The Lichens

• Lichens are symbiotic organism formed through the association of algae

and fungi. Their outer layer is made up of tightly bound filament and is
called the cortex.
• The inner layer with loosely bound hyphae together with the algal cells is
called the medulla.
• They are commonly found growing on the bark of trees, on the surface of
hard structures such as rocks or on walls of houses.
• Lichens are stable, self-supporting associations
between fungi and photobionts.
• Is a composite organism that emerges from algae or
cyanobacteria (or both) living among filaments of a
fungus in a mutually beneficial (symbiotic) relationship.
• The fungi are most commonly Ascomycota.
• The photobionts are either algae or cyanobacteria.
• The algae may be single celled or, rarely, filamentous.
• In most cases, neither partner is found living alone.
• The fungus most commonly forms the majority of the
biomass of the lichen
• Lichens are common primary colonisers of
stressful habitats.
• They are major structural determinants of
extremely cold environments, and extremely dry
climates.
• They are also seen as the surface layer of trees,
rocks, roofing tiles and exposed surfaces in many
different environments.
• In moist environments, the foliar forms are more
obvious, being attached to limbs and either
growing up or falling down from the plant.
• Lichens are not found in heavily shaded sites
because they require light to support their
photosynthesis.
• In general terms, the photobiont supplies the
association organic carbon from photosynthesis,
and the mycobiont ensures protection, and
regulates the supply of minerals and water.
• The nutritional exchange between partners is
probably much more complex than exchange of
water and minerals for organic carbon.
• For example, cyanobacteria commonly fix
dinitrogen.
• Thus the lichens containing cyanobacteria obtain
their organic nitrogen directly.
• The source of nitrogen in the absence of
fixation is presumed to be the air especially
following the reduction of dinitrogen by
lightening.
• In filamentous and gelatinous lichens, the
photobiont may form a larger proportion of
the lichen.
• In gelatinous lichens, the cyanobacterium
exudes a polysaccharide that functions to
absorb and retain water, a function normally
attributed to the fungus.
• While each lichen appears to be a homogenous structure, a
single thallus may contain several different species of
photobiont and fungus.
• Even in a single species of fungus, several different genotypes
may coexist in the one thallus.
• Thus, description of the form and function of lichens are
generalisations that need not reflect the complexity within a
single unit.
 Habit and Habitat of Lichen
• There is about 400 genera and 15,000 species of lichens,
widely found in different regions of the world.
• The plant body is thalloid; generally grows on bark of trees,
leaves, dead logs, bare rocks etc., in different habitat.
• They grow luxuriantly in the forest areas with free or less
pollution and with abundant moisture.
• Some species like Cladonia rangiferina
(reindeer moss) grows in the extremely cold
condition of Arctic tundras and Antarctic
regions.
• In India, they grow abundantly in Eastern
Himalayan regions.
• They do not grow in the highly polluted
regions like Industrial areas.
• The growth of lichen is very slow.
• Depending on the growing region, the lichens
are grouped as:
(i) Corticoles
• Growing on bark of trees, mainly in the sub-
tropical and tropical regions.
(ii) Saxicoles
• Growing on rocks, in cold climate
(iii) Terricoles
• Growing on soil, in hot climate, with sufficient
rain and dry summer
• The algal members belong to Chlorophyceae (Trebouxia,
Trentepohlia, Coccomyxa etc.), Xanthophyceae (Heterococcus)
and also Cyanobacteria (Nostoc, Scytonema etc.)
• The fungal members mainly belong to Ascomycotina and a few
to Basidiomycotina. Among the members of Ascomycotina,
Discomycetes are very common; producing huge apothecia,
others belong to Pyrenomycetes or Loculoascomycetes
 Nature of Association of Lichens
• There are three views regarding the nature of association of
algal and fungal partners in lichen
1. According to some scientists, the fungus lives parasitically,
either partially or wholly, with the algal components.
• This view gets support for the following evidences:
• (i) Presence of haustoria of fungus in algal cells of some lichen.
• (ii) On separation, the alga of lichen is able to live
independently, but the fungus cannot survive
2. According to others, they live symbiotically,
where both the partners are equally benefitted.
• The fungal member absorbs water and
mineral from atmosphere and substratum,
make available to the alga and also protects
algal cells from adverse conditions like tem-
perature etc.
• The algal member synthesises organic food
sufficient for both of them
• 3. According to another view, though the relationship is
symbiotic, the fungus shows predominance over the algal
partner, which simply lives as subordinate partner.
• It is like a master and slave relationship, termed helotism.
 Classification of Lichens
• Natural system of classification is not available for lichens.
• They are classified on the nature and kinds of fruit bodies of
the fungal partner
• Based on the structure of fruit bodies of fungal partners,
Zahlbruckner (1926) classified lichens into two main groups:
• 1. Ascolichens:
• The fungal member of this lichen belongs to Ascomycotina.
• Based on the structure of the fruit body, ascolichens are divi-
ded into two series:
• (i) Gynocarpeae:
• The fruit body is disc-shaped i.e., apothecial type. It is also
known as Discolichen (e.g., Parmelia).
(ii) Pyrenocarpeae:
• The fruit body is flask-shaped i.e., perithecial
type.
• It is also known as Pyrenolichen (e.g.,
Dermatocarport).

