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Studies on the Bryopsida of the Kolli Hills

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Chapter 3

REVIEW OF LITERATURE

. .

Interest in Bryophytes has undergone a resurgence in the last

decade. This renewed focus on the mosses, liverworts and hornworts has

converged from diverse quarters within the scientific community. With

recent advances in technology and analytical approaches to phylogeny

research, systematists have made unprecedented progress towards

reconstructing ‘tree of life’. One of the truly monumental events in the

history of life was the origin of land plants, or Embryophytes. The

bryophytes have long been considered a pivotal group positioned at or near

the base of the embryophytes and a great deal of molecular work has

recently been aimed at resolving relationships among the disparate groups

of bryophytes, and their relationship to the tracheophyte clade. At the same

time, the utility of bryophytes, especially mosses, for analysis of plant

function and development has been increasingly appreciated and

capitalized upon.

In the first part of this chapter, the present state of the art regarding

the studies on bryophytes, especially moss flora in the Indian subcontinent

will be presented, the second part will deal with antimicrobial studies on

bryophytes, and the third part phytochemical analysis will be presented.

Chapter 3 Review of Literature

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3.1 STUDIES ON BRYOLOGY

This part presents comprehensively an account on bryophytes

worldwide followed by a review of studies in India.

The first ever work dealing exclusively with taxonomy of bryophytes

was by Dillenius’ Historia Muscorum (1741), which also included algae,

lichens, lycopods, etc. Linnaeus (1753) in his Species Plantarum, which is

considered as the early study on the Bryophytes especially for Sphagnum

included some Indian mosses also. Hedwig’s (1801) Species Muscorum is

considered for the application of the principle of the priority in case of other

mosses. Mitteni’s (1869) Musci Austro-Americani and Spreuce’s (1884-

1886) Hepaticae Amazonicae et Andinae, formed the foundation for

American Bryology. During the early part of the present century, bryophytes

collected mainly by missionaries, from different parts of the world including

the tropics, were studied in Europe by Mitteni (1869), Mueller (1905-1916)

and others but they had never seen these plants in living condition.

Jussieu (1836) used the term ‘mosses’ to include true mosses.

However the remaining groups was not known at that time. It was Brown

(1866) who introduced the term ‘Bryophyta’ a Greek word derived by

combining ‘Bryon’ = moss, ‘phyton’ = plant and treated under

‘Acotyledonae’ which included algae, fungi, lichen and mosses, but not the

liverworts. Eichler (1883) divided Bryophytes into two classes, the

Hepaticae to include all liverworts and Musci to include all mosses. Engler

(1892) subdivided each of the two classes into three orders. Hepaticae was


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divided into Marchantiales, Jungermanniales and Anthocerotales and Musci

into Sphagnales, Andreales and Bryales. The anomalous position of

Anthocerotales as an order of the class Hepaticae was pointed out by many

investigators very early itself. However, it was Howe (1899) who gave the

class status to the order Anthocerotales. He named it Anthocerotes and

divided the bryophytes into three classes viz., Hepaticae, Anthocerotes and

Musci. Many eminent Hepaticologists such as Smith (1955) and Schuster

(1958, 1966) have accepted the system of Howe (1899) with a modification

of the term Anthocerotes as Anthocerotae.

The studies on the mosses of India is said to have commenced when

the road to Nepal was opened. The first step towards our knowledge of the

moss flora of this region was done by Buchanan Hamilton, a Medical

Officer in British Embassy in 1802, who first collected mosses from Nepal.

The first paper on the mosses of this region Musci Nepalensis was

published in 1808 by Hooker, who later (1818-1820) described several

species for the first time. This was followed by two papers on Indian

muscicology jointly published by Hooker and Greville (1825).

Schwaegrichen (1811-1842) included some Indian mosses in his

supplemented to Hedwig’s Species Muscorum. Royle (1839) reproduced

the catalogue of Wallich, which contains a list of 113 species of mosses

mainly from Nepal, of which 55 species listed by Royle himself from

Mussoorie (Western Himalayas). This also includes a brief note on hepatics

and liverworts. Wallich (1841) enumerated 148 mosses collected in East


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Indies, with reference to the figures of new species published in Hooker’s

Icones Plantarum. Montagne (1842a & b) published an account of the

mosses from the Nilgiris. Subsequently Griffith (1842, 1843) published an

account of a number of mosses and liverworts from Khasia hills in

Meghalaya. Collections of liverworts from the Himalayas by Wallich and

Wight were also included in the Synopsis Hepaticarum of Gottsche et al.

(1844, 1847).

The first noteworthy contribution to Indian bryology in its early days

can be found in Griffith’s (1849a & b) Notulae and Plantae Asiaticae. The

Icones Plantarum Asiaticarum, which was based on his collections during

1835-1838, was published posthumously. His discovery of the remarkable

monotypic liverwort Monoselenium tenerum Griff. attracted attention all

over the world. Mitteni and Wilson (1857) enumerated the mosses collected

by J. D. Hooker and Thomson from India. Mitteni’s (1859) Musci Indiae

Orientalis was the most important comprehensive work embodying all the

Indian species known at that time. He recorded nearly 800 species

belonging to 85 genera and 19 families including a large number of

novelties. Later Mitteni (1860, 1861) published Hepaticae Indiae Orientalis

a comprehensive treatment of Indian liverworts, which included the

diagnoses of several new plants, and a detailed list of all Indian liverworts

known till date. This work was mainly based on collections by Hooker from

the Himalayas, the Khasia hills, by Hooker and Thomson and from Sri

Lanka by Gardner and Thwaites respectively.


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Mueller (1853, 1854a & b, 1869-70, 1871, 1878) brought out several

articles on the Indian mosses. Hampe (1872-73) described some Sri

Lankan mosses based on Beccari’s collection. Mitteni (1873) published an

account of the mosses of this Island collected by Thwaites describing a

number of new species under the joint authorship of Thwaites and Mitteni.

Reference to mosses from India can also be seen in the publications of

Jaeger (1870-1879). In the post-Mitteni period much progress has been

made through direct and indirect contributions. Brotherus (1898) described

a few mosses from North Western Himalayas. Subsequently in 1899 he

published an account of 96 species of mosses from Coorg in the Western

Ghats including 20 novelties. Schiffner in 1898 brought out an exhaustive

list of species recorded till then from East India.

Stephani’s Species Hepaticarum (1906-1924) in six volumes, a world

monograph of liverworts, included description of a large number of taxa

from India, Myanmar, Sri Lanka and adjoining Indian territories such as

Nepal, Madura and the Andaman and Nicobar Islands. Even though the

status of the plants included in this treatment were questioned by the

modern bryologists, this work still remains as a valuable reference to the

present workers. Other noteworthy contributions in this period were those

of Mueller’s (1901) “Scapaniae Indiae Orientalis” and Gola’s (1914) on

Kashmir Hepatics. Contributions by Dixon (1910, 1930, 1942), Bruehl

(1931), Bartram (1955, 1960) and Robinson (1956) on Indian hepatics also

added further information.


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3.2 STUDIES ON MOSSES

The floristic study of the mosses, so richly distributed in the Indian

subcontinent has received little attention throughout the past probably on

account of non-availability of literature. Even though the Hepaticae

received attention of Indian bryologists since Kashyap (1914), researchers

on Indian mosses began only after a decade or two. However, contribution

of scientists from abroad is continued even during the early half of this

century.

