LYCOPODIUM

Sub- Division-
Lycopsida
Order-
Lycopodiales
Family-
Lycopodiaceae
 LYCOPODIUM
Distribution and Occurrence
• Commonly called:
– Club moss
– Ground pine
– Trailing over green
• Represented by 400 species, worldwide in
distribution
• In India, represented by 33 species
• Eg.- L.cernuum, L. clavatum, L.serratum,
L.volubile, L. inundatum, L. phlegmaria, L. selago
 Habit
Found in open woodlands, in moist and acidic
soils
Epiphytic eg.- L. phlegmaria
Erect and shrubby eg.- L. clavatum
Creeping forms eg.- L. cernuum- Gives out
erect branches at intervals
Erect branches highly branched and bushy
 Lycopodium clavatum Lycopodium volubile

Lycopodium phlegmaria Lycopodium

serratum
Lycopodium inundatum
 Lycopodium cernuum

Stem- Dichotomously branched- branches grow

equally or unequally
 LYCOPODIUM
External Morphology
• Plant body is the sporophyte
• Differentiated into root, stem and leaves
• Considering the variation in habit of different
species, Pritzel (1900) divided the genus
Lycopodium into two sub-genera:
– Urostachya
– Rhopalostachya
 LYCOPODIUM
UROSTACHYA RHOPALOSTACHYA
• Erect or pendant stem • Prostrate stem with upright
branches
• Stem-Dichotomously branched • Dichotomy noticed in first
• Roots arise only from base of formed branches; later
stem monopodial
• Leaves and sporophylls almost • Adventitious roots along entire
length of stem
same size • Sporophylls smaller than
• Vegetative reproduction by foliage leaves
bulbils • Sporophylls arranged to form
• Eg. L. compactum, L. selago, L. cones or strobili
• Eg. L. cernuum, L. clavatum, L.
phlegmaria densum
 LYCOPODIUM
• Root:
– First formed root is ephemeral. Older plants have
adventitious roots; they arise from the pericycle or
endodermis
• Stem:
– Weak, slender and rhizomatous
– Erect or pendant; creeping in others
– Branched- basically dichotomous; sometimes
monopodial
– Stem and branches covered with leaves
 LYCOPODIUM
Leaf
• Simple, sessile, scale like or small with median
vein- Microphylls
• Spirally arranged and dense on stem
• Decussate or whorled arrangement
 LYCOPODIUM
Leaf
Isophyllous Anisophyllous
• All leaves are of the • Different types of leaves
same type • In L. volubile, 4 rows of
leaves
– 2 lateral rows of large
and falcate leaves
– 1 ventral row of
small hair-like leaves
– 1 dorsal row of
medium sized
acicular leaves
 ANATOMY OF ROOT
• Cross section shows three distinct
regions:
– Epidermis- single layered, thin walled
cells; sometimes with unicellular hairs
– Cortex- Composed of parenchymatous
cells; older roots have thick walled cells
for mechanical support
– Stele- Protostele; monarch (one
protoxylem), diarch (two protoxylem)
or triarch (three protoxylem)
– Xylem is curved and phloem lies in
between the arms of xylem
 Anatomy of Stem
• Three zones:
– Epidermis: Protective layer; cells with thick outer
wall with cuticle
– Cortex: Nature varies with species and stem
diameter
• Homogenous: Cortex is entirely
parenchymatous
• Heterogenous: Three zones: outer chlorenchymatous,
middle parenchymatous and inner sclerenchymatous
– Endodermis – distinct in young stems
– Stele: Protostele
 STELE
• Has only primary xylem and
primary phloem
• Stele is a protostele- there is no pith at
the centre
• Xylem forms the central
core surrounded by phloem
• Xylem is exarch
• 3 types of protostele
– Actinostele- Xylem Star shaped, phloem
between arms of xylem (L. selago,
L.serratum)
– Plectostele- Xylem in the form of plates
(L. clavatum, L. companulatum)
– Mixed protostele- Xylem and phloem
uniformly distributed in T.S. (L. cernuum)
 Anatomy of Leaf
• Leaf is triangular in outline
• Outer epidermis covered by a thick
cuticle
• Stomata are present on both
surfaces- amphistomatous
• In L. companulatum and L. volubile,
stomata are hypostomatous-
confined to lower surface
• Mesophyll is undifferentiated ; made
of chlorophyllous cells
• A median concentric vascular bundle
with xylem surrounded by phloem
 REPRODUCTION
• Vegetative Reproduction
– Gemmae or Bulbils
– Fragmentation
– Formation of Resting Buds
– Formation of Root tubercles
– Formation of Adventitious Buds
• By Spores
– Homosporous condition
 Vegetative Reproduction
• Gemmae or Bulbils
– Lateral outgrowths from the stem, takes the place
of leaves
– Consists of a short reduced axis surrounded by
thick fleshy leaves with stored food material
– Falls on the ground and grows into a new plant
 Vegetative Reproduction
• Fragmentation
– Death and decay of older parts of the stem leads
to separation of younger branches which grow
into new plants
• Formation of resting buds
– Tips of apical buds of rhizome and branches store
food material and is surrounded by a bunch of
leaves. In winter, rest of the plant dies and the
resting bud develops into new plants.
 Vegetative Reproduction
• Formation of root tubercles
– Group of cells with stored food material and
protected by thick walls- germinates into new
plants
• Formation of Adventitious Buds
– In the leaves epidermal cells proliferate near the
base and grow into buds. These buds are capable
of germinating into new plants.
 Strobilus in Lycopodium
• In Urostachya, every leaf on the plant is a
sporophyll
• In Rhopalostachya, the leaves near the apices
bear sporangia and are called sporophylls
• Sometimes sporophylls are distinctly smaller
than the foliage leaves
• Aggregation of sporophylls is called strobilus
• A strobilus has a central axis on which spirally
arranged sporophylls are present
 Position of Sporangium on Sporophyll
• Axillary in position
– L. selago, L. phlegmaria
• On the dorsal side of the sporophyll-
epiphyllous
• L. cernuum, L. clavatum
• Sub-foliar, not on the sporophyll but a little
towards the sporophyll
• L. squarrosum
Sporophyll--