• 2. Basidiolichen:
• The fungal member of this lichen belongs to
Basidiomycotina e.g., Dictyonema, Corella.
• Later, Alexopoulos and Mims (1979) classified
lichens into three main groups:

i. Basidiolichen:
• The fungal partner belongs to Basidiomycetes
e.g., Dictyonema.

ii. Deuterolichen:
• The fungal partner belongs to Deuteromycetes.

iii. Ascolichen:
• The fungal partner belongs to Ascomycetes e.g.,
Parmelia, Cetraria.
 Structure of Thallus in Lichens
• The plant body of lichen is thalloid with different shapes.
• They are usually grey or greyish green in colour, but some are red,
yellow, orange or brown in colour.

• A. External Structure of Thallus:

• Based on the external morphology, general growth and nature of
attachment, three main types or forms of lichens (crustose, foliose
and fruticose) have been recognised. Later, based on detailed
structures
 Morphology of Lichens
• Lichens may be flat structures that are appressed to a surface,
leafy forms that have multiple attachments to their surface, or
foliar forms that have a single attachment to a surface with the
larger part either erect or hanging from the attachment.
• The fungus primarily determines the form, but the photobiont
may also influence it.
• Thus, complex forms are possible within one lichen.
• Hawksworth and Hill (1984) categorised the
lichens into five main types of morphological
thallus growth forms:
1. Leprose
• This is the simplest type, where the fungal
mycelium envelops either single or small
cluster of algal cells.
• The algal cell does not envelop all over by
fungal hyphae. The lichen appears as powdery
mass on the substratum, called leprose e.g.,
Lepraria incana.
• 2. Crustose
• These are encrushing lichens where thallus is
inconspicuous, flat and appears as a thin layer
or crust on substratum like barks, stones,
rocks etc.
• They are either wholly or partially embedded
in the substratum, e.g., Graphis, Lecanora,
Ochrolechia, Strigula, Rhizocarpon, Verrucaria,
Lecidia etc.
• The simplest form of lichen is a crust on the surface.
• Crustose lichens are highly variable in anatomy.
• However, they all tend to be adnate or attached directly to
their surface.
• Their growth tends to be radiate, in that the mitotic regions are
at the margins, and the centre is more likely to be dying.
• 3. Foliose
• These are leaf-like lichens, where thallus is
flat, horizontally spreading and with lobes.
• Some parts of the thallus are attached with
the substratum by means of hyphal
outgrowth, the rhizines, developed from the
lower surface e.g., Parmelia, Physcia,
Peltigera, Anaptychia, Hypogymnia, Xanthoria,
Gyrophora, Collema, Chauduria etc.
• Foliose lichens have a sheet-like structure, and are attached to
their surface by root-like rhizines.
• The thallus is highly differentiated, with the lower surface
being an absorptive tissue and the photobionts being held in a
manner that maximises photosynthesis.
• Commonly, the upper surface is fungal tissue, with the mid-
layer containing the photobiont.
• Growth takes place at the margins, and these tend to be lobed.
• These are some of the largest and perhaps most complex
lichens. The thallus generally forms flat, leaf-like lobes, with
differentiated layers of tissue, the upper and lower cortices,
forming the upper and lower surfaces.
• The lobes are commonly, but not always, appressed to the
substrate surface, but can be lifted away. The lower cortex
is often differently coloured, frequently brown or black and
usually bears rhizines. In Peltigera the lower surface is
ecorticate.