During the first half of this century Dixon’s contribution is notable. He

has published a number of new species and genera. But most of his

contributions were based on collections made by others from different parts

of the country. Dixon (1909a) described the species Brachymenium

turgidum collected by L. J. Sedgewick from Lonavala in the Western Ghats.

It was followed by the publication of a very valuable account of the mosses

from the Western Ghats collected by G. B. Savery and sent to him by L. J.

Sedgewick from various parts of the southern region of Bombay Presidency

(Dixon, 1909b), this included two new species viz., Pterobryopsis maxwelli

Cardot and Dixon and P. kavarensis Cardot & Dixon. In 1910, Dixon

described the new genus Merceyopsis Broth. & Dixon, closely related to

Merceya in the arrangement of cells in the nerve of the leaf. Sedgewick

(1910, 1911, 1913) compiled a list of 71 mosses collected by him, Maxvel,

Kirtikar and Woodrow from Mahabaleswar, Kanara, Panchgani and

Purandhar in Western India and identified by Dixon. In 1911, Dixon


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proposed a new genus Hyphilopsis, based on a collection by Sedgewick

from Purandhar and also described Orthotrichum griffithii Mitt. & Dixon,

Brachymenium fischeri and Bryum sachyadense. In 1912, the new genus

Bryosedgewickia of Entodontaceae from Purandhar was proposed by Dixon

and described three new species each of Pogonatum, Forsstroemia and

Lindbergia alongwith a list of many mosses from various parts of India. He

in 1914a enumerated 42 species belonging to 34 genera of mosses from

Abor hills of Assam. Dixon (1914) reported 58 species of mosses including

40 genera collected by C.E.C. Fischer and others from South India and

Ceylon, which include three new species viz., Campylopus pseudogracilis

Cardot & Dixon, Taxithelium vivicolor Broth. & Dixon, Barbella questei

Cardot & Dixon, and two new varieties viz., Stereophyllum ligulatum

(C. Muell.) A. Jaeger var. sedgwickii Broth. & Dixon and Levierella

fabroniaceae var. dilatinerve Cardot & Dixon.

In 1921, Dixon assigned Sedgewick’s collections from North Kanara

into 43 species of mosses under 27 genera. During 1922, he proposed

seven new genera of mosses of which Beddomiella funarioides Dixon was

from the Nilgiri hills. Dixon (1925) described Grimmia somervellii Dixon

from the Himalayas based on the collections made by T. H. Somervell, and

later (Dixon, 1928) proposed a new genus Homaliopsis from Palni hills,

collected by Foreau. In 1931 Dixon published six new genera, of which

Nanothecium a member of Entodontaceae was collected by Foreau in 1929

from the Kannan Devan Hills in Travancore, Orthotheciadelphus


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(Entodontaceae) collected from North-west Himalayas by R. L. Bardhwar in

July 1928 and Hydrocryphae (Cryphaceae) from Abor hills in Assam,

collected by F. Kingdon ward. In 1937, Dixon reported 208 species of

mosses of Naga hills in Assam based on the collection made by Dr. N. L.

Bor between 1933-1936 which include one new genus and 40 new species.

Dixon and Badhwar (1938) described 17 new mosses from North-west

Himalayas. In 1942, Dixon instituted 16 new species of acrocarpous

mosses from North-western Himalayas.

Brotherus (1909-1910) described a new genus Hageniella Broth. with

H. sikkimensis Broth. from Sikkim and later he (Brotherus, 1928) provided

an account of 106 species and 6 varieties of the mosses of Kashmir

collected by Borelli and Duthie.

Varde (1922, 1923a & b, 1924) published an account of mosses

collected by Foreau, Andre, Roine and Velle from Madura in Southern

India. Subsequently Varde (1925) described 14 new species and reported

32 mosses new to Southern India based on the collections made by Foreau

from the then Madura district. Varde (1928) and Dixon and Varde (1927)

described a large number of new species and proposed two new genera

viz., Foreauella and Trigonodictyon.

In 1929, Bruehl and Sarkar described the mosses of Bengal. Blatter

in 1929 listed the mosses collected from the Bombay Presidency. Fleischer

(1929) while listing some mosses from Southern India, described the new

species Thuidium dodabettense.


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Foreau (1930, 1931) presented some notes of the bryological

geography for the Presidency of Madras, which includes a family-wise list

of 323 species and varieties, of which 4 genera, 95 species and 15

varieties were the new discoveries during 1908-1927 from this area. Later,

he (1961, 1964) compiled the result of the survey on the Moss Flora of

Palni hills and enumerated 424 species. Bruehl (1931) published a census

of Indian mosses in which he enumerated 2471 species including about

1500 species of the Indian continent.

Gupta (1933) studied the structure of Physcomitriellopsis indica Dixon

collected from Banares. Sharma (1949) published an account of the Indian

Sphagnum including the endemism of species and their economic

importance. He enumerated 17 species of Sphagnum, out of this seven

were indigenous. The majority of them occur in the eastern Himalayas,

Sikkim, Bhuttan and Khasi hills. Collections of mosses by Steward from

North Western Himalayas were enumerated by Bartram (1955). Ochi in

1956 supplemented this list. Deb (1955) reported 35 species of mosses

belonging to 28 genera and 17 families, from Manipur with a note on their

habitats. Chopra et al. (1956) published a preliminary list of mosses of

Mussorie consisting of 143 species belonging to 77 genera and 27 families.

Noguchi (1958) studied a collection of mosses from Southern India,

made by T.S. Rao from Kodaikanal, Coonoor and Ootacamund and

enumerated 27 taxa including a new variety in his revision of the family


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Trachypodaceae of India. Chopra and Sharma (1959) studied the life

history of Oligotrichum semilamellatum.

Gangulee (1957, 1959, 1960, 1961) in his series of papers on the

mosses of eastern India described the members of order Fissidentales in

1957, Ditrichaceae (1959), Dicranaceae (1960) and Leucobryaceae (1961)

respectively. Gangulee (1985) in his Handbook of Indian Mosses

enumerated 100 species including. Acrocarpous and Pleurocarpous

mosses with colour plates. Chopra in 1960 listed 158 species of mosses

belonging to 68 genera under 25 families from Nainital. The genus Alonia

was recorded for the first time from India by Wadhwa and Vohra (1963)

from Garhwal with a detailed account of A. rigida. Foreau in 1964 reported

Bryum coronatum Schwaegr. from Idukki district of Kerala and

B. curyphyllum Dixon & P. Varde, Philonotis subrigida var. adpressa Cardot

& P. Varde and Physcomytrium insigne Dixon & P. Varde from Kerala but

without mentioning the precise localities.

Ochi (1964) enumerated some species of the family Bryaceae and

Bartramiaceae from Darjeeling and Sikkim including one new species and a

new variety. Robinson in 1956 described Theriotia kashmirensis Robinson

and Tortula websteri Robinson together with a list of 60 species including

40 mosses and 20 liverworts collected from Kashmir. Srivastava (1966)

listed 137 species of mosses belonging to 70 genera and 27 families from

Kumaun. Chopra and Kana (1966) reported 17 species of mosses from

Delhi, of which Bryum klinggraeffi was a new record to India.


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Vohra and wadhwa in 1964 reported 60 species of mosses under 35

genera and 13 families collected from Nilkanth and Chaukhamb. Wadhwa

and Vohra in 1965 reported 36 species of mosses under 26 genera and 15

families based on collections of expedition to Chooyu.

