are frequently used as top soil conditioners to provide the perfect medium for retainment of nutrients and their gradual release following drying. They offer good fertilizer, particularly when mixed with throw away products like sewage or fish offal. The cyanophycean flora of some peat mosses provides extra fixed nitrogen. Their antimicrobial activities check fungal growth along with the preservation of moisture. These remarkable properties well suited for air layering, good culture media and potting mixes. In forestry the role of peat is also significant as mosses are frequently used to culture juvenile seedlings. In food industry peat mosses are extensively used in mushroom and morels culture at commercial level. Diminutive moss taxa are used customarily and effectively in container gardens for instance, bonsai, where a variety of mosses are used to imitate diverse features of minuscule landscapes. Use of moss taxa in terraria is the most difficult practice due to the threat of fungal contamination; however, with proper aeration and vigilant selection of moss species this problem can be counteracted. Overall, in the field of horticulture, the use of bryophytes, especially mosses will remain obligatory.
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