• In foliose lichens with multiple branches of the thallus that

may stand away from the substrate, the differentiated
lower cortex distinguishes them from fruticose lichens, e.g.
Evernia prunastri, in which the thallus lobes are white
beneath, and Pseudevernia furfuracea, in which the
undersides are black when mature.
Parmotrema perlatum upper surface
4. Fruticose (Frutex, Shrub)
• These are shrubby lichens, where thalli are
well developed, cylindrical branched, shrub-
like, either grow erect (Cladonia) or hang from
the substratum (Usnea).
• They are attached to the substratum by a
basal disc e.g., Cladonla, Usnea, Letharia,
Alectonia
• A fruticose lichen is a form of lichen fungi that is
characterized by a coral-like shrubby or bushy
growth structure. It is composed of a thallus and
a holdfast. It is formed from a symbiotic
relationship of a photobiont such as
cyanobacteria and two mycobionts.
• Fruticose lichen is composed of a complex
vegetation structure, and characterized by an
ascending, bushy or pendulous appearance.
• While lichen communities are mainly controlled
by water and light, vegetative dispersal and
filamentous growth in fruticose lichen is often
associated with areas of low elevation. Fruticose
lichens can endure high degrees of desiccation.
• Fruticose or umbilicate lichens are attached to their surface by
a holdfast.
• The main body of the lichen is either erect or pendulous, and
commonly highly branched.
• Growth takes place at the ends of the “stems” and may be
quite complex
Fruticose Lichen Letharia vulpina
• 5. Filamentous
• In this type, algal members are filamentous and well-
developed.
• The algal filaments remain ensheathed or covered by only a
few fungal hyphae.
• Here algal member remains as dominant partner, called
filamentous type, e.g., Racodium, Ephebe, Cystocoleus etc.)
 Internal Structure of Lichen Thallus
• Based on the distribution of algal member inside the thallus,
the lichens are divided into two types; Homoisomerous or
Homomerous and Heteromerous.
• 1. Homoisomerous
• Here the fungal hyphae and the algal cells are more or less
uniformly distributed throughout the thallus.
• The algal members belong to Cyanophyta.
• This type of orientation is found in crustose lichens.
• Both the partners intermingle and form thin outer protective
layer, e.g., Leptogium, Collema etc
• 2. Heteromerous
• Here the thallus is differentiated into four distinct layers upper
cortex, algal zone, medulla, and lower cortex.
• The algal members are restricted in the algal zone only.
• This type of orientation is found in foliose and fruticose lichens
e.g., Physcia, Parmelia etc.
 Reproduction in Lichens
• Many lichens reproduce asexually either by vegetative
reproduction or through the dispersal of diaspores containing
algal and fungal cells
• Soredia (sing. Soredium) are small groups of algal cells
surrounded by fungal filaments that form structures called
soralia from which the soredia can be dispersed by wind
• Another form of diaspores are isidia, elongated outgrowths
from thallus that break off for mechanical dispersal e.g
Fruticose lichens
• Manparther before a functional lichen can
formy lichen fungi also reproduce sexually in a
manner typical of fungi producing spores that
are presumably the result of sexual fusion and
meiosis
• Following dispersal, such fungal spores must
meet with a compatible algal
• Spores are produced in spore-producing
bodies (Ascocarp)
• Five common spore body types are
Apothecium, perithecium and Cleistothecium,
gymnothecium and pseudothecium
 (i) Apothecium
• This is a wide, open saucer-shaped, or cup shaped fruit body.
• It is sessile and fleshy' Structure of Apothecium chiefly consist of
three parts, namely hymenium (upper concave surface),
hypothecium, and excipulum.
• The asci (spore-bearing cells) are present in the hymenium layer.
• The asci are freely exposed at maturity.
• An example are the members of Dictyomycetes.
• Here the fertile layer is free, so that many spores can be dispersed
simultaneously.
• E.g The genera Helvella and Gyromitra
 (ii) Perithecium
• These are flask shaped structures opening by a pore or ostiole
(short papilla opening by a circular pore) through which the
ascospores escape.
• The ostiolar canal may be lined by hair-like structures called
periphyses.
• The unitunicate asci are usually cylindrical in shape, borne on a
stipe (stalk), released from a pore, developed from the inner wall of
the perithecium and arise from a basal plectenchyma-centrum.
• Examples are members of Sphaeriales and Hypocreales. Perithecia
are also found in Xylaria (Dead Man’s Fingers, Candle Snuff) and
Nectria.
• Perithecia of Nectria
 (iii) Cleistothecium
• A cleistothecium is a globose, completely closed fruit body
with no special opening to the outside.
• The ascomatal wall is called peridium and may typically
consists of densely interwoven hyphae or
pseudoparenchyma cells.
• It may be covered with hyphal outgrowth called
appendages.
• The asci are globose, deliquescent, and scattered
throughout the interior cavity i.e. as in Eurotium or arising
in tufts from the basal region of ascocarps as in Erysiphe.
• In this case the ascocarp is round with the hymenium
enclosed, so the spores do not automatically get released,
and fungi with cleistothecia have had to develop new
strategies to disseminate their spores.
• The truffles, for instance, have solved this
problem by attracting animals such as wild
boars, which break open the tasty ascocarps
and spread the spores over a wide area.
• Cleistothecia are found mostly in fungi that
have little room available for their ascocarps,
for instance those that live under tree bark, or
underground like truffle
 (iv) Gymnothecium

• Similar to a cleistothecium, a gymnothecium is a completely

enclosed structure containing globose or pear-shaped,
deliquescent asci.
• However, unlike the cleistothecium, the peridial wall of a
gymnothecium consists of a loosely woven "tuft" of hyphae,
often ornamented with elaborate coils or spines.
• Examples are the Gymnoascus, Talaromyces and the
dermatophyte Arthroderma.
 (v) Pseudothecium
• This is similar to a perithecium, but the asci are not regularly
organised into a hymenium and they are bitunicate, having a
double wall that expands when it takes up water and shoots
the enclosed spores out suddenly to disperse them.
• Example species are Apple scab (Venturia inaequalis) and the
horse chestnut disease Guignardia aesculi.
Ecological Importance of Lichens
1. Used as food by some wild animals especially under dry conditions, e.g
Squirrel, Reindeers.
2. Some are able to fix the nutrient Nitrogen due to their Cyanobacterial
partners.
3. Usefull in soil formation through their Rhizospheres which penetrate and
crack rocks thereby forming soils.
4. May offer habitat to some wild fauna such as birds or small animals. They
also provide good camouflage to some insects and small animals.

