One of
the archegonia at the female shoot tip will develop into the
sporophyte after fertilisation. It
grows out from the perianth as a stalk topped by spore capsule,
which is usually darkly coloured
and glossy when mature and may be more-or-less spherical but is
usually elongated. In some
forms, the young sporophyte begins its development completely
enclosed in gametophyte tissue
which grows up from the stem to enclose the sporophyte, forming a
marsupium. This protective
sheath of stem tissue may bear leaves or bracts and may replace a
perianth in some forms which
lack this structure and may occur at the end of the shoot or hang
down from the branch at right
angles and may be a sizeable structure.
Spore
Dispersal
The
wall of the spore capsule is two to several cell layers thick and
the walls of the cells develop
bands of thickening. The spores are produced by meiosis and so are
haploid. As the ripe
capsule dries out, these bands of thickenings, which are deposited
in very specific patterns,
create stresses in the shrinking tissues which causes the capsule
to rupture and to split into four
valves (above right). Bearing the capsule at the end of a stalk
enables the spores to be released
above the boundary layer of still air, assisting dispersal. This
is further assisted by the discharge
mechanism. In most leafy liverworts, the spores are discharged by
a water rupture mechanism.
Interspersed with the spores are swollen, elongated cells called elaters. These clear,
water-filled
elaters have a double spiral band of wall-thickenings on the
inside of the cell wall. The elaters
may be free or anchored to the inside walls of the spore capsule (sporangium) in liverworts, but
are anchored at their base in this mechanism. As they dry out, the
spiral thickenings try to
contract, like stretched springs eventually breaking the water
column inside which cavitates as it
suddenly fills with pockets of gas and the elater instantaneously
untwists, tearing itself free from
the sporangium wall, flying and rotating into the air, spinning
off attached spores.
In the thalloid liverworts, Marchantia and Pellia (see: bryophytes introduction) each
elater has a
single spiral thickening, which is much weaker and wriggle about
as they slowly untwist upon
drying. This hygroscopic mechanism slowly fluffs up the spore mass
which can then be more
easily dispersed by wind. In some leafy liverworts, such as Frullania, each elater spans the
sporangium and is firmly attached to the sporangium floor at one
end and to the roof at the other.
As the four valves of the capsule bend back, as the capsule dries
and opens, the elaters are
momentarily stretched and each has a single spiral of thickening
and is essentially a stretched
spring in the wall of a water-filled tube. In less than a second
the elaters are torn free at their
lower ends, which moves in an arc (as when a stretched spring is
bent backwards and then
released from one end) flinging out the spores!
Spore production is prolific. An estimated 24 000 spores occur in
each capsule of Lophocolea
cuspidata.
Under suitable conditions a spore will germinate into a young
gametophyte. The
pattern of sporeling development depends on
species and, for example, the young
gametophyte may pass through either a filamentous stage, a
disc-like stage or a ball-of-cells
stage. Eventually the three-sided apical cell is formed. This may
generate at first a shoot with
juvenile leaves, which may be single cells or a row of cells, and
then later shoots which appear
more adult-like, before the gametophyte finally puts out mature
shoots.
Some leafy liverworts can also reproduce asexually by specialised
means, apart from incidental
fragmentation or separation of parts due to death of stem
segments, some have deciduous
propagules: branches, leaves or
perianths (Gymnocolea
inflata),
which readily detach and
regenerate into new plants. In Frullania
fragifolia,
the large antical lobes of the bilobed leaves
detach, leaving only the smaller postical lobes. Some have more
specialised asexual propagules
called gemmae, which are structures
specialised only for reproduction, having no other specific
function in the adult plant. In Lophozia
ventricosa,
gemmae are produced at the tips of leaves,
which may bear clusters of gemmae along their margins, each gemma
consists of one or two cells
with dense cytoplasm which detach easily.
Plagiochila
(Featherwort) - an example of a leafy liverwort
Plagiochila is a beautiful leafy liverwort known as
Featherwort. This specimen is from the British isles in a locale
where two similar species occur: Plagiochila porelloides
(Lesser Featherwort) and Plagiocila asplenioides (Greater
Featherwort). These two species can be hard to distinguish. P.
asplenioides is the larger of the two, forming shoots up to
12 cm long and 9 mm wide, with leaves up to 4.5 mm long. The
smaller P. porellioides forms shoots up to 7 cm long, up
to 6 mm wide and with leaves up to 3 mm long. Lesser Featherwort
has frequent thread-like branches that bind adjacent shoots
together to form a mat, whereas Greater Featherwort lacks
threadlike branches.
A woodland population of Plagiochila asplenioides growing
among mosses (in particular the large moss Plagiomnium
undulatum) in winter. This site is also shared with Early
purple Orchids (Orchis mascula), Bugle (Ajuga reptans)
and Bluebells (Hyacinthoides non-scripta). which were just
beginning to put up new leaves at this time.
This specimen was slightly over 7 cm in
length (the photo above shows about 1 cm of shoot length) up to
9 mm wide and with leaves up to 4 mm or more in length and so is
compatible with P. asplenioides. Furthermore, this specimen bore three branches
that where similar to the main shoot. Plagiochila
has greatly reduced amphigastria (underleaves), which are
generally considered absent. this liverwort is succubous, meaning
the back margin (nearest the apex in this case) lies beneath the
front margin of the next leaf nearest the apex. The front margin
folds back and extends down the stem, forming a line where it
joins the stem (insertion line) that curves slightly and ends at
about the midline of the stem.
As well as reproducing sexually, Plagiochila can
propagate asexually by shedding leaves which then regenerate into
new plants.
Above: bunches of rhizoids can be seen on the nodes of the
underside of the stem. According to one source, the leafy portions
of the stems of P. asplenioides lack rhizoids, but I have
not yet verified this.
The leaves of both species usually have minute teeth along their
margins,but these teeth are often too minute to see without a
microscope, as in this specimen.
The perianth of Plagiochila, when it forms, consists of
two modified leaf-like structures or valves and is laterally
compressed.
Plagiochila porelloides is found on woodland banks,
streamsides, and sheltered areas on upland slopes. P.
asplenioides is the more common species in the south and
east of Britain and is found in damp turf, woodlands, chalky
slopes, fens, stream banks, hedgerows, and on rocks and rotting
wood.
Frullania
Recommended
Further Reading
Watson,
E.V. 1964. The Structure and life of Bryophytes. Hutchinson
University Library.
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Diplophyllum%20albicans.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Frullania%20tamarisci.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Frullania%20fragilifolia.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Lophozia%20ventricosa.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Gymnocolea%20inflata.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Lophocolea%20bidentata.pdf
http://rbg-web2.rbge.org.uk/bbs/Activities/liverworts/Lophocolea%20heterophylla.pdf
Article created: 26 Nov 2016
Article updated: 21 Jan 2020