Chopra in 1967 studied the relationship between mosses and

liverworts and concluded that the differences between the two groups

outweigh the resemblances and hence neither of the taxa were derived

from the other. Srinivasan in 1968 gave a general account of ecology and

distribution of Indian liverworts and mosses. Further he listed species which

are common with the various continents and insular areas. Dabhade in

1969 reinvestigated the four species of Bryum from Western India providing

their taxonomic details, habit and range of distribution. Later, he (1970)

reported Funaria nutans (Mitt.) Broth. from Khandala as a new record to

Western India. Haji Mohamed (1986) reported 42 species of mosses from

Mahabaleswar, Pune and Khandale in north Western Ghats.

Vohra (1969a) described the new species Habrodon kashmiriensis

Vohra, along with illustrated account of six other species from Kashmir,

Tehri and Dehra Dun. Subsequently Vohra (1969b, 1970) enumerated 165

species of mosses from Western Himalayas, of these three species were

new to India and five were new to Western Himalayas.

Raghavan and Wadhawa (1968) reported 28 species of mosses

belonging to 21 genera and 16 families from Agumbe-Hulical ranges in the

Shimoga district of Karnataka state. This list included two new records to


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India and Southern India. They in 1970 studied and presented a key to

mosses of Agumbe including two species viz., Bryosedgwickia densa

(Hook.) Bizot & P. Vard and Dendropogonella rufescens (A. Schimp.) Britt.,

as new records for India.

The report of the genus Buxbaumia (Udar et al., 1970b) from the

Western Himalayas was an addition to the moss flora of India. They in the

same year (1970c) described B. himalayensis Udar et al., which was

growing on rottenwood. Vohra in 1974 described a new species of

Plagiothecium viz., P. dehradunense from the northwestern Himalayas.

Srinivasan in 1974 published an account of the anatomical and

morphological aspects of 64 species of mosses, collected from Palni, Gingi,

Yercaud, Thirupathi, Kodaikanal, Nilgiris, Thiruvananthapuram and Mysore.

Chopra (1975) published his monumental work Taxonomy of Indian

Mosses which includes nearly 2000 species belonging to 329 genera under

56 families. He also included the collections from the neighbouring regions

such as Pakistan, Nepal, Bhutan and Western and South-eastern Tibet as

most of the species extend to other regions also.

Vohra in 1977 critically reviewed the studies on bryophytes done in

Botanical survey of India with reference to Musci and provided the location

of large number of collections of Indian mosses, deposited in foreign

countries. During his study on the order Hypnobryales (Musci) in the

Himalayas, Vohra (1978) described a new species, Lescuraea

darjeelingensis and a new variety, Haplocladium microphyllum ssp.


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capillatum var. bhutanicum. Subsequently Vohra (1979) made a few new

combinations in Hypnobryales and added two species to Indian bryophytic

flora. Gangulee (1969-80) made valuable contributions on the mosses of

eastern India and adjacent regions. His work, Mosses of Eastern India and

Adjacent Regions - A Monograph, provided a well illustrated, detailed

taxonomic account, and the distribution map of each species dealt with.

This is the first of its kind in India and certainly forms the basic reference

manual to study the Indian mosses.

Vohra (1981a) reported the Indian endemic species Pogonatum

hexagonum Mitt. from Silent Valley of Kerala State. Subsequently, Vohra

(1981b) mentioned about his 250 collections of mosses from Mulla Periyar

and its catchment area in the Idukki district of Kerala State. Chopra and

Kumar in 1981 published a well illustrated taxonomic account of 65 mosses

of Western Himalayas and adjacent plains. Bhatla in 1981 studied the

effect of pH in gametangial formation in the moss Bryum argenteum Hedw.

and found that the change in pH of the medium was one of the after-effects

of the gametangial formation, rather than its cause, i.e., the change in pH

has no role in the onset of reproduction phase.

Vohra et al. in 1982 published a list of 83 species of mosses collected

from Silent Valley. They could also locate the fruiting material of

Handeliobryum setschwanicum Broth. for the first time. Vohra in 1983 gave

a detailed description on the Hypnobryales suborder Leskeineae of the

Himalayas. He described and illustrated 42 genera and 161 species from


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Himalayas out of the 45 genera and 177 species occurring in India. Among

the 161 species, 44 are endemic in this area. Buck in 1984 provided the

taxonomical and nomenclatural notes on the west Indian Hypanaceae

which also include two new species and three varietal combinations.

Pant and Tewari (1983, 1938) made an assessment of bryophytic

vegetation of Nainital and surroundings based on substrate preferences of

genera and species. Bapna in 1980 reported the occurrence of Fissidens in

Rajasthan.

Chopra and Bhatla (1983) found that bryophytes exhibit maximal

gametangial initiations in a definite pH range and the nutritional status of

the medium has variable effect on gametangial induction. Vasishta and

Chopra (1986) in the experimental studies on bryophytes in India provided

a review of studies on gemmae, apogamy, apospory, callus induction and

its differentiation and protoplast isolation and culture. Kumra (1995)

conducted in vitro studies on Bryum coronatum and Funaria hygrometrica

Hedw.

Cytochemical studies on the developing sporophyte in the moss

Physcomitrium cyathicarpum Mitteni were carried out by Ekalavya (1987).

Mehra (1988) carried out studies on the effect of some growth regulators on

the formation and behaviour of protonemal gemmae in the moss Hyophila

involuta (Hook.) A. Jaeger and found that the auxins delay the germination

of gemmae. Babbar in the same year (1988) conducted similar studies in

Philonotis lancifolia Mitt. and Trematodon brevicalyx. The development and


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behaviour of protonemal gemmae in vitro in two gemmiferous mosses,

Trematodon brevicalyx and Semibarbula orientalis has been made by

Babber and Chopra in 1995. Mitra (1995) studied the morphology of moss

protonema for understanding the morphogenetic changes, which occur

during growth and development of protonema.

Ellis (1992a) made studies on the species Fissidens ceylonensis

Doxy & Molk. and compared with related taxa such as F. ceylonensis var.

acutifolius Dixon & P. Varde and F. kawarensis Dixon. He in the same year

(1992b) studied the taxonomy of some taxa of Calymperaceae occurring in

the Indian Peninsula.

Vohra and Kar (1996) reported 37 species of mosses from Great

Nicobar Island. Among them Taxithelium verneiri (Duby) Besch., was a new

record for India and 17 species occurring in the mainland were new

additions to the Island flora. Leucophanes nicobaricum C. Muell. &

Gangulee and Trichosteleum punctipapillosum Gangulee, endemic to the

Island so far represented by the type material only, have been recollected

after more than a century. Ochyra (1998) discussed the identity of two

species of Hygroamblystegium endemic to northern India.

Narayan et al. (2001) illustrated, morphotaxonomic details of eight

species of mosses belonging to six genera and six families from Great

Himalayan National Park, Himachal Pradesh.

Fossil remains and fossil studies on this group are rare. Pant and

Singh (1991) attempted to distinguish the spores of Hepaticae and


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Anthocerotales from iso- or microspores of Pteridophytes and pollen grains

of Gymnosperms and Angiosperms for the identification of possible fossil

spore dispersal of bryophytes reported from different geological horizons.

Chandra (1995) recovered a good assemblage of bryophytes from the early

Permian sediments of India as impressions along with the typical

Glossopteris flora for the first time. Both Hepaticae and Musci are

represented by newly designated form genera and species. Remarks on the

naming of the fossil bryophytes, the possibility of preservation, spore

distribution, probable habitat and evolutionary aspects have also been

discussed by him. Pant (1995) discussed the palynology of bryophytes.