Economic Importance of Lichens

1. Bioindicators of air pollution. Different lichens are known to be sensistive
to different types of air pollutants and thus their presence or absence in a
particular habitat could be an indication of air pollution by particular
pollutants.
2. Source of dyes used in colouring fabrics or yarns. Lichens have different
types of pigments in their fungal partners which make them bear different
colours, i.e yellow, orange, grey, brown, green etc. These pigments are
normally extracted and used in colouring of fabrics.
3. Some Lichens are known to produce some organic acids and essential oils
that are used in making perfumes. Oakmoss is an example of Lichen species
whose oils is used in making very expensive perfumes.
4. Some Lichens produce organic acid that are effective as antibiotics. Some
are also known to antiviral.
 BRYOPHYTES AND PTERIDOPHYTES
• Bryophyta is a Greek term (Bryon = moss, phyton
= plant), used as a collective name for such lower
plants which have amphibious habitat.
• They are able to live on land as well as in water.
The group, therefore, goes well with the name of
“amphibians of plant kingdom” owing to the
amphibious habitat of these plants.
• Pteridophytes are non-flowering but spore-
bearing plants.
• They constitute the most primitive group of
plants with a vascular system (xylem and
phloem).
 Bryopytes
• Plant scientists recognize two kinds of land
plants, namely, bryophytes, or nonvascular
land plants and tracheophytes,or vascular land
plants
• Bryophytes are small, herbaceous plants that
grow closely packed together in mats or
cushions on rocks, soil, or as epiphytes on the
trunks and leaves of forest trees.
• Known as Lower non vascular plants
• They are a group of simple and primitive plants. They grow in
moist and shady places forming a dense mass or mat on damp
soil, rock, logs and other substrata.
• They are basically terrestrial forms, but require the presence of
water for completing the life cycle. Hence, they are commonly
described as 'amphibians of the plant kingdom'.
• Bryophytes are distinguished from
tracheophytes by two important characters.
First, in all bryophytes the ecologically
persistent, photosynthetic phase of the life
cycle is the haploid, gametophyte generation
rather than the diploid sporophyte;
• Bryophyte sporophytes are very short-lived,
are attached to and nutritionally dependent
on their gametophytes and consist of only an
unbranched stalk, or seta, and a single,
terminal sporangium
 Reproduction in Bryophytes
• The bryophytes reproduce both by vegetative and sexual methods.
• Vegetative reproduction is by special structures such as tubers,
gemmae, etc.
• In sexual reproduction, the male reproductive organ is called
antheridium which produces the male gametes called antherozoids.
The female reproductive organ, the archegonium produces the
non-motile egg.
• The reproductive organs (both male and female) are borne on the
dorsal surface of the thallus in the median groove.
 Alternation of Generation
• It is a process where the plant passes through two successive
generations – gametophytic with haploid (n) chromosomes and
sporophytic with diploid (2n) chromosomes- to complete its life
history.
• The sex organs (the antheridia and archegonia) develop on the
gametophyte. These produce the gametes which fuse (fertilization)
to form a zygote.
• The zygote divides to form an embryo which is not liberated but is
retained within the archegonium where it develops into a
sporophyte.
• The sporophyte is physiologically and physically dependent on the
gametophyte. The sporophyte consists of a foot, seta, and capsule.
• In the capsule, spores are formed after meiosis.
• These spores (called meiospores), are all similar in shape, size and
ratio, and hence the plant is homosporous. The non-motile spores
germinate directly into new gametophytic plants.
 Classification of Bryophytes
• The bryophytes are divided into three classes;

Class: Hepaticopsida (Marchantiophyta-Liverworts)