Strobilus--
 Lycopodium - Strobilus,
Sporophylls and
Sporangia

Longitudinal
section of
strobilus

Upper (adaxial) side of sporophyll

 Structure of Sporangium
• Reniform or Kidney shaped
• Has a stalk and a capsule
• Capsule is unilocular
• Sporangial wall is three layered
• Innermost layer is the tapetum
• Inside sporogenous mother cells undergoes
meiosis and produce tetrads of spores
• Sporangia are homosporous
 Structure of the Spore
• Lycopodium is homosporous
• Occur in tetrads
• Tetrahedral in shape with rounded or
semicircular base
• Unicellular and range in size
from 0.03-0.05mm diam.
• Has a triradiate ridge, 2 layers of cell
wall
• Spore wall may be smooth
or variously sculptured
• Has a single haploid nucleus
• Cytoplasm filled with food
reserve material
 Dehiscence of Sporangium
• Elongation of the internodes of strobilus
• Sporophylls spread out exposing sporangia
• A line of cells are differentiated on the sporangial
wall called stomium
• Inner walls of cells of stomium are thick and
lignified
• Exposed sporangia lose water and dry
• Split appears in the stomium and sporangia open
into two valves
• Air disseminates the spores
 Germination of the Spore
• Time taken for germination varies from a few
days to several years after their liberation from
the sporangium
• Quickly germinating spores produce aerial short
lived and green prothalli
– L. cernuum, L. inundatum
• When spores take longer time to germinate, they
get buried under the soil and produce colourless
subterranean prothalli that are large, tuberous
and long lived
 Germination of the Spore
• The spore germinates to produce the
gametophyte
• The gametophyte is called the
prothallus
• It produces the antheridia and
archegonia
• It is free living independent of the
sporophyte
• 2 regions- lower region attaches to
the soil
 Gametophyte
• Three types of Prothalli are present in
Lycopodium
– Cernuum Type
– Clavatum Type
– Phlegmaria Type
 CERNUUM TYPE
• L. cernuum, L. inundatum
• Erect cylindrical body, 2-3 mm long
• Grows on the surface of the ground
• Colourless basal portion buried in the soil
• Lobed generative zone-green and bearing sex organs at
the base of the lobes
• Rhizoids restricted to lower buried portion
• Endophytic fungus is present in the basal portion
• Meristematic tissue present as a rim around upper part
• Prothallus is independent and prepares its own food
Longitudinal section of subterranean
gametophyte of Lycopodium
complanatum

Slide 31
 CLAVATUM TYPE
• Spore germination is delayed for a long time (one to
many years), thus the prothallus has a longer lifespan.
• Here the prothalli are fleshy, non-green, totally
saprophytic and completely subterranean and
perennial in nature.
• Development takes place beneath the surface of the
ground or within a layer of humus.
• The prothalli are large and may be up to 2 centimeters
in length. They may be top-shaped with a convolute
margin or carrot shaped
• The top of the prothallus are lobed and the sex organs
and the growing embryos are located on these lobes.
• All the gametophytic cells are parenchymatous
Second type of Prothallus

Longitudinal section of subterranean

gametophyte of Lycopodium clavatum
 PHLEGMARIA TYPE
• The pro­thalli are aerial but saprophytic in
nature, grow on tree trunks below a coating of
humus.
• This type is found in epiphytic species of
Lycopodium (e.g., L. phlegmaria).
• Here the spore germination is immediate and
the gametophyte grows for only one season.
• The prothallus consists of a short, tuberous
cen­tral part from which a number of
colourless, slender and cylindrical branches
develop in an irregular fashion.
• These branches bear sex organs and they are
usually surrounded by glandular hairs called
paraphysis.
Photosynthetic lobes Gametophyte diversity
Gameto- in Lycopodium spp.
Fungal phyte of L.
portion inundatum

Subterranean Embryo
-Rhizoids gametophyte of L. Antheridium
Archegonium
annotinum

Meristem
Sperm Storage
tissue
Palisade tissue--- Hyphal
tissue
Rhizoid

Slide 31

Longitudinal section of subterranean

Longitudinal section of subterranean
gametophyte of Lycopodium
gametophyte of Lycopodium clavatum
complanatum
 Vegetative Reproduction of the
Prothallus
• By formation of gemmae
– Borne on branches of prothallus
– Singly or in clusters
– Thick walled and stores food materials
– Capable of perennation
• By progressive death and decay of older parts
• Adventitious buds arise from injured parts of
the lobes
 PROTHALLI
• Prothalli are monoecious- male
antheridia and female archegonia
are borne on the same thallus
• ANTHERIDIUM
– Usually sunken in the prothallus;
position visible by a small bulge
– Has a single layered jacket; an
operculum cell
– As water enters, jacket rupture and
biflagellate antherozoids are
liberated
 PROTHALLI
• ARCHEGONIA
– Flask shaped, having basal venter and a neck
– Has 4-8 neck canal cells
– Venter has a venter canal cell and an egg cell
 FERTILIZATION
• When archegonium matures, tip of neck cells
split, neck canal cells and venter canal cell
disintegrate and form a mucilaginous
substance
• Antherozoids enter the archegonium and one
fuses with the egg to form the oospore
• The diploid oospore germinates and produces
the sporophytic generation