Ecological studies on this group is very meager in India. Negi and

Gadgil (1997) studied the species diversity and community ecology of

mosses of Garhwal Himalaya and concluded that the microhabitat and

altitude seem to be the major ecological factors governing species diversity

and composition. Negi (2001) studied the ecology of bryophytes of Chopta-

Tunganath in the Garhwal Himalaya.

Nath et al. (2000) discussed the role of bryophytes in soil

management and in rock binding.

SEM studies of spores of two species of Bryum and seven species of

Fissidens from the Western Ghats have been done by Dhabade and Pat il

(2001). Parihar (2001) presented a method of preparation of a perpetual

taxonomic cross-reference dictionary of the bryophytes, for this a system of

numeration of each taxon has been adopted. Chaudhary and Deora (2001)


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provided an illustrated account of 25 species of mosses belonging to 18

genera and 8 families from Mt. Abu. Among them 23 species occur in

Western Himalaya, 19 in Eastern Himalaya and 15 in South India. Pant

(2001) highlighted the diverse habitat for bryophytes and given a checklist

of the bryo-diversity of Kumaun Himalaya pointing the need to explore and

study the diverse species.

Saxena et al. (2001) highlighted the usefulness of bryophytes as

subjects of biomonitoring studies and called for the establishment of

bryophyte bank for environmental monitoring in India. Tewari (2001)

described the distribution of “copper mosses” Scopelophila ligulata

(Spruce) Spruce and S. cataractae (Mitt.) Broth. in Kumaun Himalaya and

mentioned the plant-substrate relationship and indicator value of these two

species. Kapoor and Bhatla (2001) studied the biochemical regulation of

protonema differentiation in Funaria hygrometrica.

Pandey et al. (2001) studied the accumulation of heavy metals such

as copper, chromium, lead, zinc, nickel and cadmium in bryophytes from

the banks of Ganga. They found high concentration of heavy metals and

which may be accumulated from the polluted waters of the river. Singh

(2001) discussed the status, vulnerability and conservation of the Indian

liverworts. An action plan for the conservation and monitoring has been

suggested and need for capacity building in its taxonomy and integration of

the regional herbaria is also emphasized. Vashistha (2001) reviewed the

heavy metal pollution and the role of bryophytes in reducing the metal


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pollution. Saxena (2001) monitored the impact of heavy metals such as Pb,

Ni, Cr and Hg using bryophytes in Nainital.

Saini and Vashistha (2001) studied the effects of some auxins (IAA,

NAA, NDA, 2,4-D) and cytokinins (BAP, 2iP and Kinetin) on the protonemal

growth and bud formation in the moss Anoectangium bicolor Rev. & Cardot.

Uniyal (2001) made cytological observations on 4 species of Philonotis viz.,

P. angusta Mitt., P. falcate (Hook.) Mitt. P. Fontana (Hedw.) Brid. and

P. turneriana (Schwaegr.) Mitt.

Banerjee (2001) made a review on the antimicrobial activities of

bryophytes, which shows that nearly 200 species of bryophytes have been

surveyed so far to detect such activity, of which 53-76% of the plants have

shown positive results. Mosses are the most studied one followed by

thalloid and leafy liverworts.

Daniel and Daniel (2003) added Fissidens grifithii to Indian bryoflora

from Kanyakumari district of Tamil Nadu. He in the same year described six

species viz., Fissidens kalimpongensis Gangulee, F. leptopelma Dixon,

Leptolejeunea sikkimensis Udar &U. S. Awasthi, L. jhimalayensis Pande &

Misra, Radula madagascariensis Gottsche and Leucobryum juniperoideum

(Brid.) C. Muell. as new record to Peninsular India.

Nair et al. (2004) recorded Bryum tuberosum Mohamed & Damanhuri,

which was known earlier from the Peninsular Malaysia, as a new record for

India from Uduppi of Karnataka State. Singh and Nath (2004) added an

epiphytic liverwort Frullania rotundistipula Steph. from Khasi hills,


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Meghalaya to the Indian bryoflora. Singh and Singh (2004) added Lejeunea

flava (Swartz) Nees to the bryoflora of Western Himalaya, a species

already known from Eastern Himalaya to Indian flora.

Pande and Joshi (2004) assessed the species composition,

phytosociology, pattern of biomass and net primary production of bryoflora

growing on decaying logs in the evergreen conifer forest of silver fir of

Nanda Devi Biosphere reserve in Central Himalaya.

The sequenced genome of the moss Physcomitrella patens provides

a powerful tool for comparative analyses of land plant genomes. In parallel,

several tools for studying gene function have been developed in P. patens,

including RNA interference, inducible promoters and gene targeting, a

unique attribute of this plant system. Transcriptomic analyses illustrate

commonalities among plant lineages in gene content, structure, and

regulation. Transgenic studies show that the regulatory factors Abscisic

Acid Insensitive-3 (ABI-3) and Leafy (LFY) have molecular functions that

are conserved between moss and angiosperms, in spite of the fact that they

function in non-homologous tissues. Future work in P. patens will contribute

to our understanding of the molecular basis of plant development and

evolution (Quatrano et al., 2007).

Szczepaniak and Biziuk (2003) studied on mosses and lichen, in spite

of all disadvantages, are good tools for air pollution monitoring, but best

results could be achieved while using both of them together, because of

differences in their metal uptake and retention. The researcher dealing with


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31

biomonitoring faces many difficulties; for example, in the use of lichens,

those of similar composition are not easy to find, because of the differences

caused by the tree on which the lichens are growing.

Meyer (1940) worked on leafy gametophytes of Physcomitrium

turbinatum (Michx.) Brid. develop from primary protonemata submerged in

a liquid nutrient medium, and neither direct exposure to air nor attachment

to a solid substratum is essential for that development. However, both the

rate of gametophores development and the number of leafy plants formed

are less than on a solid nutrient substrate. Gametophytes developed in

water cultures show pronounced morphological modifications when

compared to plants grown normally in air. Growth of plants in water cultures

indicates that the normal nutrition of protonemata as well as that required

for the development of leafy gametophores is entirely autotrophic in the

species used.

Historical account of the Indian Moss Flora (Gangulee, 1969) would

be better understood by dividing it into four different zones as follows:

1. Eastern Zone - including Eastern Himalayan region, Bengal, Orissa,

Sikkim, Bhutan, NEFA (Arunachal Pradesh), Andaman-Nicobar and

Burma.

2. North-West Zone - including north-west Himalayan region, Punjab,

Kumaon, Kulu Manali Valley, Simla, Mussourie, Dehra Dun, Naini Tal,

Kashmir, etc.


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3. South Indian zone - including Nilgiris, Ootacammund, Palni Hills, Jog

Falls, Eastern Ghats and Ceylon.

4. Western Zone - including Western Ghats (Sahyadri ranges), Satpura

ranges, Mt. Abu.