– these include Marchantia and Riccia.
• Gametophytic plant body is either thalloid or foliose. If foliose, the
lateral appendages (leaves) are without mid-rib. Always
dorsiventral.
• Rhizoids without septa.
• Each cell in the thallus contains many chloroplasts; the chloroplasts
are without pyrenoi.
• Sex organs are embedded in the dorsal surface.
• Sporophyte may be simple (e.g., Riccia) having only a capsule, or
differentiated into root, seta and capsule (e.g., Marchantia, Pallia
and Porella etc.)
• Capsule lacks columella.
• Constitute more than 8,000 species of small, nonvascular,
spore-producing land plants
• They include the thallose liverworts that show branching,
ribbonlike gametophytes and the leafy liverworts (mainly in
the order Jungermanniales).
• Liverworts are distributed worldwide, though
most commonly in the tropics.
• Thallose liverworts grow commonly on moist soil
or damp rocks, while leafy liverworts are found in
similar habitats as well as on tree trunks in damp
woods.
• The thallus of thallose liverworts resembles a
lobed liver—hence the common name liverwort
(“liver plant”).
• Filamentous structures called rhizoids anchor
most liverworts to their substrata, except for the
few genera that are aquatic.
• Constitute more than 8,000 species of small, nonvascular,
spore-producing land plants
• They include the thallose liverworts that show branching,
ribbonlike gametophytes and the leafy liverworts (mainly in
the order Jungermanniales).
• Liverworts are distributed worldwide, though
most commonly in the tropics.
• Thallose liverworts grow commonly on moist soil
or damp rocks, while leafy liverworts are found in
similar habitats as well as on tree trunks in damp
woods.
• The thallus of thallose liverworts resembles a
lobed liver—hence the common name liverwort
(“liver plant”).
• Filamentous structures called rhizoids anchor
most liverworts to their substrata, except for the
few genera that are aquatic.
Class: Anthocerotopsida (Anthocerotophyta)
(Hornworts)
e.g. Anthoceros.
• Gametophytic plant body is simple, thalloid;
thallus dorsiventra without air cambers, shows no
internal differentiation of tissues.
• Hornworts may be found worldwide, though they
tend to grow only in places that are damp or
humid.
• Some species grow in large numbers as tiny
weeds in the soil of gardens and cultivated fields.
• Large tropical and sub-tropical species of
Dendroceros may be found growing on the bark
of trees.
• The group's common name "hornwort" refers to
the tall narrow sporophytes which are embedded
in the top of the plant.
• As in other bryophytes, the sporophyte remains
attached to its parent gametophyte throughout
its life, but unlike these other plants, the
sporophyte continues to grow throughout its life
• This happens as a group of cells at the base of the
horn divide repeatedly.
• This continuous growth from a near-basal
meristem is unique among plants to hornworts.
• The plant body of a hornwort is a haploid gametophyte stage.
• This stage usually grows as a thin rosette or ribbon-like thallus
between one and five centimeters in diameter.
• Each cell of the thallus usually contains just one chloroplast.
• Many hornworts develop internal mucilage-filled cavities when
groups of cells break down.
• These cavities are invaded by photosynthetic cyanobacteria,
especially species of Nostoc.
• Such colonies of bacteria growing inside the thallus give the
hornwort a distinctive blue-green color.
• The horn-shaped sporophyte grows from an archegonium
embedded deep in the gametophyte.
• The sporophyte of a hornwort is unusual in that it grows from a
meristem near its base, instead of from its tip the way other
plants do.
• Unlike liverworts, most hornworts have true stomata on their
sporophyte as mosses do.
• The exceptions are the genera Notothylas and
Megaceros, which do not have stomata.
• When the sporophyte is mature, it has a
multicellular outer layer, a central rod-like
columella running up the center, and a layer of
tissue in between that produces spores and
pseudo-elaters.
• The pseudo-elaters are multi-cellular, unlike the
elaters of liverworts.
• They have helical thickenings that change shape
in response to drying out; they twist and thereby
help to disperse the spores.
• When a spore germinates, it produces a flat
thalloid plant with a greasy blue-green color
and odd morphology.
• The best way to recognize a hornwort, and
especially to tell it apart from a liverwort or
fern gametophyte, is to look at the plant
under a low-power microscope; hornworts
will generally have a single large chloroplast
per cell.
• Hornworts also possess a large number of unique
traits that are not found in any other land plants.
These include the following:
• (1) Zygote and sporophyte development: the
orientation of the first zygote division is
longitudinal, unlike in other land plants except
leptosporangiate ferns.
• The sporophyte differs from mosses and
liverworts in that it lacks a seta and continuously
and progressively produces spores upwardly from
a basal meristem, being essentially a growing
sporangium.
• 2) Sporophytes bear stomata, which may be
homologous to those of tracheophytes
• (3) Chloroplast: it is the only extant land plant
lineage, together with some lycophytes that
has a single chloroplast (or just a few) per cell.
• The chloroplast of several hornwort species
may contain a pyrenoid, a structure also found
in many streptophyte algae and other algal
lineages but not in other land plants.
• 4) Hornwort plastid genomes have also one of
the highest RNA editing rates amongst land
plants
• (5) Symbiosis: hornworts establish symbiotic relationships with
endophytic cyanobacteria and various fungal partners
(mycorrhiza).
• A small number of studies have provided insight into hornwort
morphology and growth but detailed molecular and genetic
studies examining hornwort development are lacking.
Class: Bryopsida (Musci)-Bryophyta (Mosses)
e.g., Funaria, Moss, etc.
• Mosses are small (a few centimeters tall) herbaceous (non-
woody) plants that absorb water and nutrients mainly through
their leaves and harvest carbon dioxide and sunlight to create
food by photosynthesis.
• They differ from vascular plants in lacking water-bearing xylem
tracheids or vessels.
• As in liverworts and hornworts, the haploid gametophyte
generation is the dominant phase of the life cycle.
• This contrasts with the pattern in all vascular plants (seed plants
and pteridophytes), where the diploid sporophyte generation is
dominant.
• Mosses reproduce using spores, not seeds and have no flowers.
• Moss gametophytes have stems which may be
simple or branched and upright or prostrate.
• Their leaves are simple, usually only a single layer
of cells with no internal air spaces, often with
thicker midribs.
• They do not have proper roots, but have
threadlike rhizoids that anchor them to their
substrate.
• Mosses do not absorb water or nutrients from
their substrate through their rhizoids.
• They can be distinguished from liverworts
(Marchantiophyta or Hepaticae) by their multi-
cellular rhizoids.
• Spore-bearing capsules or sporangia of mosses are borne
singly on long, unbranched stems, thereby distinguishing them
from the polysporangiophytes, which include all vascular
plants.
• The spore-bearing sporophytes (i.e. the diploid multicellular
generation) are short-lived and dependent on the
gametophyte for water supply and nutrition
 Life cycle of mosses
• The life cycle of mosses shows two distinct phases namely a
haploid gametophytic phase and a diploid sporophytic
phase alternating with each other.
• The adult plant body represents the gametophyte.
• A short-lived sporophyte occurs as a parasite on the
gametophyte.
• Mosses spread in multiple ways, but unlike flowering
plants, they depend on moisture to sexually reproduce.
• Mosses reproduce by spores, which are analogous to the
flowering plant's seed; however, moss spores are single
celled and more primitive than the seed
• Vegetative reproduction may sometimes occur by
fragmentation. However, sexual reproduction is common
and is of oogamous type.
(i) The Gametophyte

