GENERAL I ARTICLE

Uses of Bryophytes
D K Saxena and Hari nder
D K Saxena is at t he
De par t me nt of Bot any,
Barielly College, Bareilly.
Hi s current res earch
i nt eres t s are in t he
det oxi f i cat i on of met al s
by bryophytes, and t hei r
rel evance to
biomonitoring/
biomapping st udi es.
Hari nder i s at pres ent in
t he Department of PG
Di pl oma of Envi ronmen-
tal Management, Bareilly
College, Bareilly and has
be e n involved in res earch
on t he bi omoni t ori ng
pot ent i al of bryophyt es.
Keywords
Liverworts, mosses, bi o-i ndi ca-
t ors, ec o- f r i endl y pr oduct s,
medi ci nal plants.
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Al t ho ug h bryophyt es are a mo ng t he o l de s t l and pl ant s , t he i r
us e f ul ne s s i s rel at i vel y u n k n o wn to mo s t pe opl e . Bryo-
phyt e s are us e d i n pha r ma c e ut i c a l product s , i n hor t i c ul -
ture, for h o u s e h o l d purpos es , and are al s o e c o l o g i c a l l y
i mport ant . T h e mul t i f ar i ous us e s and a ppl i c a t i o ns o f t he
bryophyt e fl ora are be i ng i nc r e a s i ng l y r e c o g ni z e d around
t he worl d. Th e i r pot e nt i al i n t he b i o ma p p i n g o f at mo-
s phe r i c pr e c i pi t at i on i s al s o e no r mo us . In t hi s art i cl e, we
bri ef l y e x a mi n e s o me o f t he us e s o f t hi s a nc i e nt group o f
s i mpl e pl ant s .
The liverworts, hornwort s and mosses are small, l ow-growi ng
pl ant s t hat const i t ut e t he phyl um Bryophyt a. Bryophyt es have
nei t her true stems, leaves nor roots, nor a vascular system.
Consequent l y, t hey are confi ned to damp shaded areas wi t h hi gh
humi di t y, and f r equent rainfall. Most br yophyt es are ei t her
liverworts or mosses. Li verwort s grow hori zont al l y, and are
fl at t ened or ' leafy' , whereas mosses have an upr i ght stalk wi t h
spirally arranged leaf-like structures. The pl eurocarpous mosses
(carpet -formi ng) are charact eri zed by ext ensi ve branchi ng and
lateral sporophyt e placement, compared to t he terminal sporophy-
tes in acrocarpous (erect) mosses. Pl eur ocar pous mosses consti-
t ut e t he maj or groups of mosses. Though t i ny, br yophyt es modi f y
t hei r mi cro-cl i mat e and serve to conserve moi st ure, check soil
erosi on on hilly slopes, and also serve as a seed bed for forest
cover. They are now i ncreasi ngl y bei ng used for purposes rang-
ing from pol l ut i on moni t or i ng to new sources of pharmaceut i cal
product s. Present uses of bryophyt es i ncl ude ecology, hort i cul -
ture, const ruct i on, househol d uses, medi ci ne and even food.
E c o l o g i c a l U s e s
Li verwort s and mosses have been f ound to be good i ndi cat ors of
envi ronment al condi t i ons. The occurrence of certain aquat i c
RESONANCE I June 2004
GENERAL I ARTICLE
mosses can be used as an indicator of calcium and nutrient
content in water. Some bryophytes grow only in a narrow and
specific pH range and, therefore, their presence can be used as an
indicator of soil pH. Bryophytes envelope the forest floor and
tree trunks and aid in moisture conservation. Mosses like
Atrichum, Pogonatum, Trematodon, Pohlia, Nardia and Blasia play
a role as inhibitors of soil erosion due to their trample-resistant
structure and their regenerative ability. Acrocarpous mosses
like Rhodobryum and Dicranum prevent soil erosion of the slopes
of hills because the netted and webbed protonemata cover the
exposed substrata. In pastures of Nova Scotia in Canada, it has
been seen that white spruce germinates most prolifically in
carpets of Polytrichum. Many mosses, especiallyHypnum imponens,
provide excellent seeding beds for a variety of coniferous tree
species. This is likely to be due to the role of bryophytes in
providing moisture, appropriate temperature, and also organic
matter and minerals after the death of bryophytes. Thus, bryo-
phytes play an important role in the maintenance and replenish-
ment of forest cover.