Moss Flora of Eastern India

The Eastern Indian Zone comprising Nepal, Burma, NEFA, Assam,

was first explored by Dr. Francis Buchanan Hamilton (1802-1803) from

Kathmandu valley or Nepal. During years 1815 to 1835, Dr. Nathaniel

Wallich made extensive collection of mosses of Nepal, Lower Burma and

published them in his famous “Wallich Catalogue” (1826-1832), listing 114

mosses. A complete list with notes on these mosses was published by

W. H. Harvey and J. D. Hooker (1840). Many of them were also illustrated

by J. D. Hooker (1840, 1857) in “Icones Plantarum”, based on the collection

of mosses from Assam and Nepal. After Dr. Wallich, further exploration was

made by Dr. Griffith (1835-1845) in Assam, Khasia, Burma, Bhutan, Sikkim,

Central India and Malacca. He was serving as an Assistant Surgeon in East

India Company, but due to his premature death in 1845 his work was

published posthumously by J. McCleland (1849) as “Notulae and Plantas

Asiaticae”, and illustrations of them in a supplement to “Icones Planarum

Asisticarum”. Out of 9000 species of plants described in this work 101 are

mosses; some other were also described from Afghanistan in “Afghan

Catalogue”.


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The next sizable work on the moss flora from Rajmahal Hills, Karen

Hills, Arakans, Pegu in Burma and of Andamans was by Kurz (1867-1871),

who had been working as a Curator at the Royal Botanical Garden, Sibpur

in 1894. Another part of Eastern region viz. Manipur was surveyed by Dr.

George Watt (1881-1882), Gammie (1896-1897), Miss M. Roberts (1893),

Oliver (1896-1899), Dr. King (1893-1902), Bryotheca E. Levier (1881-1904),

explored mosses from far Eastern Himalayas. This material was determined

by Brotherus (1907-1908) at the British Museum. The Moss flora of Bilaspur

and Amarkantak was surveyed by Martin (1899-1901) in the beginning of

the 20th century, and that of Darjeeling, abor Hills, sadiya region of NEFA

by Miller (1901), Miss Craig (1910), Mrs. Roper and Miss Shephard (1911-

1912) and Burkill (1911-1912). During the last sixty to seventy years quite a

large number of Muscologists have showed increasing interest in the

Eastern Himalayan regions, especially Dr. T. H. Somervell (1924) who

collected a rare moss Angstromla julacea (Hook.) Mitt. at an altitude of

6530 m from Rongbuk Valley during British Mount Everest Expedition.

Captain Kingdon Ward (1928) collected in the Upper Assam and Burma. Dr.

Bor (1933-1936) has collected 208 species of moss in Aka Hills of NEFA,

Dr. Kerstan (1935) at Darjeeling and Dr. Troll (1936) in Sikkim. F. Ludlow,

G. Sheriff and G. Taylor of the British Museum (1939 and 1949) collected

mosses from South-East Tibet and Bhutan and A. C. Cole (1944) from

Upper Assam and NEFA. These were described by Robinson (1964).

R. S. Chopra (1956) had made some collection of mosses from Darjeeling.


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Another Botanical party led by Norkett (1961) made extensive collection of

mosses in Nepal and adjacent area of Eastern India in 1961. Dr. K. Yoda

(1963), Norkett (1961-62) of British Museum, London, A. J. Sharp and

Z. Iwatsuki (1965), etc. also explored the moss flora of Eastern Himalayan

region. Norkett and Gangulee have made further large collections from this

region in 1966 (Gangulee, 1969).

Moss Flora of North-West Himalayas

Nathniel Wallich (1815-1835) was the first to make collection of

mosses from Western Himalayas. He published their list in “Wallich

Catalogue” in 1828-1832. In 1956, Thomson made a large collection of

mosses in Uttar Pradesh. Later Gamble (1896) explored the moss flora of

Simla, Mussories and Dehra Dun. Gollan (1896), the then Superintendent

of Botanical Garden at Mussorie, Duthie (1900) the then Superintendent of

Botanical Garden at Saharanpur and other like Radhalal, Bahadur,

Kabirkhan, Inayatkhan made collection of mosses from the North-West

Himalayan regions. In 1998, Brotherus, who had worked on the moss flora

of this region of Himalayas published them in Acta Societies, Scientiarum

Fenica Helsingfors. Badhwar made a collection of mosses of this region

and a paper on them was published by Dixon and Badhwar in 1938. They

brought out a number of new genera and species (Gangulee, 1969; Chopra,

1975). In 1961 and 1966, A. H. Norkett of the British Museum, London,

made intensive collections in the North-West Himalayan region and

published “A Note Himalayan Mosses”. For the last 35 years R. S. Chopra


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35

has been working on them with a view to monographic from entitled,

“Taxonomy of Indian Mosses” from different localities in North -West

Himalayas such as Simla, Mussorie, Dehra Dun, Naini Tal, Shiwalik Hills,

Kashmir etc. (1975). His student Vora (1960) has worked on the moss flora

of Mussorie and partly of Kashmir. Wadhwa (1958) published a paper on

the Mosses of Simla. Srivastava (1966) has surveyed the moss flora of

Naini Tal in Kumaon valley of Himalayas, and Chopra and Kumar (1981)

published an illustrated account of the mosses of Western Himalayas and

adjacent plants.

Moss Flora of South India

Very little work has been done on the mosses of South India.

Brotherus (1899) had made a preliminary survey of the mosses of South

India and has published his account in the Records of Botanical Survey of

India (1899). The mosses he examined were collected by Dr. Walker in

Coorg and Srilanka (Ceylon). Then Fisher and others (1910) had made a

collection of mosses of South India and Ceylon and they were published by

Dixon in 1914. Dixon (1921) further found a number of new moss genera

and species. He has also described mosses from North Kanara now in

Karnatak State. In 1931 Bruhl published “A Census of Mosses from South

India” (Gangulee, 1969). Later, Father Foreau (1961) and (1964) made

extensive collections of Mosses of Nilgiris and Palni Hills especially at

Ootcammund and published “Moss Flora of Palni Hills”. Recently, Norkett

(1966-1970) had made bryological collections at Ootacummand, Nandi Hill,


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36

Kovalam, Thekadi, Nagercoil, Manjoli and other places at the end of

Western Ghats in Mysore and Kerala border. Dabhade (1971) had collected

a number of hepatics and mosses from Jog Falls in Karnatak.

Moss Flora of Western India

In the 19th century, Mosses of Western Ghats were collected from

diverse localities by enthusiastic botanists like Woodrow (1895), Surgeon

General Kirtikar (1897), Sedgwick (1908), Rev. Fr. Blatter (1905) and a few

others. Kirtikar (1897) had collected them at Mahabaleshwar, Matheran and

Bombay and they were published in Birdwood’s “Catalogue of the Flora of

Matheran and Mahabaleshwar”, 1897. Maxwell (1908) and Dixon (1921)

made collection from the forests of Karnataka district and Sedgwick (1910,

1911, 1913) from different localities in Western Ghats such as Purandar,

Poona, Mahabaleshwar, Lonavala, Khandala, Trimbakeshwar, etc. They

were published in three different papers on the “Mosses of Western Ghats”.

Gammie (1905-1910) and Burns collected mosses from Sakar Pathar or

Lonavala and adjacent area. Dixon (1910-1921) who had special interest in

the mosses of Sahayadris or Western Ghats discovered several new

genera and species like Merceyopsis sp. (1910), Hyophilopsis sp. (1911),

Bryosedgewickia kirtikari Card. et. Dix. (1912), etc. After Dixon, there has

been a big gap in studies on moss flora of Western Ghats. Thereafter

Dabhade and Norkett (1961) had collected them at Mahabaleshwar,

Khandala, Purander Fort, Kankeshwar, Kasara, Amba Ghat, etc. in Western

Ghats. Norkett (1970) discovered, Trachycarpidium sp. characterized by


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37

spiny surfaced capsule from Khanadala. Incidently this genus is first record

from Asia. Dabhade (1968) reported Entosthodon nutans, Mitt. and

Archidium indicum C. Muell. which were new to South and Western India.

