Bryophytes

Bryophytes

Plant scientists recognize two kinds of land plants: bryophytes (nonvascular land plants) and tracheophytes (vascular land plants). Bryophytes are small, herbaceous plants that grow closely packed together in mats or cushions on rocks or soil or as epiphytes on the trunks and leaves of forest trees. Bryophytes are distinguished from tracheophytes by two important characteristics. 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. Second, bryophytes never form xylem tissue, the special lignin-containing, water-conducting tissue that is found in the sporophytes of all vascular plants. At one time, all bryophytes were placed in a single phylum, intermediate in position between algae and vascular plants. Modern studies of cell ultra-structure and molecular biology, however, confirm that bryophytes comprise three separate evolutionary lineages , today recognized as mosses (phylum Bryophyta), liverworts (phylum Marchantiophyta), and hornworts (phylum Anthocerotophyta). Following a detailed analysis of land plant relationships, Paul Kenrick and Peter R. Crane proposed that the three groups of bryophytes represent a structural level in plant evolution, identified by their monosporangiate life cycle. Within the bryophytes, liverworts are the geologically oldest group, sharing a fossil record with the oldest vascular plants (Rhyniophytes) in the Devonian era.

Mosses

Of the three phyla of bryophytes, greatest species diversity is found in the mosses, with up to fifteen thousand species recognized. A moss begins its life cycle when haploid spores, which are produced in the sporophyte capsule, land on a moist substrate and begin to germinate. From the one-celled spore a highly branched system of filaments , called the protonema, develops. Cell specialization occurs within the protonema to form a horizontal system of reddish-brown anchoring filaments and upright green 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 specialized green filaments form leafy buds that will ultimately form the adult gametophyte shoots. Numerous shoots typically develop from each protonema so that, in fact, 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.

As is typical of bryophytes, mosses produce large, multicellular sex organs for reproduction. Many bryophytes are unisexual, or sexually dioicous . In mosses male sex organs, called antheridia, are produced in clusters at the tips of shoots or branches on the male plants; female sex organs, the archegonia, are produced in similar fashion on female plants. Numerous motile sperm are produced by mitosis inside the brightly colored, club-shaped antheridia

DISTINGUISHING CHARACTERISTICS OF MOSSES, LIVERWORTS, AND HORNWORTS
Characteristics	Mosses (Bryophyta)	Liverworts (Marchantiophyta)	Hornworts (Anthocerotophyta)
Protonema	Filamentous, forming many buds	Globose, forming one bud	Globose, forming one bud
Gametophyte form	Leafy shoot simple or with air chambers	Leafy shoot or thallus; thallus	Simple thallus
Leaf arrangement	Leaves in spirals	Leaves in three rows	Not Applicable
Leaf form	Leaves undivided, midvein present	Leaves divided into two-plus lobes, no midvein	Not Applicable
Special organelles	None	Oil bodies	Single plastids with pyrenoids
Water-conducting cells	Present in both gametophyte and sporophyte	Present only in a few simple thalloid forms	Absent
Rhizoids	Brown, multicellular	Hyaline, one-celled	Hyaline, one-celled
Gametangial position	Apical clusters (leafy forms)	Apical clusters (leafy forms) or on upper surface of thallus	Sunken in thallus, scattered
Stomates	Present on sporophyte capsule	Absent in both generations	Present in both sporophyte and gametophyte
Seta	Photosynthetic, emergent from gametophyte early in development	Hyaline, elongating just prior to spore release	Absent
Capsule	Complex with operculum, theca, and neck; of fixed size	Undifferentiated, spherical, or elongate; of fixed size	Undifferentiated, horn-shaped; growing continuously from a basal meristem
Sterile cells in capsule	Columella	Spirally thickened elaters	Columella and pseudoelaters
Capsule dehiscence	At operculum and peristome teeth	Into four valves	Into two valves

while a single egg develops in the base of each vase-shaped archegonium. As the sperm mature, the antheridium swells and bursts open. Drops of rainwater falling into the cluster of open antheridia splash the sperm to nearby females. Beating their two whiplash flagellae , the sperm are able to move short distances in the water film that covers the plants to the open necks of the archegonia. Slimy mucilage secretions in the archegonial neck help pull the sperm downward to the egg. The closely packed arrangement of the individual moss plants greatly facilitates fertilization. Rain forest bryophytes that hang in long festoons from the trees rely on torrential winds with the rain to transport their sperm from tree to tree, while the small pygmy mosses of exposed, ephemeral habitats depend on the drops of morning dew to move their sperm. Regardless of where they grow, all bryophytes require water for sperm dispersal and subsequent fertilization.

Embryonic growth of the sporophyte begins within the archegonium soon after fertilization. At its base, or foot, the growing embryo forms a nutrient transfer zone, or placenta, with the gametophyte. Both organic nutrients and water move from the gametophyte into the sporophyte as it continues to grow. In mosses the sporophyte stalk, or seta, tears the archegonial enclosure early in development, leaving only the foot and the very base of the seta embedded in the gametophyte. The upper part of the archegonium remains over the tip of the sporophyte as a caplike calyptra. Sporophyte growth ends with the formation of a sporangium (the capsule) at the tip of the seta. Within the capsule, water-resistant haploid spores are formed by meiosis. As the mature capsule swells, the calyptra falls away. This allows the capsule to dry and break open at its tip when the spores are mature. Special membranous structures, called peristome teeth, that are folded down into the spore mass now bend outward, flinging the spores into the drying winds. Moss spores can travel great distances on the winds, even moving between continents on the jet streams. Their walls are highly protective, allowing some spores to remain viable for up to forty years. Of course, if the spore lands in a suitable, moist habitat, germination will begin the cycle all over again.