Some bryophytes
grow only in a
narrow and
specific pH range
and, therefore,
their presence can
be used as an
indicator of soil pH.
Minerals supplied by rain, by leaching of the canopy, and by
through-fall are trapped by bryophytes. Thus, bryophytes play
an important role in retaining minerals that might otherwise be
quickly leached from the soil. Mosses easily accumulate K, Ca
and Mg from rainfall. Yet, at the same time, they do not compete
for phosphorous in the soil. Bryophytes also play an important
role in iron-ore deposition. Nitrogen is often a limiting nutrient
for plant growth, especially in cold regions of the northern
hemisphere. Even small contributions from biological nitrogen
fixation may therefore be of importance to ecosystems. Some
mosses (Sphagnum) and liverworts provide suitable substrates
for the biological fixation of nitrogen in association with
cyanobacteria. These cyanobacteria seem to behave symbioti-
cally in these plants. Indeed, such moss symbionts are respon-
sible for most of the nitrogen fixation in artic and sub-artic
ecosystems.
Some bryophyte species have been found to be closely associated
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G E N E R A L [ A R T I C L E
The suitability of
liverworts and
mosses as
bioindicators is
mainly due to their
simple thalloid or
one-cell thick
structure, lack of
cuticle or epidermis,
resulting in greater
absorption and
accumulation of
nutrients and
pollutants directly
from the atmosphere.
with particular mineral deposits. For example, Merceya,
Mielichhoferia elongata, and M. mielichhoferia are known as copper
mosses because they grow in copper rich soil. Such species can
be used as indicator plants.
Mosses are also good indicators of acid rain, because they lack a
protective epidermis and cuticle and, hence, are more suscep-
tible than the vascular plants. In general, studies have shown
that liverworts and mosses are very sensitive to air pollution. In
polluted areas, standard transplantation of certain mosses has
been found to be quite useful for monitoring the intensity and
trend of air pollution. By making regular survey and counting
abundance and frequency of species of mosses, an IAP (Index of
Atmospheric Purity) can be calculated. Bryophytes have also
been used to monitor airborne pollution caused by emissions
from various sources. Similarly, aquatic bryophytes A mblystegium
riparium, Fontinalis antipyretica, F. squamosa, Eurhynchium
riparioides, and Scapania undulata are now used to monitor water
pollution.
The suitability of liverworts and mosses as bioindicators is
mainly due to their simple thalloid or one-cell thick structure,
lack of cuticle or epidermis, resulting in greater absorption and
accumulation of nutrients and pollutants directly from the at-
mosphere. Moreover, mosses can be stored for several years
without noticeable deterioration and old specimens can easily
be chemically analysed. Several studies have indicated that
mosses accumulate heavy metals such as Pb, Fe, Zn and Ni in
high concentrations. Moreover, bryophytes retain their toxic
load for a long time after death because of slow release of
accumulated substances. Thus, they can serve as ' environmental
specimen banks' and are being used for this purpose.
Hort i cul t ural Us e s
There is a long tradition of use of bryophytes in horticulture as
soil additives, ornamental material for cultivation, and in Japa-
nese gardens (Figure 1). Bryophytes are particularly popular
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GENERAL t ARTICLE
because of their high water holding capacity and
permeability to air. Peat is an important soil con-
ditioner and is commonl y used for agricultural
and horticultural purposes around the world.