Raghavan and Wadhwa (1968), while listing the mosses of Agumbe,

recorded two new species Dendropogonella rufescens (Schimp). Britt. and

Bryosedgwickia densa (Hook.) Biz. et P. Vard and Kaji Mohamed et al.

(1985) had listed 26 mosses from North-West of Western Ghats.

3.3 ANTIMICROBIAL STUDIES ON BRYOPHYTES

In recent years there has been a commendable progress made in

bryological researches in the country, especially in mosses. This has been

not only in taxonomy and morphology of this group but also in such

fascinating fields as cytology, tissue culture, morphogenesis, physiology

and pharmacognosical and pharmacological studies. There has been

number of works has been done in antimicrobial works on liverworts in

mosses.

Banerjee and Sen (1979) were concerned with an examination of the

antibiotic activity of 52 species (in 40 genera) of the bryophytes. The plants

were extracted in water, methanol, ethanol, ether and acetone and tested

against 12 microorganisms, including 3 Gram positive (Staphylococcus

aureus, a penicillin-sensitive and penicillin-resistant strain, Sarcina lutea

and Bacillus subtilis), 5 Gram negative (Escherichia coli, Salmonella typhi,

Vibrio cholerae, Klebsiella pneumoniae and Pseudomonas aeruginosa),

one acid-fast bacterium (Mycobacterium phlei) and 3 fungi (Curvularia


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38

lunata, Aspergillus niger and Helminthosporium oryzae). Solubility data and

antibiotic spectra of the active plants indicate the occurrence of a variety of

antibiotic substances among bryophytes. Out of 52 species of bryophytes,

29 (56%) were active against at least one of the test bacteria, but none

possessed any antifungal property. The moss Brachythecium procumbens

and the liverworts Asterella sanguina and Marchantia paleacea showed the

broadest spectrum of antibiotic activity. Among the test organisms used

Salmonella typhi was found to be most sensitive.

The first in vivo experiments for antifungal activity have been

performed at Bonn University by Jan-Peter Frahm (2004). Alcoholic

extracts of all twenty bryophytes used had an effect on a variety of crops

infected with different fungi. Two liverworts showed systemic effects. Based

on these results, commercial products from bryophytes have been

developed and are sold in Germany. Bryophyte extract also has effects on

human pathogenic fungi and may cure skin diseases. This same extract

shows antifeedant effects against slugs.

Semra Ülhan et al. (2006) studied and described the antimicrobial

activity of 2 extracts of Palustriella commutata (Hedw.) Ochyra, collected

from the S.Ndiken Mountains, Eskisehir, Turkey. These extracts were first

prepared with acetone and methanol solvents under reflux. The

antimicrobial activity of the extracts was then assessed using the disc

diffusion method against 11 bacteria, 1 yeast, and 8 moulds. The

antimicrobial test results revealed that the acetone extract had a potential


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39

activity against 9 test bacteria. While some gram-positive bacteria tested

were sensitive (Bacillus mycoides, B. cereus, B. subtilis, and Micrococcus

luteus), all gram-negative bacteria tested (Klebsiella pneumoniae, Yersinia

enterocolitica, Pseudomonas aeruginosa, Escherichia coli, and

Enterobacter aerogenes) were sensitive to the acetone extract. Both

extracts were inactive against yeast and mould strains. P. commutata

possesses a new antimicrobial compound that has an effect against gram-

negative and gram-positive bacteria.

The study of Vuotto (2000) was designed to evaluate the antibacterial

and antioxidant activities of an aqueous extract from the tropical Feijoa

sellowiana Berg. fruit which is widely used for human food. The extract was

tested against Gram-positive and Gram-negative bacteria by a broth

dilution test and on human whole blood leukocytes, as well as isolated

neutrophils using a chemiluminescence (CL) assay. The extract inhibited

bacterial growth; Pseudomonas aeruginosa, Enterobacter aerogenes and

Enterobacter cloacae were the most sensitive. The fruit extract significantly

decreased CL emission from human whole blood phagocytes and isolated

polymorphonuclear leukocytes whether they were activated or not by

soluble or phagocytic stimuli. F. sellowiana showed both antibacterial and

antioxidant properties and therefore its extract might be used as a new

multifaceted drug.

The synthesis and potent antiprotozoal activity of 14-hydroxy-

lunularin, a natural hydroxybibenzyl bryophyte constituent is reported by


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Roldosa et al. (2008). 14-Hydroxylunularin was highly active in vitro assays

against culture and intracellular forms of Leishmania spp. and

Trypanosoma cruzi, in absence of cytotoxicity against mammalian cells.

Preliminary structure - activity relationship studies showed that the reported

bioactivity depends on hybridization at the carbon - carbon bridge, position

and number of free hydroxy group on the aromatic rings. The reported

results were also in agreement with the in silico prediction using Non-

Stochastic Quadratic Fingerprints-based algorithms. The same compound

also showed antiprotozoal activity in Leishmania spp. infected mice by oral

and subcutaneous administration routes, with an optimal treatment of a

daily subcutaneous administration of 10 mg/kg of body weight for 15 days.

This study suggested that 14-hydroxylunularin may be chosen as a new

candidate in the development of leishmanicidal therapy.

Plagiochasma appendiculatum (Aytoniaceae) of the order

Marchantiales is widely used in the form of paste ethnomedicinally by

Gaddi tribe in Kangra valley for treating skin diseases. In this context,

antimicrobical potential of Plagiochasma appendiculatum against a wide

range of microorganisms was studied. To validate the ethnotherapeutic

claims of the plant in skin diseases, wound healing activity was studied,

besides antioxidant activity to understand the mechanism of wound healing

activity. The plant (alchoholic and aqueous extract) showed significant

antibacterial and antifungal activity against almost all the organisms:

Micrococcus luteus, Bacillus subtilis, Bacillus cereus, Staphylococcus


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41

aureus, Streptococcus pneumoniae, Enterobacter aerogenes, Escherichia

coli, Klebsiella pneumoniae, Proteus mirabilis, Pseudomonas aeruginosa,

Salmonella typhimurium, and eight fungi Candida albicans and

Cryptococcus albidus - dimorphic fungi, Trichophyton rubrum -

dermatophyte fungi, Aspergillus niger, Aspergillus flavus, Aspergillus

spinulosus, Aspergillus terreus and Aspergillus nidulans - systemic fungi,

with especially good activity against the dermatophyte (Trichophyton

rubrum) and some infectious bacteria (Escherichia coli, Proteus mirabilis

and Salmonella typhimurium) with an MIC of 2.5 μg/disc. The results show

that Plagiochasma appendiculatum extract has potent wound healing

capacity as evident from the wound contraction and increased tensile

strength. The results also indicated that Plagiochasma appendiculatum

extract possesses potent antioxidant activity by inhibiting lipid peroxidation

and increase in the superoxide dismutase (SOD) and Catalase activity

(Singha et al., 2006).