Liverworts and Hornworts

Liverworts and hornworts are like mosses in the fundamental features of their life cycle, but differ greatly in organization of their mature game-tophytes and sporophytes. Liverwort gametophytes can be either leafy shoots or flattened thalli . In the leafy forms the leaves are arranged on the stem in one ventral and two lateral rows or ranks, rather than in spirals like the mosses. The leaves are one cell-layer thick throughout, never have a mid-vein, and are usually divided into two or more parts called lobes. The ventral leaves, which actually lie against the substrate (soil or other support), are usually much smaller than the lateral leaves and are hidden by the stem. Anchoring rhizoids, which arise near the ventral leaves, are colorless and unicellular. The flattened ribbonlike to leaflike thallus of the thallose liver-worts can be either simple or structurally differentiated into a system of dorsal air chambers and ventral storage tissues. In the latter type the dorsal epidermis of the thallus is punctuated with scattered pores that open into the air chambers. Liverworts synthesize a vast array of volatile oils, which they store in unique organelles called oil bodies. These compounds impart an often spicy aroma to the plants and seem to discourage animals from feeding on them. Many of these compounds have potential as antimicrobial or anticancer pharmaceuticals.

Liverworts.

Liverwort sporophytes develop completely enclosed within gametophyte tissues until their capsules are ready to open. The seta, which is initially very short, consists of small, thin-walled hyaline cells. Just prior to capsule opening, the seta cells lengthen, thereby increasing the length of the seta up to twenty times its original dimensions. This rapid elongation pushes the darkly pigmented capsule and upper part of the whitish seta out of the gametophytic tissues. With drying, the capsule opens by splitting into four segments, or valves. The spores are dispersed into the winds by the twisting motions of numerous intermixed sterile cells called elaters. In contrast to mosses, which disperse their spores over several days, liverworts disperse the entire spore mass of a single capsule in just a few minutes.

Hornworts.

Hornworts resemble some liverworts in having simple, unspecialized thalloid gametophytes, but they differ in many other characters. For example, colonies of the symbiotic cyanobacterium Nostoc fill small cavities that are scattered throughout the ventral part of the hornwort thallus. When the thallus is viewed from above, these colonies appear as scattered blue-green dots. The cyanobacterium converts nitrogen gas from the air into ammonium, which the hornwort requires in its metabolism, and the hornwort secretes carbohydrate-containing mucilage, which supports the growth of the cyanobacterium. Hornworts also differ from all other land plants in having only one large, algal-like chloroplast in each thallus cell. Hornworts get their name from their long, horn-shaped sporophytes. As in other bryophytes, the sporophyte is anchored in the gametophyte by a foot through which nutrient transfer from gametophyte to sporophyte occurs. The rest of the sporophyte, however, is actually an elongate sporangium in which meiosis and spore development take place. At the base of the sporangium, just above the foot, is a mitotically active meristem , which adds new cells to the spore-producing zone throughout the life span of the sporophyte. In fact, the sporangium can be releasing spores at its apex at the same time that new spores are being produced by meiosis at its base. Spore release in hornworts takes place gradually over a long period of time, and the spores are mostly dispersed by water movements rather than by wind.

Mosses, liverworts, and hornworts are found throughout the world in a variety of habitats. They flourish particularly well in moist, humid forests like the fog forests of the Pacific Northwest or the montane rain forests of the Southern Hemisphere. Their ecological roles are many. They provide seed beds for the larger plants of the community , they capture and recycle nutrients that are washed with rainwater from the canopy, and they bind the soil to keep it from eroding. In the Northern Hemisphere peatlands, wetlands often dominated by the moss Sphagnum, are particularly important bryophyte communities. This moss has exceptional water-holding capacity, and when dried and compressed forms a coal-like fuel. Throughout northern Europe, Asia, and North America, peat has been harvested for centuries for both fuel consumption and horticultural uses, and today peat lands are managed as a sustainable resource.

see also Evolution of Plants; Gametophyte; Nitrogen Fixation; Peat Bogs; Reproduction, Alternation of Generations and; Sporophyte.

Barbara Crandall-Stotler

Bibliography

Crandall-Stotler, Barbara. "Morphogenetic Designs and a Theory of Bryophyte Origins and Divergence." BioScience 30 (1980): 580-85.

Hébant, Charles. The Conducting Tissues of Bryophytes. Vaduz: J. Cramer, 1977.

Kenrick, Paul, and Peter R. Crane. The Origin and Early Diversification of Land Plants: A Cladistic Study. Washington, DC: Smithsonian Institution Press, 1997.

Miller, Norton G. "Bogs, Bales and BTUs: A Primer on Peat." Horticulture 59 (1981):38-45.

Schofield, W. B. Introduction to Bryology. New York: Macmillan, 1985.

Shaw, Jonathon A., and Bernard Goffinet, eds. The Biology of Bryophytes. Cambridge, England: Cambridge University Press, 2000.