Bryophytes have also been used for green house
crops, potted ornamental plants and seedlings,
and in garden soil. Gardeners use Sphagnum in air
layering, a met hod of propagating plants. Fresh
Sphagnum is also mixed with the s6il or spread
over the ground as a mulch. It helps to maintain
moisture and prevent growth of weeds; it is per-
meable to air and has elasticity, making it an ideal growth
medium. Use of mosses in Japan as ornamental plants for culture
is very popular. Landscape tray is an alternative horticultural art
of Japan in which several mosses like Polytrichum commune,
Leucobryum neilgherrense andBartramia pomiformis are used. Mosses
also provide an important element for bonsai, help to stabilize
the soil and are useful in retaining moisture. In addition to this,
the moss looks graceful and gives a green cooling touch. In
Japan, mosses have frequently been used in moss gardens be-
cause they give a quiet beaut y and ancient look to gardens by
clothing tree trunks, rocks and stone. A very popular moss
garden is at the Buddhist temple in Kyoto, an ancient capital of
Japan.
Moss Industry
The physical structure of peat is highly absorbent and perme-
able and it has been found to absorb metals, and therefore,
Sphagnum is used as an effective filtering and adsorption agent
for the treatment of waste water and effluent of factories with
acid and toxic discharge containing heavy metals (Ag, Cu, Cd,
Hg, Fe, Sb and Pb), and organic substances such as oils, deter-
gents, dyes and micro-organisms. Use of peat moss in this role
appears economical for developing countries. It involves contact
of waste water with peat, drying the peat by mechanical pressure,
and finally burning the peat and reclaiming the metals. Peat can
also be used as an effective adsorption agent for oil spills. Active
RESONANCE J June 2004
Figure 1. Moss gardens in
Japan.
In Japan, mosses
have frequently been
used in moss
gardens because
they give a quiet
beauty and ancient
look to gardens by
clothing tree trunks,
rocks and stone. A
very popular moss
garden is at the
Buddhist temple in
Kyoto, an ancient
capital of Japan.
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GENERAL ]ARTICLE
The commercial
production of
Sphagnum, which
holds up to 30 times
its weight in water,
has been going on for
over 150 years in the
USA.
Figure 2. Moss carpets.
Courtesy: Michel Chiaffredo,
M/S Bryotech, France.
carbon is an important adsorptive substance in many chemical
industries and it can be produced from peat (Sphagnum). A
number of different high quality products are now commercially
being produced to serve client needs in the field of pollution
control and bioremediation. These products are used for the
biotreatment of oil spills in marshes, mangroves, and wetlands.
Conventional sorbents work slowly when applied to a spill and
may subsequently release the oil, violating regulatory leachate
standards. Some of the bryophyte-derived products can absorb
up to 12 times their weight, and require less storage space t han
conventionally used materials. Transportation and handling of
bryophyte-detived products are also easier, and disposal costs
are lower than conventional products. Some of these bryophyte-
derived products are ' hydrophobic' (do not absorb water) and
' oleophilic' (absorb oil), making t hem especially useful for oil
spills on waterways. Some of the other fluids that can be absor-
bed by these bryophyte-derived products are acetone, benzene,
butanol, carbon tetracholoride, chloroform, corn oils, cutting
oils, diesel fuels, ethyl benzene, ethyl ether, gasoline, jet fuels,
kerosene, methanol, motor oils, oil based paint, oil base ink,
paraffin oils, toluene, styrene, tetrachloroethane, xylenes, and
vinyl acetate.
The commercial production of Sphagnum, which holds up to 30
times its weight in water, has been going on for over 150 years in
the USA. Sphagnum is an excellent material for shi pment of
plants and fresh vegetables and flowers, for hydrophonics gar-
dening, and for storage of roots and bulbs. It
is also used in the manufacturing of insulator
sheets for houses. Moss industries in France
manufacture moss carpets in various sizes
(Figure 2). They are easy to fix along the
roads, lawns, play grounds, etc. In Sri Lanka,
a wide range ofeco-friendly products such as
coir pots, coir fiber pith (coco - peat), moss
sticks, hanging wire baskets and basket lin-
ers are made using bryophytes (Figure 3).