The National Cancer Institute’s record of bryophytes (mosses,

liverworts, and hornworts) screened for biologically active chemicals is

reviewed. In 1977, an extract of a moss from Maryland, Polytrichum

ohioense, showed cytotoxicity in KB cell culture, but antitumor activity was

not discovered in bryophytes until 1980. This led to a greater emphasis in

collecting and screening bryophytes during 1980-1981. Samples weighing

0.5–2 kg (dried) were collected for each of 208 species and varieties.

These included 184 species in 97 genera of mosses, 23 species in 16


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genera of liverworts, and 1 hornwort. The methodologies for f ield work,

extraction, and screening are briefly described. Extracts of 75 species were

toxic; extracts of 43 species were active. Activity was especially noted in

the moss families Thuidiaceae, Mniaceae, Neckeraceae, Hypnaceae,

Brachytheciaceae, Polytrichaceae, Dicranaceae, and Grimmiaceae. The

authors suggest that bryophytes are a promising source for discovery of

novel biologically active compounds. The possibility that biological activity

in bryophytes may be the result of allelopathy, or the reputed ab ility of

bryophytes to accumulate toxic substances, is considered (Spjut et al.,

1986).

Neelam Mewari and Padma Kumar (2008) were worked on the crude

methanol and flavonoid (free and bound) extracts of Marchantia

polymorpha L. (Marchantiaceae) and were screened against three bacterial

strains, viz., Escherichia coli, Proteus mirabilis (Gram negative), and

Staphylococcus aureus (Gram positive), and four fungal strains, viz.,

Aspergillus flavus, A. niger, Candida albicans , and Trychophyton

mentagrophytes. Disk diffusion and microbroth dilution techniques were

performed for evaluation of antimicrobial activity of the extracts. Total

activity of extracts (ml) against each sensitive pathogen was also

evaluated. Methanol and free flavonoid extract showed best activity against

S. aureus (IZ 20.6 and 19.6 mm, MIC 0.281 and 0.312 mg/ml, MBC 1.125

and 0.312 mg/ml, respectively). However, all the microorganisms were

found to be sensitive against the extracts tested. Total activity for


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P. mirabilis and S. aureus for methanol extract was found to be the same

(124 ml), but was maximum for free flavonoid against C. albicans (199 ml).

Bryophytes, the previous land plants posses medicinally important

bioactive compounds but with little information. Traditionally the bryophytes

plants posses some bioactive components and therefore used throughout

the world as drugs and remedies to cure the various diseases. Evaluation

of antimicrobial effect of mentioned bryophytes on some pathogenic

microorganisms. Different Solvent fractions of selected bryophytes were

obtained and dried in vacuum. Antimicrobial effect of these fractions was

determined by agar diffusion method on different pathogenic

microorganisms. The result was then compared with the standard

antibiotics ampicillin and nystatin (10 g/ml). Bodade et al. (2008) studied

with the bryophyte extracts and were found to be active against at least one

of the test organisms except Racomitrium crispulum. The extracts that

displayed antibacterial activity were neither always effective against the

same organisms, nor consistent in magnitude of inhibition. Ethanolic,

acetone and chloroform extracts were found to be more effective on

Escherichia coli and Staphylococcus aureus. Among the fungi Aspergillus

niger was most sensitive to the ethanolic extract of Plagiochasma

appendiculatum and Bryum argentium.

Sphagnum plants grow in natural, species-poor carpets at low pH but

without any known substantial fungal disease. To investigate this

phenomenon, we analysed bacterial populations associated with two


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Sphagnum species with different ecological behaviour, namely

S. magellanicum and S. fallax, from three sites in Germany and three in

Norway, with a special focus on the functional group of antagonists. The

screening of 493 bacterial isolates for antagonistic activity against fungal

pathogens resulted in 237 (48%) active isolates. We found a higher

proportion of antagonists for S. magellanicum (24%) than we did for

S. fallax (19%) in general. The majority of the antagonists belonged to the

genera Serratia (15%), Burkholderia (13.5%), Staphylococcus (13.5%), and

Pseudomonas (10%). In contrast to the high moss specificity found for

antagonistic bacteria, Burkholderia as well as Serratia isolates with highly

similar molecular fingerprints as ascertained by BOX-PCR for both

Sphagnum species were found. Interestingly, a high proportion of

antagonists, for example Staphylococcus, Hafnia, Yersinia, and Pantoea,

were identified as strains that are known as facultative pathogens of

humans. Sphagnum plants represent an ecological niche not only for

diverse and extraordinary microbial populations with a high potential for

biological control of plant pathogens but also for opportunistic human

pathogens (Opelt et al., 2007).

Disc diffusion assay was used to screen for antibacterial activity of

ethanolic extracts of 60 bryophytes belonging to 39 genera, including 38

liverworts, one hornwort, and 21 mosses from China and Mongolia. Out of

60 bryophytes, 56 species (93.3%), including all liverworts tested, have

detectable antibacterial activity against at least two of the selected seven


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45

bacteria, including four Gram positive (Bacillus megaterium, Bacillus

subtilis, Bacillus thuringiensis, and Staphylococcus aureus) and three Gram

negative (Escherichia coli, Pseudomonas aeruginosa, and Pseudomonas

putida). Seventeen species (two mosses and 15 liverworts) are active

against all seven selected bacterial species. Out of 21 mosses, four

species did not show positive evidence. Antibacterial activity was

particularly prominent in the members of Conocephalum, Frullania,

Herbertus, Marchantia, Mastigophora, and Porella. The antibacterial activity

of the alcoholic extracts of Bazzania tridens, Herbertus aduncus, Porella

densifolia, Polytrichum commune, and Thuidium kanedae, expressed as

MICs (minimal inhibitory concentration) and MBCs (minimal bactericidal

concentration), were compared with three reference antibiotic drugs. Out of

the seven bacteria tested, Staphylococcus aureus is most resistant to the

extracts of both liverworts and mosses. Pseudomonas putida is most

sensitive to the extracts of mosses, and Bacillus subtilis is most sensitive to

the extract of liverworts. The broad spectrum of antibacterial activity shown

in the present study suggests that most liverworts are worthy of further

investigation for the nature of their definitive antibacterial compounds and

other potentially biologically active ingredients. The investigation of oil

bodies of the tested liverworts reveals that there is no correlation between

the antibacterial activity and the size and numbers of oil bodies (Zhu Rui -

Liang et al., 2006).


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46

An acetone extract of the moss Pleurochaete squarrosa was tested

against eleven bacterial strains, some of which are pathogenous for man.

The extract was active on some Gram-negative strains. Antibacterial

activity of the extract, expressed as MICs, was compared with three

reference antibiotics. Acute toxicity assay was performed in Balb C mice

(Basile et al., 1998a).

Meenakshi Singh et al. (2007) evaluated the antimicrobial activity of

ethanolic extracts of 15 Indian mosses. The antibacterial activity of

ethanolic extracts was investigated against five G(+) and six G(−) bacterial

strains. Antimycotic activity was assayed against 8 fungi. Sphagnum

junghuhnianum, Barbula javanica, Barbula arcuata, Brachythecium

populeum, Brachythecium rutabulum, Mnium marginatum and Entodon cf

rubicundus were found to be most active against all the organisms.

The antimicrobial activity of Bryum argenteum ethanol extracts was

evaluated by microdilution method against four bacterial (Escherichia coli,

Bacillus subtilis, Micrococcus luteus and Staphilococcus aureus) and four

fungal species (Aspergillus niger, Penicillium ochrochloron, Candida

albicans and Trichophyton mentagrophyes). All the investigated ethanol

extracts have been proved to be active against all bacteria and fungi tested

(Sabovljevica et al., 2006).

