Bryophyte

Bryophytes are an informal group

consisting of three divisions of non-
vascular land plants (embryophytes): the
liverworts, hornworts and mosses.[1]
They are characteristically limited in size
and prefer moist habitats although they
can survive in drier environments.[2] The
bryophytes consist of about 20,000 plant
species.[3][4] Bryophytes produce
enclosed reproductive structures
(gametangia and sporangia), but they do
not produce flowers or seeds. They
reproduce via spores.[5] Bryophytes are
usually considered to be a paraphyletic
group and not a monophyletic group,
although some studies have produced
contrary results. Regardless of their
status, the name is convenient and
remains in use as an informal collective
term. The term "bryophyte" comes from
Greek βρύον, bryon "tree-moss, oyster-
green" and φυτόν, phyton "plant".

Marchantia, an example of a liverwort.

Terminology
The term "Bryophyta" was first suggested
by Braun in 1864.[6] G.M. Smith placed
this group between Algae and
Pteridophyta.[7]

Features
The defining features of bryophytes are:

Their life cycles are dominated by the

gametophyte stage
Their sporophytes are unbranched
They do not have a true vascular tissue
containing lignin (although some have
specialized tissues for the transport of
water)[8]
Habitat
Bryophytes exist in a wide variety of
habitats. They can be found growing in a
range of temperatures (cold arctics and
in hot deserts), elevations (sea-level to
alpine), and moisture (dry deserts to wet
rainforests).[9]

Bryophytes can grow where vascularized

plants cannot because they do not
depend on roots for an uptake of
nutrients from soil. Bryophytes can
survive on rocks and bare soil.[9]

Life cycle
The life cycle of a dioicous bryophyte. The
gametophyte (haploid) structures are shown in
green, the sporophyte (diploid) in brown.

Like all land plants (embryophytes),

bryophytes have life cycles with
alternation of generations.[10] In each
cycle, a haploid gametophyte, each of
whose cells contains a fixed number of
unpaired chromosomes, alternates with a
diploid sporophyte, whose cell contain
two sets of paired chromosomes.
Gametophytes produce haploid sperm
and eggs which fuse to form diploid
zygotes that grow into sporophytes.
Sporophytes produce haploid spores by
meiosis, that grow into gametophytes.

Bryophytes are gametophyte

dominant,[11] meaning that the more
prominent, longer-lived plant is the
haploid gametophyte. The diploid
sporophytes appear only occasionally
and remain attached to and nutritionally
dependent on the gametophyte.[12] In
bryophytes, the sporophytes are always
unbranched and produce a single
sporangium (spore producing capsule),
but each gametophyte can give rise to
several sporophytes at once.

The sporophyte develops differently in

the three groups. Both mosses and
hornworts have a meristem zone where
cell division occur. In hornworts, the
meristem starts at the base where the
foot ends, and the division of cells is
pushing the sporophyte body upwards. In
mosses, the meristem is located
between the capsule and the top of the
stalk (seta), and produce cells downward,
elongating the stalk and elevates the
capsule. In liverworts the meristem is
absent and the elongation of the
sporophyte is caused almost exclusively
by cell expansion.[13]

Liverworts, mosses and hornworts spend

most of their lives as gametophytes.
Gametangia (gamete-producing organs),
archegonia and antheridia, are produced
on the gametophytes, sometimes at the
tips of shoots, in the axils of leaves or
hidden under thalli. Some bryophytes,
such as the liverwort Marchantia, create
elaborate structures to bear the
gametangia that are called
gametangiophores. Sperm are flagellated
and must swim from the antheridia that
produce them to archegonia which may
be on a different plant. Arthropods can
assist in transfer of sperm.[14]

Fertilized eggs become zygotes, which

develop into sporophyte embryos inside
the archegonia. Mature sporophytes
remain attached to the gametophyte.
They consist of a stalk called a seta and
a single sporangium or capsule. Inside
the sporangium, haploid spores are
produced by meiosis. These are
dispersed, most commonly by wind, and
if they land in a suitable environment can
develop into a new gametophyte. Thus
bryophytes disperse by a combination of
swimming sperm and spores, in a
manner similar to lycophytes, ferns and
other cryptogams.