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RESONANCE I June 2004
GENERAL I ARTICLE
Bryophyt es as Fuel
Liverworts and mosses have long been tried and used as a fuel in
developed countries like Finland, Sweden, Ireland, West Ger-
many, Poland and Soviet Union. Peat is suitable for production
of low and intermediate BTU gas as well as hydrogen, ethylene,
natural gas, methanol and Fisher Tropsch gasoline. Peat mosses
are best suited for the production of methane, and peat is likely
to become an important source of fuel for production of heat,
methane, or electricity in the future. Peat has rapid regenera-
tion, can be easily harvested, has low sulfur content, and its
heating value is superior to that of wood.
Figure 3. Various products
made from bryophytes: (a)
hanging wire baskets (b)
moss sticks (c) basket lin-
ers (d) coir pots.
Reproduced with permission.
Source:
http://www, rex. industries @
lanka.com.ltd
Hous e Cons t ruct i on
In parts of the globe where woody plants are scarce and bryo-
phytes common, these tiny plants are used in the construction of
houses and their furnishings. At Kapkot in the Himalayas,
villagers use moss mats with shrubs, grasses, and bamboo to
make a pharki, a kind of door placed at the openings of their
temporary huts. Sphagnum peat has been developed as a new
construction material through the use of binders for solidifica-
tion and strengthening, resulting in new products like ' peatcrete'
In parts of the globe
where woody plants
are scarce and
bryophytes common,
these tiny plants are
used in the
construction of
houses and their
furnishings.
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GENERAL J ARTICLE
In the past, mosses
were used as
material for bedding,
packing, plugging
and stuffing owing to
their soft elastic
texture, and also
because they are not
easily attacked by
insects and micro-
organisms.
and ' peat wood' . Peat cret e is low cost, easy for sawi ng and nail-
ing, and can be cast and mol ded i nt o any shape. Thi s xnew
pr oduct is easy to carry to any dest i nat i on where t ransport at i on
is a pr obl em and can be used in bui l di ng material.
Hous ehol d Us e s
Mosses have been used for decorative purposes in Japan, En-
gland, France, Fi nl and and America. Or nament al wat er fl owers
are made f r om dri ed pl ant s of Climacium japonicum and sol d in
Japan. Pr obabl y the most useful househol d moss is Sphagnum,
largely because of its absorbent pr oper t y and as an insulator.
Mosses are woven i nt o mats and sol d in many parts of India. In
t he alpine hi ghl ands of t he Nor t hwest Hi mal ayas, Indi ans make
beddi ng, mat t resses, cushi ons and pi l l ows by st uffi ng mosses
into coarse l i nen sacks or by spreadi ng t hem on t he muddy floor.
The Hi mal ayans also use mosses as i nsect repel l ent s when stor-
ing food. Local mosses and l i verwort s are dried, made i nt o a
coarse powder t hat is spri nkl ed over grains and ot her goods to be
st ored in containers. Several i nsect ant i -feedant s have now been
f ound in di verse bryophyt es, maki ng t hi s group of pl ant s a
useful source of insecticides and i nsect repellents.
Sphagnum is now even bei ng used for cushi oni ng t he foot and
absorbi ng moi st ur e and odors in hi ki ng boots. In Ger many,
Sphagnum mi xed wi t h wool has been used to prepare a cheap
cloth. Women in t he villages of Kumaon, India, make head
cushi ons of mosses (Hylocomium, Hypnum, Trachypodopsis) to
carry vessels of wat er by stuffing t he mosses i nt o cl ot h sacks. I n
the past, mosses were used as mat eri al for beddi ng, packi ng,
plugging and st uffi ng owi ng to t hei r soft elastic texture, and also
because t hey are n o t easily at t acked by insects and mi cro-
organisms. In India, Sphagnum, Hypnum cupressiforme, Macro-
thamnium submacrocarpum, Neckera crenulata, Trachypodopsis cris-
panda and Thuidium tamarisceUum are used for packi ng of appl es
and pl ums in t he west ern Himalayas. Nur ser ymen in Indi a oft en
use wet Sphagnum for sendi ng or suppl yi ng live plants.