• Spores are housed in the brown capsule that sits on the

seta.
• As the spores ripen they are dispersed from the
capsule, and some land in areas where there is enough
moisture for them to grow.
• From the one-celled spore, a highly branched system of
filaments, called the protonema (Pl. Protonemata),
develops.
• Cell specialization occurs within the protonema to form
a horizontal system of reddish-brown, anchoring
filaments, called caulonemal filaments and upright,
green filaments, called chloronemal filaments.
• Each protonema, which superficially resembles a
filamentous alga, can spread over several centimeters
to form a fuzzy green film over its substrate.
• As the protonema grows, some cells of the caulonemal
filaments specialize to form leafy buds that will
ultimately form the adult gametophyte shoots
• Protonema is a transitory stage in the life of a moss,
but from the protonema grows the gametophore
("gamete-bearer") that is structurally differentiated
into stems and leaves.
• Numerous shoots typically develop from each
protonema so that a single spore can give rise to a
whole clump of moss plants.
• Each leafy shoot continues to grow apically,
producing leaves in spiral arrangement on an
elongating stem.
• In many mosses the stem is differentiated into
a central strand of thin-walled water-
conducting cells, called hydroids, surrounded
by a parenchymatous cortex and a thick-
walled epidermis
• The leaves taper from a broad base to a
pointed apex and have lamina that are only
one-cell layer thick.
• A hydroid-containing midvein often extends
from the stem into the leaf.
• Near the base of the shoot, reddish-brown,
multicellular rhizoids emerge from the stem to
anchor the moss to its substrate.
• Water and mineral nutrients required for the
moss to grow are absorbed, not by the
rhizoids,but rather by the thin leaves of the
plant as rain water washes through the moss
cushion.
• Mosses produce large, multicellular sex organs
for reproduction.
• From the tips of the gametophore stems or
branches develop the sex organs of the mosses.
• The female organs are known as archegonia (sing.
archegonium) and are protected by a group of
modified leaves known as the perichaetum
(plural, perichaeta).
• The archegonia are small flask-shaped clumps of
cells with an open neck (venter) down which the
male sperm swim.
• The male organs are known as antheridia (sing.
antheridium) and are enclosed by modified
leaves called the perigonium (pl. perigonia).
• The antheridia have a club-shaped body and a stalk. They
produce flagellated male gametes called antherozoids or
sperms
• The surrounding leaves in some mosses form a splash cup,
allowing the sperm contained in the cup to be splashed to
neighboring stalks by falling water droplets.
• Mosses can be either dioecious or monoecious.
• In dioecious mosses, male and female sex organs are borne on
different gametophyte plants.
• In monoecious (also called autoecious) mosses, both are borne
on the same plant.
• In the presence of water, sperm from the antheridia swim to
the archegonia and fertilization occurs to form a zygote which
develops into a diploid sporophyte.
• The sperm of mosses is biflagellate, i.e. they have two flagellae
that aid in propulsion.
• Since the sperm must swim to the archegonium, fertilization
cannot occur without water.
• Some species (for example Mnium hornum or several species
of Polytrichum) keep their antheridia in so called 'splash cups',
bowl-like structures on the shoot tips that propel the sperm
several decimeters when water droplets hit it, increasing the
fertilization distance.
(ii) The Sporophyte
• Zygote represents the first cell of the sporophytic phase.
• It divides and develops into a sporophytic plant body called
sporogonium.
• The wall of the archegonialventer forms a protective
covering to the sporogonium, called calyptra.
• The sporophyte body (sporangium) comprises a long stalk,
called a seta, and a capsule capped by a cap called the
operculum.
• The capsule and operculum are in turn sheathed by a
haploid calyptra which is the remains of the archegonial
venter
• It remains attached throughout its life to the gametophytic
host with the help of foot.
• It absorbs nutrients directly from the gametophyte.
• The sporogonium produces haploid spores
(meiospores) which get released from the capsule.
• The calyptra usually falls off when the capsule is
mature. Within the capsule, spore-producing cells
undergo meiosis to form haploid spores, upon which
the cycle can start again. The mouth of the capsule is
usually ringed by a set of teeth called peristome. This
may be absent in some mosses
• The spores represent the first cells of gametophytic
generation.
• They germinate to produce new gametophytes either
directly or through a juvenile stage called protonema.
• It takes about a quarter to half a year for the
sporophyte to mature.
