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 (1964). G.M. Smith placed this
group between Algae and Pteridophyta.[6]

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)[7]

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).[8]

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.[8]

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.[9] 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,[10]

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.[11] 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.[12]

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.[13]

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.[14]

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.[14]

Monoicous plants are necessarily

hermaphroditic, meaning that the same
plant has both sexes.[14] 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).[14][15] Dioicous
plants are unisexual, meaning that the
same plant has only one sex.[14] All four
patterns (autoicous, paroicous, synoicous
and dioicous) occur in species of the
moss genus Bryum.[15]

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,[16]
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.[17][18][19]

The three bryophyte clades (which may be

treated as divisions) are the
Marchantiophyta (liverworts), Bryophyta
(mosses) and Anthocerotophyta
(hornworts).[20] 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.[19][21] 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.[22] The
issue remains unresolved.
Paraphyletic view …

  liverworts
 
  mosses
  bryop
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.[23][24] 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.[25] The contrast is shown in a
slightly different cladogram:[26]

land plants   liverworts
   
    mosses
  
    hornworts
  
"prot
  such
  Horn
  polysporangiophytes Agla
   
  trach
  vasc
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:[22]
   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.[22] Other studies
suggest a monophyletic group comprising
liverworts and mosses, with hornworts
being sister to vascular plants.[27]

Evolution
There have probably been several different
terrestrialization events, in which originally
aquatic organisms colonized the land, just
within the lineage of the Viridiplantae.[28]
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.[29][1][30][31] 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.[32] Like
green algae and land plants, bryophytes
also produce starch stored in the plastids
and contain cellulose in their walls.[32]
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.[33] The bryophytes and
vascular plants (embryophytes) also have
embryonic development which is not seen
in green algae.[32] 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.[32]
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.[33]

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
Structure Large, with chlorophyll Large, with 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 spores Elaters Peristome teeth Pseudo-elaters

Columella Absent Present Present

Dehiscence Longitudinal or irregular Transverse Longitudinal

Uses
This article is in list format, but may read better
as prose. Learn more

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.[34]
Bryophytes are used in pollution studies to
indicate soil pollution (such as the
presence of heavy metals), air pollution,
and UV-B radiation.[34] Gardens in Japan
are designed with moss to create peaceful
sanctuaries.[34] Some bryophytes have
been found to produce natural pesticides.
The liverwort, Plagiochila, produces a
chemical that is poisonous to mice.[34]
Other bryophytes produce chemicals that
are antifeedants which protect them from
being eaten by slugs.[34] When Phythium
sphagnum is sprinkled on the soil of
germinating seeds, it inhibits growth of
"damping off fungus" which would
otherwise kill young seedlings.[35]

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.[34] Also, because of its
antibiotic properties, Sphagnum was used
as a surgical dressing in World War I.[34]

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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Look up bryophyte in Wiktionary, the

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Wikisource has the text of the 1911
Encyclopædia Britannica article
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 .
Retrieved from
"https://en.wikipedia.org/w/index.php?
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