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G E N E R A L I A R T I C L E
Me d i c i n a l U s e s
The Chi nese and t he nat i ve Ameri cans have used vari ous moss
speci es like Philonotis, Bryum, Mnium, cr ushed i nt o a ki nd of
past e and appl i ed as a poultice. In India, t he bur ned ash of
mosses mi xed wi t h fat and honey is used as an oi nt ment for cuts,
bur ns and wounds in t he Hi mal ayan region. The l i verwort
Marchantia polymorpha is also used as a medi ci ne for boils and
abscesses, perhaps because t he young archegoni ophore resembl es
a boi l when it fi rst emerges from t he-t hal l us. The rosette-
f or mi ng Riccia spp., are used as an external appl i cat i on to cure
ri ngworm. Chi nese t radi t i onal medi ci ne names 40 ki nds of
br yophyt es t hat have been used to treat illnesses of t he cardio-
vascul ar syst em, tonsillitis, bronchi t i s, t ympani t i s, cystitis, as
well as ski n diseases and burns. It has been shown t hat an extract
ofRhodobryum giganteum can increase aorta bl ood t ransi t by upt o
30% i n animals. Tr ansgeni c PhyscomitreUa are now bei ng used to
pr oduc e ' bl ood- c l ot t i ng f act or I X' , for t he t r e a t me nt of
' haemophi l i a' B. Preparat i ons of calcined peat have l ong been
regarded as effective and cheap germicides; peat wat er possesses
ast ri ngent and ant i sept i c properties. ' Sphagnol ' , a distillate of
Peat Tar, is useful in eczema, psoriasis, pruri t us, hemorrhoi ds,
chi l bl ai ns, scabies, acne and ot her forms of ski n diseases, and is
also beneficial for allaying i rri t at i on arising f r om i nsect bites.
One of t he features t hat hel ps bryophyt es survi ve and mai nt ai n
t hei r place in t oday' s flora is t hei r cont ent of bi ol ogi cal l y active
compounds. These prot ect t he ot herwi se delicate pl ant s not onl y
f r om fungi and ot her mi croorgani sms but also f r om insects and
slugs. I n a way, t hese bi ochemi cal compounds make up for t he
lack of a t hi ck cut i cl e and bark in bryophyt es.
Li ver wor t s are not suscept i bl e to fungal diseases, and ant i mi cro-
bial act i vi t y has been seen in extracts ofReboulia andPallaviciania,
possi bl y due to l unul ari c acid, f ound in l i verwort s but not in
mosses. A few moss genera like Atrichum, Dicranum, Minium,
Polytrichum and Sphagnum also possess ant i bi ot i cal l y active sub-
stances. It was f ound t hat a met hanol extract f r om H. aduncus
Li verwort s are not
suscept i bl e to
fungal di seases,
and ant i mi crobi al
act i vi t y has been
seen in ext r act s of
Reboulia and
Pallaviciania,
possi bl y due to
l unul ari c aci d,
f ound in l i verwort s
but not in mosses.
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GENERAL I ARTICLE
Fi gure 4. Ant i mi cr obi al ac.
t i vi t i es of bryophyt es: Ef.
f ect of enti re l i verwort ex-
t ract on tomato pl ant i n-
f ect ed wi t h Phyt opt hor a
i nf est ans; (a) treated wi t h
ext ract (b) wi t hout extract.
Figures 4 and 5 reproduced with
permission.
Source:www.bryologie.uni-bonn.
de/english/contentGB/Aktuelles
GB/fungizidgb.htm
Fi gure 5. Ant i . f eedant eff-
ect of bryophyt es. (a) Leaf
t reat ed wi t h f i verwort ex-
t ract r emai ned unt ouched,
whereas (b) the unt reat ed
(control ) l eaf was devoured
compl et el y by a sl ug.
64
inhibited the growth of pathogenic fungi Botrytis cinerea, Rhizoc-
tonia solani and Pythium debaryanum, whereas petroleum-ether
extracts of Barbula and Timmiella species were found to be active
agai nst bot h gram-posi t i ve and gr am- negat i ve bact eri a.