Basile et al. (1998b) evaluated the action of Lunularia cruciata

(Bryophyta) acetone extract against 13 bacterial and 2 fungal strains. On

pharmacological screening, substantial antibacterial activities were


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47

observed for the acetone extract of the mature gametophyte of L. cruciata

while antifungal activity was not observed on the strains tested. Adult thalli

of L. cruciata, collected in the Botanical Gardens of Naples, underwent

extraction with acetone. Inhibition of bacterial growth was compared with

that of Na-cefotaxime, benzyl penicillin, and tetracycline, while the

antifungal activity was compared with that of griseofulvin. The minimum

inhibitory concentration (MIC) and minimum bactericidal concentration

(MBC) were determined for the extract showing exclusively bacteriostatic

activity.

The antibacterial activity of methanol extracts of ten moss species

and fractions prepared from 80% methanol extract of Hylocomium

splendens were evaluated by disk diffusion method. Nine moss species

showed antibacterial activity against Gram (+) bacteria, in particular

H. splendens and its ethyl acetate fractions showed stronger activity.

Enhancement of antibacterial activity against Staphylococci by UV-A light

irradiation was demonstrated in the extracts of Bartramia pomiformis,

Ceratodon purpureus and Neckera douglasii (Kanga et al., 2007).

An acetone extract from the moss Rhynchostegium riparioides was

tested against 11 bacterial strains, some of which are pathogenic. The

extract was active on some Gram-negative strains. The antibacterial

activity of the extract, expressed as MICs, was compared with three

reference antibiotics (Basile et al., 1998c).


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3.4 PHYTOCHEMICAL STUDIES ON BRYOPHYTES

Only 30 years ago, the chemistry of bryophytes was virtually

unknown. Recent research on the biology of bryophytes and progress is

analytical techniques has resulted in a deeper knowledge about the

chemical constituents of bryophytes, although our understanding of their

biochemical processes, especially biosynthetic pathways, compared to

vascular plants, is still rather poor. So this chapter aimed to present it in

tandem with new developments in the phytochemistry of bryophytes.

Seven pure flavonoids were isolated and identified from five moss

species. The flavonoids were the flavones apigenin, apigenin-7-O-

triglycoside, lucenin-2, luteolin-7-O-neohesperidoside, saponarine and

vitexin; and the biflavonoid bartramiaflavone. Some of these flavonoids

were shown to have pronounced antibacterial effects against Enterobacter

cloaceae, E. aerogenes and Pseudomonas aeruginosa (minimal

bacteriostatic concentration MBC in the range of 4–2048 μg/ml) (Basile et

al., 1999).

Bryophytes contain a large number of terpenoids and phenolic

compounds. Recent investigations relating to the chemical constituents

found in 36 Japanese, 3 New Zealand, 2 European, 1 Argentinean and

1 Taiwanese liverworts and 2 Japanese mosses and their biological activity

are discussed. The chemosystematics of some liverworts as well as the

chemical relationship between liverworts and mosses, and bryophytes and

ferns are also discussed (Asakawa, 2001).


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49

Asakawa (2007) investigated and produced a great variety of

lipophilic terpenoids, aromatic compounds, and acetogenins from

liverworts. Many of these constituents have characteristic scents,

pungency, and bitterness, and display a quite extraordinary array of

bioactivities and medicinal properties. These expressions of biological

activity are summarized and discussed, and examples are given of the

potential of certain lead compounds for structure-activity studies and

synthesis.

Dried mosses (five moss species) were progressively extracted

and subjected to a four-step Craig distribution. Seven pure flavonoids

were isolated and identified. The flavonoids were the flavones apigenin,

apigenin-7-O-triglycoside, lucenin-2, luteolin-7-O-neohesperidoside,

saponarine and vitexin; and the biflavonoid bartramiaflavone and they were

submitted to biological tests. The tests were performed in vitro on spore

germination and protonemal growth of the moss Tortula muralis and on

seed germination and root growth of Raphanus sativus. Flavonoids caused

a decrease in the percentage of spore germination, protonemal

development and root growth. In addition they caused morphological

alterations, such as forked tips, swollen apices, rounded cells and early

formation of brood cells in the protonemata. Data were discussed in

relation to the presence of allelochemicals in mosses (Basile et al., 2003).

The essential oils of mosses of the genera Mnium, Plagiomnium,

Homalia, Plagiothecium and Taxiphyllum (Musci) have been investigated by


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50

gas chromatography and mass spectrometry. The new sesquiterpenes

(+)-10-epi-muurola-4,11-diene and 10,11-dihydro-α-cuparenone were

isolated by preparative gas chromatography and identified as major

constituents of the hydrodistillation products of Mnium hornum (Hedw.)

using NMR and mass spectrometry. In addition, (+)-dauca-8,11-diene and

two new butenolides, 3,4,5-trimethyl-5-pentyl-5H-furan-2-one and 3,4-

dimethyl-5-pentyl-5H-furan-2-one were identified as constituents in

Plagiomnium undulatum (Hedw.) T. Kop. Although the amounts of volatiles

present in the investigated mosses are generally smaller than in liverworts,

the spectrum of terpenoid compounds is similar. The investigated mosses

also generate aliphatic compounds of greater abundance and structural

variety (Saritasa et al., 2001).

Toyota in 1998 analysed the chemical constituents of diethyl ether

extract of the moss Floribundaria aurea subsp. nipponica and yielded

polypoda-7,13,17,21-tetraene. This is the first example of its isolation from

a moss, although it has been isolated from ferns. Further purification of the

extract afforded dammara-17Z, 21-diene in addition to diploptene, hop-22

(20)-ene. The structure of dammara-17Z, 21-diene was established by

extensive 2D NMR techniques. From gametophytes of Mnium hornum have

been isolated a sophorotriose caffeate, three biflavonoids, among them the

first biflavone methyl ether from a moss, and six derivatives of 4,2 ′-epoxy-

3-phenylcoumarin - a group of isoflavone-related compounds - that has

been found so far only in some seed plants. The structures of all


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51

compounds were elucidated and confirmed spectroscopically (Brinkmeier,

et al., 1999).

Two unusual flavonones coupled with styryl units, communins

A and B, and a new benzonaphthoxanthenone, ohioensin H, together with

11 known compounds, were isolated from the moss Polytrichum commune.

The structures of were assigned by spectroscopic data interpretation. The

new compounds were evaluated for cytotoxicity against a small panel of

cancer cell lines (Peng Fu et al., 2009).

2α-Cinnamoyloxy-6β-acetoxybornane has been isolated from the

liverwort Conocephalum conicum (Conocephalaceae), while Plagiochila

barteri and P. terebrans (Plagiochilaceae) furnished ent-spathulenol,

marchantins C and H, 1(10),14-halimadien-13ξ-ol, and trifarienol B.

Although the bisbibenzyl-type isoplagiochins have been reported to occur in

some Plagiochila species, the presence of marchantins in the genus is very

rare. All of the isolated compounds were tested for α-glucosidase inhibitory

activity but only marchantin C showed moderate activity (Harinantenaina,

2007).

Huaifang Guo et al. in 2007 did a detailed analysis of mass spectra

and generated bis(bibenzyl) compounds in bryophytes under electron

ionization time-of-flight (EI-TOF) and electrospray ionization triple-

quadrupole (ESI-TQ) mass spectrometry conditions.

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