Sexuality …

The arrangement of antheridia and

archegonia on an individual bryophyte
plant is usually constant within a species,
although in some species it may depend
on environmental conditions. The main
division is between species in which the
antheridia and archegonia occur on the
same plant and those in which they
occur on different plants. The term
monoicous may be used where
antheridia and archegonia occur on the
same gametophyte and the term
dioicous where they occur on different
gametophytes.[15]

In seed plants, "monoecious" is used

where flowers with anthers
(microsporangia) and flowers with ovules
(megasporangia) occur on the same
sporophyte and "dioecious" where they
occur on different sporophytes. These
terms occasionally may be used instead
of "monoicous" and "dioicous" to
describe bryophyte gametophytes.
"Monoecious" and "monoicous" are both
derived from the Greek for "one house",
"dioecious" and "dioicous" from the Greek
for two houses. The use of the "oicy"
terminology refers to the gametophyte
sexuality of bryophytes as distinct from
the sporophyte sexuality of seed
plants.[15]

Monoicous plants are necessarily

hermaphroditic, meaning that the same
plant has both sexes.[15] The exact
arrangement of the antheridia and
archegonia in monoicous plants varies.
They may be borne on different shoots
(autoicous or autoecious), on the same
shoot but not together in a common
structure (paroicous or paroecious), or
together in a common "inflorescence"
(synoicous or synoecious).[15][16]
Dioicous plants are unisexual, meaning
that the same plant has only one sex.[15]
All four patterns (autoicous, paroicous,
synoicous and dioicous) occur in species
of the moss genus Bryum.[16]

Classification and phylogeny

Hornworts include those bryophytes that are

believed to be the closest living relatives of the
vascular plants.
Mosses are one group of bryophytes.

Traditionally, all living land plants without

vascular tissues were classified in a
single taxonomic group, often a division
(or phylum). More recently, phylogenetic
research has questioned whether the
bryophytes form a monophyletic group
and thus whether they should form a
single taxon. Although a 2005 study
supported the traditional view that the
bryophytes form a monophyletic
group,[17] by 2010 a broad consensus
had emerged among systematists that
bryophytes as a whole are not a natural
group (i.e., are paraphyletic), although
each of the three extant (living) groups is
monophyletic.[18][19][20]

The three bryophyte clades (which may

be treated as divisions) are the
Marchantiophyta (liverworts), Bryophyta
(mosses) and Anthocerotophyta
(hornworts).[21] The vascular plants or
tracheophytes form a fourth, unranked
clade of land plants called the
"Polysporangiophyta". In this analysis,
hornworts are sister to vascular plants
and liverworts are sister to all other land
plants, including the hornworts and
mosses.[20][22] Phylogenetic studies
continue to produce conflicting results.
In particular those based on gene
sequences suggest the bryophytes are
paraphyletic, whereas those based on the
amino acid translations of the same
genes suggest they are monophyletic. A
2014 study concluded that composition
biases were responsible for these
differences and that the bryophytes are
monophyletic.[23] The issue remains
unresolved.

Paraphyletic view …
   liverworts
 
  mosses
  bryo
embryophytes
      hornworts
     
    vascular plants
 

Liverworts are included in the bryophyte group

When extinct plants are taken into

account, the picture is slightly altered.
Some extinct land plants, such as the
horneophytes, are not bryophytes, but
also are not vascular plants because, like
bryophytes, they do not have true
vascular tissue. A different distinction is
needed. In bryophytes, the sporophyte is
a simple unbranched structure with a
single spore-forming organ
(sporangium). In all other land plants, the
polysporangiophytes, the sporophyte is
branched and carries many
sporangia.[24][25] It has been argued that
this contrast between bryophytes and
other land plants is less misleading than
the traditional one of non-vascular versus
vascular plant, since many mosses have
well-developed water-conducting
vessels.[26] The contrast is shown in a
slightly different cladogram:[27]
land plants   liverworts
   
    mosses
  
  
hornworts
  
  polysporangiophytes   "pro
      suc
Hor
Agl

  trac
  vas

The term "bryophyte" thus refers to a

grade of lineages defined primarily by
what they lack. Compared to other living
land plants, they lack vascular tissue
containing lignin and branched
sporophytes bearing multiple sporangia.
The prominence of the gametophyte in
the life cycle is also a shared feature of
the three bryophyte lineages (extant
vascular plants are all sporophyte
dominant).

Other views …

An alternative phylogeny, based on

amino acids rather than genes, shows
bryophytes as a monophyletic group:[23]
   hornworts
 
bryophytes   liverworts
     
embryophytes     mosses
   
  vascular
  plants

If this phylogeny proves correct, then the

complex sporophyte of living vascular
plants might have evolved independently
of the simpler unbranched sporophyte
present in bryophytes.[23] Other studies
suggest a monophyletic group
comprising liverworts and mosses, with
hornworts being sister to vascular
plants.[28]
Evolution
There have probably been several
different terrestrialization events, in
which originally aquatic organisms
colonized the land, just within the lineage
of the Viridiplantae.[29] Between 510 -
630 million years ago, however, land
plants evolved from aquatic plants,
specifically green algae. Molecular
phylogenetic studies conclude that
bryophytes are the earliest diverging
lineages of the extant land
plants.[30][1][31][32] They provide insights
into the migration of plants from aquatic
environments to land. A number of
physical features link bryophytes to both
land plants and aquatic plants.