 Life and Growth forms in mosses
• Life form is a useful concept in bryophyte ecology because of
the "exceptionally high dependence of bryophytes on transient
external water supplies.
• For bryophytes, it is not the individual that forms the ecological
unit, but rather the clonal or colonial life form
• The life form is so constructed as to minimize evaporative loss
while maximizing photosynthetic light capture
• Growth form is the overall character of a plant and explains it
can only be determined by detailed morphological analysis.
• It is a purely a morphological term
• Life form is more encompassing and describes the result of life
conditions, including growth form, influence of environment,
an assemblage of individuals
• Life form embodies all the selection pressures that are brought
to bear upon a species.
• It is the organization of a plant in correspondence with its life
conditions. Hence, life forms are genetically determined.
• Growth forms are influenced by the environment.
• There three known moss growth forms: Acrocarpous (orthotropic)
mosses, Pleurocarpous (plagiotropic) mosses and Cladocarpous mosses.
• (a) Acrocarpous mosses (acros: highest; karpos: fruit)
• -Are those which grow upright as individual plants, either separately or
very close together to form a turf, tuft, or cushion.
• In this group, the leaves nearly always have a costa, and the sporophyte
grows from the tip or highest part .
• Some of the common acrocarpous genera are Dicranum, Tortella, Barbula,
Mnium, Bryum, Polytrichum.
• Perichaetial leaves not usually visible
• The Acrocarpous (orthotropic) mosses can be
further
• divided into the protonema mosses with short
or non-existent shoots that wither after the
sporophyte is
• produced, and turf mosses with upright
shoots that bear new shoots after the
sporophyte forms and subsequently bear
further archegonia and more sporophytes;
these new growths are the innovations
Polytrichastrum ohioense is a moss with an acrocarpous
growth form
• The same species may exhibit more than one
growth form. For example, in some
populations Hylocomium splendens exhibits
monopodial growth (single central axis with
apical growth).
• However, some populations can continue by
sympodial growth (growth produced by lateral
buds just behind apex).
(b) Cladocarpous: is distinct from pleurocarpy,
with perichaetia terminal on lateral branches
and with juvenile leaf development similar to
that on vegetative branches; perichaetial
branches have lateral primordia that
potentially develop subperichaetial branches.
• It is a type of pleurocarpy having sporophytes
borne terminally on short lateral branches, as
in Fontinalis
(c) Pleurocarpous mosses (pleura: side; karpos: fruit).
• Are the prostrate or creeping plants on ground, wood, or rock surfaces.
• Some of these add new intertwining or overlaid growth each year to form
mats.
• The leaves are usually without costa and the sporophyte grows from the
side of one of the branches of the plant
• The perichaetial leaves are longer and often quite different from the
regular leaves.
• Many of the pleurocarpous mosses are known popularly as "feather
mosses" and "fern mosses", including the genera Hypnum, Thuidium,
Amblystegium, Brachythecium, Plagiothecium, Hylocomium.
• The Pleurocarpous (plagiotropic) mosses
include thread mosses e.g. Leskeaceae , some
Amblystegiaceae, with little difference
between the main stem and lateral branches,
comb mosses (e.g. Hypnaceae,
Brachytheciaceae, Meteoriaceae ), with a
strong main shoot with many simple or
branched lateral branches, and the creeping-
shoot mosses (e.g. Leucodon, Antitrichia,
Climaciaceae, Hypnodendraceae), with
rhizomatous main shoots that give rise to
upright main shoots.
 Comparison with other plant groups
• Bryophytes resemble Algae in their structure and mode of
life. They are probably more akin to green Algae
(Chlorophyceae) in the features listed in the box below:
• -The plant body is thalloid
• -Plants are autotrophic
• -Reserve food material is starch
• -Vascular tissues are absent
• -True roots are absent
• -Gametophyte is the dominant phase of the life cycle
• -Spermatozoids are motile and flagellated
• -Cell wall is made up of cellulose
• The above similarities put Bryophytes near green Algae
(Chlorophyceae) but there are certain differences which
put the two groups apart. These differences include:
• -The Bryophytes are mostly terrestrial with preference to
damp shady places, whereas the Algae are mostly aquatic.
• -Asexual reproduction is completely absent in Bryophytes
(but vegetative reproduction is sometimes common). In
algae, Asexual reproduction is the common method during
which zoospores are formed. Other types of spores are
aplanospores and akinetes.
• -In Bryophytes, sexual reproduction is invariably of
Oogamous type; while in Algae, sexual reproduction varies