Plagiochila stevensoniana proved to inhibit dermatophytic organ-
isms like Trichophyton mentagrophytes, Candida albicans and Ba-
cillus subtilis. This antibiotic activity mi ght be attributed to the
presence of non-ionized organic acids and polyphenolic com-
pounds. In the case of several bryophyte species, the active
ingredients responsible for anti-microbial effects have been
identified, e.g. Polygodial from Porella, Norpiguisone from
Conocephalum conicum, and Lunul ari n from Lunularia cruciata.
Green-house tests have shown that plants (tomato, pepper, cu-
cumber, wheat) treated with liverwort extract were distinctly
less affected (if at all) by fungus infections byPhytophtora infestans
than untreated plants (Figure 4). However, as yet, these extracts
have not actually been applied in agricultural settings.
Many bryophyte extracts act like stomach poison on animal
pests. The slug Arion lusitanicus was offered lettuce with liver-
wort extract and with pure solvent as con-
trol. The untreated (control) leaf was de-
voured completely (Figure 5b) whereas the
leaf treated with liverwort extract remai ned
untouched (Figure 5a). This leads us to the
conclusion that liverwort extract is not only
a potent fungicide and bactericide, but also
a weak biocide (stomach poison) against
animal pests.
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GENERAL J ARTICLE
Ant i -l eukemi c activity has also been demonst rat ed in several
compounds from leafy liverworts. A new ent eudesmanol i de,
di pl ophyl l i n, was isolated from Diplophyllin albicans and D.
taxifolium. Di pl ophyl l i n, havi ng an al pha-met hyl ene lactone
uni t , showed significant activity (ED50 4-6/~g/ml) against hu-
man epi der moi d carci noma (KB cell culture). Its optical enanti-
omer, derived chemically from t he compound isoanantolactone,
exhi bi t ed anticancer activity (ED50 20-37/~g/ml). Compounds
from Plagiochila fasciculata (New Zealand) seemed to i nhi bi t
P388 cells (leukaemia).
Dur i ng t he first world war, t he Germans used Sphagnum exten-
sively for dressing wounds. Sphagnum pads are superior to cot-
t on for dressing wounds as t hey absorb liquids 3-4 times as much
as cot t on, renderi ng frequent change of dressing unnecessary.
Moreover, t hey are economical, cooler, softer and less irritating
to t he ski n t han cotton. A pad of Sphagnum moss absorbs t he
discharge in lateral directions, as well as i mmedi at el y above t he
wound, and holds it unt i l fully saturated in all parts of t he
dressing before allowing any to escape. For t he preparation of
t he dressing, moss after being dried is carefully pi cked over and
put up loosely in small, flat t hi n musl i n bags (2 oz. of t he moss to
each bag, 10 inches by 14 inches).
Suggested Reading
[1] D K Saxena, inProceedings of XIIIInternationalBotanical Congress
Abstracts, 1981, p. 292, Sydney.
[2] J M Gl i me and D K Saxena, Uses of Bryophytes, Today and Tomorrow
Pr i nt er s and Publishers, New Delhi, India, 1990.
[3] D K Saxena and A Saxena, Role of plant tissue culture in biodiversity
conservation and economic development, Gyanodaya Prakashan, Nai ni t al ,
I ndi a, pp. 605-621, 2002.
[4] G Pant and S D Tewari, Bryologist., Voi.92, pp.120-122, 1989.
[5] G Pant , Indian J. Forestry, Vol.4, No.3, pp. 186-190, 1981.
[6] K G Gupt a and B Singh, Res. Bull. Punjab Univ., Vol.22, pp. 237-239,
1971.
[7] H Ando, BullBiol. Soc. Hiroshima Univ., Vol.7, No. 7, pp. 23-26,1957.
[8] H Ding, Medical spore-bearingplants of China, Shanghai, p. 499, 1982.
Address for Correspondence
D K Saxena and Horinder
Deparlmenl of Botany
Bareilly College
Bareilly 243 005, UP, Indio.
Email: dksoxena@hotmail.com
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