Similarities to algae and vascular

plants
…

Green algae, bryophytes and vascular

plants all have chlorophyll a and b, and
the chloroplast structures are similar.[33]
Like green algae and land plants,
bryophytes also produce starch stored in
the plastids and contain cellulose in their
walls.[33] Distinct adaptations observed in
bryophytes have allowed plants to
colonize Earth's terrestrial environments.
To prevent desiccation of plant tissues in
a terrestrial environment, a waxy cuticle
covering the soft tissue of the plant may
be present, providing protection. In
hornworts and mosses, stomata provide
gas exchange between the atmosphere
and an internal intercellular space
system. The development of gametangia
provided further protection specifically
for gametes, the zygote and the
developing sporophyte.[34] The
bryophytes and vascular plants
(embryophytes) also have embryonic
development which is not seen in green
algae.[33] While bryophytes have no truly
vascularized tissue, they do have organs
that are specialized for specific
functions, analogous for example to the
functions of leaves and stems in
vascular land plants.[33]

Bryophytes depend on water for

reproduction and survival. In common
with ferns and lycophytes, a thin layer of
water is required on the surface of the
plant to enable the movement of the
flagellated sperm between
gametophytes and the fertilization of an
egg.[34]

Comparative morphology
Summary of the morphological
characteristics of the gametophytes of
the three groups of bryophytes:
 Liverworts Mosses Hornworts

Structure Thalloid or foliose Foliose Thalloid

Symmetry Dorsiventral or radial Radial Dorsiventral

Rhizoids Unicellular Pluricellular Unicellular

Chloroplasts/cell Many Many One

Protonemata Reduced Present Absent

Gametangia
(antheridia Superficial Superficial Immersed
and archegonia)

Summary of the morphological

characteristics of the sporophytes of the
three groups of bryophytes:

Liverworts Mosses Hornworts

Stomata Absent Present Present

Small, without Large, with Large, with

Structure
chlorophyll chlorophyll chlorophyll

Persistence Ephemeral Persistent Persistent

Growth Defined Defined Continuous

Seta Present Present Absent

Differentiated
Capsule form Simple (operculum, Elongated
peristome)

Maturation of
Simultaneous Simultaneous Graduate
spores

Dispersion of
Elaters Peristome teeth Pseudo-elaters
spores

Columella Absent Present Present

Dehiscence Longitudinal or irregular Transverse Longitudinal

Uses
Environmental

Soil Conditioning
Bioindicators
Moss gardens
Pesticides

Characteristics of bryophytes make them

useful to the environment. Depending on
the specific plant texture, bryophytes
have been shown to help improve the
water retention and air space within
soil.[35] Bryophytes are used in pollution
studies to indicate soil pollution (such as
the presence of heavy metals), air
pollution, and UV-B radiation.[35] Gardens
in Japan are designed with moss to
create peaceful sanctuaries.[35] Some
bryophytes have been found to produce
natural pesticides. The liverwort,
Plagiochila, produces a chemical that is
poisonous to mice.[35] Other bryophytes
produce chemicals that are antifeedants
which protect them from being eaten by
slugs.[35] When Phythium sphagnum is
sprinkled on the soil of germinating
seeds, it inhibits growth of "damping off
fungus" which would otherwise kill young
seedlings.[36]
Moss peat is made from Sphagnum

Commercial

Fuel
Packaging
Wound Dressing

Peat is a fuel produced from dried

bryophytes, typically Sphagnum.
Bryophytes' antibiotic properties and
ability to retain water make them a useful
packaging material for vegetables,
flowers, and bulbs.[35] Also, because of
its antibiotic properties, Sphagnum was
used as a surgical dressing in World War
I.[35]
See also
Anthocerotophyta (hornworts)
Bryophyta (mosses)
Embryophyte
Marchantiophyta (liverworts)
Plant sexuality
List of British county and local
bryophyte floras

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External links

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Bryophyta.
Andrew's Moss Site Photos of
bryophytes
27-May-2013 Centuries-old frozen
plants revived , 400-year-old bryophyte
specimens left behind by retreating
glaciers in Canada are brought back to
life in the laboratory.
Farge, Catherine La; Williams, Krista H.;
England, John H. (22 May 2013).
"Regeneration of Little Ice Age
bryophytes emerging from a polar
glacier with implications of totipotency
in extreme environments" .
Proceedings of the National Academy
of Sciences. 110 (24): 9839–9844.
Bibcode:2013PNAS..110.9839L .
 doi:10.1073/pnas.1304199110 .
PMC 3683725 . PMID 23716658 .
Magill, R. E., ed. (1990). Glossarium
polyglottum bryologiae. A multilingual
glossary for bryology. Monographs in
Systematic Botany from the Missouri
Botanical Garden, v. 33, 297 pp. Online
version: Internet Archive .

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