from Isogamy, Anisogamy, to Oogamous types.
• -In Bryophytes, the zygote germinates into sporophyte
immediately after fertilization without being liberated
and undergoing any resting period. In algae, the zygospore
is liberated and undergoes a resting period.
• -The Bryophytes exhibit sharply defined heteromorphic
type of alternation of generation. In Bryophytes, the
sporophyte, though well-developed is dependent physically
and nutritionally on the gametophyte. In Algae, the
sporophyte constitutes the reduced and independent stage
in the life cycle.
 Importance of Bryophytes
• 1. Ecological importance:
• The liverworts, mosses and lichens are supposed
to be the pioneers in establishing vegetation
where other vegetation seems to be practically
impossible.
• They colonize the barren rocks and exposed areas
of hills, and make them suitable for growing
angiospermic and other plants by depositing
humus soil and plant debris.
• In the beginning the forms and grasses grow, and
ultimately shrubs and trees also establish, and
the whole area converts into dense wood.
• However the Sphagnum plants are of great
ecological importance.
• When these plants establish themselves in
some lake or other areas full of water, sooner
or later they cover the whole surface of the
water.
• Due to deposition of plant debris the surface
may be raised
• The Sphagnum plants along with other
hydrophytes form a dense surface covering
over the water below.
• This covering gives the appearance of the soil
from the surface.
• These areas are known as quacking bogs.
Later on these bogs are converted into
swamps.
• Ultimately these swamps are replaced by the
forest growth of mesophytic type.
• A few bryophyotes play an important role in checking the soil
erosion.
• They are capable of holding the soil by their extensive carpets,
and prevent the soil erosion to some extent
• In the tropical rainforest, 'moss balls' form in
the higher elevations.
• Here they can absorb great quantities of rain
and release water slowly into the atmosphere
or ground.
• These balls support numbers of invertebrates
and smaller organisms.
• Basically these layers have created a second
'ground' or terra high in the tropical canopy, a
world recently discovered with walkways and
ladders that span the canopy.
• 2. Packing material
• Most of the mosses are used as packing
material after being dried.
• They make a fairly good packing material in
the case of glass ware and other fragile goods.
• Especially the dried peat mosses (Sphagnum
spp.) are used to pack bulbs, cuttings and
seedlings for shipment.
3. Used in seed beds:
• Since the peat mosses have remarkable power
to absorb and hold water like a sponge, they
are extensively used in seed beds and green
houses to root cutting.
• The peat mosses (Sphagna) are also used to
maintain high soil acidity required by certain
plants.
4. As a source of fuel
• The peat is also a potential source of coal.
• Dried peat may be used as fuel.
• In Ireland, Scotland and other European
countries the peat is used for fuel.
• In colder parts of the world where peat
reaches its greatest development, the lower
layers of peat become carbonized, and after
the ages have passed, becomes available to
human kind in the form of coal.
5. Absorbent bandages
• The Sphagnum plants are slightly antiseptic and
possess superior absorptive power.
• On account of these properties they may be used
for filling absorbent bandages in place of cotton,
in the hospitals.
• Because of its absorbent and antiseptic
properties it was used for dressings during World
War I, to make pillows for resting of wounded
members for soldiers transported to hospitals
from battlefields, and recently as filling material
for sanitary napkins.
6. Construction
• Wrapping and wall paper was made from
Sphagnum in the early 1900's
• More recently, sphagnum has been used as
construction material
• Peatcrete, a mixture of Sphagnum, Portland
cement and water is commonly used in
mixtures of soil to loosen compact soils and
increase water-holding capacity.
7. Landscaping
• In Japan, mosses have traditionally been used
as base cover for bonsai (an ornamental tree
or shrub grown in a pot and artificially
prevented from reaching its normal size).
• The art of growing ornamental, artificially
dwarfed trees or shrubs in small terraria, in
miniature landscapes and in the designs of the
famous Japanese gardens.
8. Traditional medicine
• In folk medicine, particularly in China close to 40
species are used in various remedies.
• Mosses have been used to treat cardiovascular
diseases, nervous postration, in poultice to treat
skin rashes, burns, infections and bites.
• A chemical analysis of the moss Rhodobryum
giganteum indicates that it contains volatile oils,
lactones and amino acids.
• An extract of this moss tested on white mice
increased by 30% the rate of blood flow in the
aorta