Plant scientists recognize two kinds of land plants, namely, bryophytes, or nonvascular land plants and tracheophytes,or vascular land plants. Bryophytes are small, herbaceous plants that grow closely packed together in mats or cushions on rocks, soil, or as epiphytes on the trunks and leaves of forest trees. Bryophytes are distinguished from tracheophytes by two important characters. 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, bryophytes were placed in a single phylum, intermediate in position between algae and vascular plants. Modern studies of cell ultrastructure and molecular biology, however,confirm that bryophytes comprise three separate evolutionary lineages, which are today recognized as mosses (phylum Bryophyta), liverworts (phylum Marchantiophyta) and hornworts (phylum Anthocerotophyta). Following a detailed analysis of land plant relationships, Kenrick and Crane (1998) proposed that the three groups of bryophytes represent a grade or structural level in plant evolution, identified by their "monosporangiate" life cycle. Within this the geologically oldest group, sharing a fossil record with the oldest vascular plants in the Devonian era.
Of the three phyla of bryophytes, greatest species diversity is found in the mosses, with up to 15,000 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, called caulonemal filaments and upright, green filaments, called chloronemal 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 caulonemal filaments specialize to 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 and 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 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 rain water falling into the cluster of open antheridia splash the sperm to near-by 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. Slimey 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 cap-like calyptra. Sporophyte growth ends with the formation of a sporangium or capsule at the tip of the seta. Within the capsule, water-resistant 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. 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 40 years. Of course, if the spore lands in a suitable, moist habitat, germination will begin the cycle all over again.
Liverworts and hornworts are like mosses in the fundamental features of their life cycle, but differ greatly in organization of their mature gametophytes 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 midvein and are usually divided into two or more parts called lobes. The ventral leaves, which actually lie against the substrate, 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 ribbon-like to leaf-like thallus of the thallose liverworts 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 pharmecuticals.
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 upto 20 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 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 peatlands are managed as a sustainable resource.
- Crandall-Stotler, Barbara. "Morphogenetic designs and a theory of bryophyte origins and divergence." BioScience 30(1980): 580-585.
- 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, D. C.: 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
Distinguishing Characters of Mosses (Phylum Bryophyta), Liverworts (Phylum Marchantiophyta) and Hornworts
Filamentous, forming many buds
Globose, forming one bud
Globose, forming one bud
Leafy shoot or thallus; thallus simple or with air chambers
Leaves in spirals
Leaves in three rows
Leaves undivided, midvein present
Leaves divided into 2+ lobes, no midvein
Single plastids with pyrenoids
Water conducting cells
Present in both gametophytes and sporophytes
Present only in a few simple thalloid forms
Apical clusters (leafy forms) or on upper surface of thallus
Sunken in thallus, scattered
Present on sporophyte capsule
Absent in both generations
Present in both sporophyte and gametophyte
Photosynthetic, emergent from gametophyte early in development
Hyaline, elongating just prior to spore release
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
Spirally thickened elaters
Columella and pseudoelaters
At operculum and peristome teeth
Into 4 valves
Into 2 valves
Prepared by : Barbara Crandall-Stotler, Department of Plant Biology, Southern Illinois University,
Carbondale, IL 62901-6509
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