Contents 1 Habitat 2 Life cycle 2.1 Sexuality 3 Classification and phylogeny 3.1 Paraphyletic view 3.2 Other views 4 Evolution 4.1 Similarities to land plants 4.2 Similarities to aquatic plants 5 Comparative morphology 6 Uses 7 See also 8 References 9 Bibliography 10 External links

Habitat[edit] 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).[5] 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.[5]

Life cycle[edit] See also: Alternation of generations 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. 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, meaning that the more prominent, longer-lived plant is the haploid gametophyte.[6] The diploid sporophytes appear only occasionally and remain attached to and nutritionally dependent on the gametophyte. In bryophytes, the sporophytes are always unbranched and produce a single sporangium (spore producing capsule). 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.[7] 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. Further information: Liverwort § Life cycle, Moss § Life cycle, and Hornwort § Life cycle Sexuality[edit] 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.[8] 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 is said to have the advantage of emphasizing the difference between the gametophyte sexuality of bryophytes and the sporophyte sexuality of seed plants.[8] Monoicous plants are necessarily bisexual (or hermaphroditic), meaning that the same plant has both sexes.[8] 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).[8][9] Dioicous plants are unisexual, meaning that the same plant has only one sex.[8] All four patterns (autoicous, paroicous, synoicous and dioicous) occur in species of the moss genus Bryum.[9]

Classification and phylogeny[edit] 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,[10] 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.[11][12][13] The three bryophyte clades are the Divisions Marchantiophyta (liverworts), Bryophyta (mosses) and Anthocerotophyta (hornworts).[14] 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.[13][15] 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.[16] The issue remains unresolved. Paraphyletic view[edit] embryophytes  liverworts   mosses   hornworts   vascular plants bryophytes 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.[17][18] 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.[19] The contrast is shown in a slightly different cladogram:[20] land plants liverworts mosses hornworts polysporangiophytes "protracheophytes", such as Horneophyton or Aglaophyton tracheophytes or vascular plants 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[edit] An alternative phylogeny, based on amino acids rather than genes, shows bryophytes as a monophyletic group:[21] embryophytes bryophytes hornworts liverworts 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.[16] Other studies suggest a monophyletic group comprising liverworts and mosses, with hornworts being sister to vascular plants.[22]

Evolution[edit] Between 510 - 630 million years ago, 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.[23][1][24][25] 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 land plants[edit] 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 provides protection. The development of gametangia provided further protection specifically for gametes.[26] They also have embryonic development which is a significant adaptation seen in land plants and not green algae.[27] While bryophytes have no truly vascularized tissue, they do have organs that have specific functions, similar to those functions of leaves and stems in higher level land plants.[27] Similarities to aquatic plants[edit] Bryophytes also exhibit connections to their aquatic ancestry. They share various features with their green algae ancestors. Both green algae and bryophytes have chlorophyll a and b, and the chloroplast structures are similar.[27] Like algae and land plants, bryophytes also produce starch and contain cellulose in their walls.[27] Bryophytes depend on water for reproduction and survival. A thin layer of water is required on the surface of the plant to enable the movement of sperm between gametophytes and the fertilization of an egg.[26]

Comparative morphology[edit] 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 and archegonia) Superficial Superficial Immersed Summary of the morphological characteristics of the sporophytes of the three groups of bryophytes: Liverworts Mosses Hornworts Stomata Absent Present Present Structure Small, without chlorophyll Large, with chlorophyll Large, with chlorophyll Persistence Ephemeral Persistent Persistent Growth Defined Defined Continuous Seta Present Present Absent Capsule form Simple Differentiated (operculum, peristome) Elongated Maturation of spores Simultaneous Simultaneous Graduate Dispersion of spores Elaters Peristome teeth Pseudo-elaters Columella Absent Present Present Dehiscence Longitudinal or irregular Transverse Longitudinal

Uses[edit] This article is in a list format that may be better presented using prose. You can help by converting this article to prose, if appropriate. Editing help is available. (April 2017) 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.[28] Bryophytes are used in pollution studies to indicate soil pollution (such as the presence of heavy metals), air pollution, and UV-B radiation.[28] Gardens in Japan are designed with moss to create peacful sanctuaries.[28] Some bryophytes have been found to produce natural pesticides. The liverwort, Plagiochila, produces a chemical that is poisonous to mice.[28] Other bryophytes produce chemicals that are antifeedants which protect them from being eaten by slugs.[28] When Phythium sphagnum is sprinkled on the soil of germinating seeds, it inhibits growth of "damping off fungus" which would otherwise kill young seedlings.[29] Moss peat is made from Sphagnum Commercial Fuel Packaging Wound Dressing Peat is a fuel that is 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.[28] Also, because of the antibiotic properties, sphagnum was used as a surgical dressing in World War I.[28]

See also[edit] Anthocerotophyta (hornworts) Bryophyta (mosses) Embryophyte Marchantiophyta (liverworts) Plant sexuality List of British county and local bryophyte floras

References[edit] ^ a b "Reviews glossary". Retrieved 2009-03-26.  ^ Levetin, Estelle; McMahon, Karen (2012). Plants and Society. New York, NY: McGraw-Hill. p. 139. ISBN 978-0-07-352422-1.  ^ "Bryophytes (Mosses and liverworts) — The Plant List". Retrieved 2017-04-11.  ^ "What are Bryophytes". Retrieved 2017-04-11.  ^ a b "Habitats - ecology - bryophyte". Retrieved 2017-04-12.  ^ "Bryophytes - introduction". Retrieved 2009-05-31.  ^ Cronberg N, Natcheva R, Hedlund K (2006). "Microarthropods Mediate Sperm Transfer in Mosses". Science. 313 (5791): 1255. doi:10.1126/science.1128707. PMID 16946062.  ^ a b c d e Glime, J.M. & Bisang, I. (2014). "Sexuality: Its Determination (Ch. 3-1)" (PDF). In Glime, J.M. Bryophyte Ecology. Volume 1 Physiological Ecology. Michigan Technological University and the International Association of Bryologists. Retrieved 2014-11-09.  ^ a b Watson, E.V. (1981). British Mosses and Liverworts (3rd ed.). Cambridge University Press. p. 7.  (Watson uses the "oecy" terms rather than the "oicy" terms.) ^ Goremykin, V. V. & Hellwig, F. H. (2005). "Evidence for the most basal split in land plants dividing bryophyte and tracheophyte lineages". Plant Systematics and Evolution. 254 (1–2): 93–103. doi:10.1007/s00606-005-0337-1.  ^ Konrat, M.; Shaw, A. J.; Renzaglia, K. S. (2010). "A special issue of Phytotaxa dedicated to Bryophytes: The closest living relatives of early land plants". Phytotaxa. 9: 5–10.  ^ Troitsky AV, Ignatov MS, Bobrova VK, Milyutina IA (December 2007). "Contribution of genosystematics to current concepts of phylogeny and classification of bryophytes". Biochemistry Mosc. 72 (12): 1368–76. doi:10.1134/S0006297907120115. PMID 18205621.  ^ a b Knoop, Volker (2010). "Looking for sense in the nonsense: a short review of non-coding organellar DNA elucidating the phylogeny of bryophytes". Tropical Bryology. 31: 51–60.  ^ "GLOSSARY B". Retrieved 2009-03-26.  ^ Qiu, Y. L.; Li, L.; Wang, B.; et al. (October 2006). "The deepest divergences in land plants inferred from phylogenomic evidence". Proceedings of the National Academy of Sciences. 103 (42): 15511–6. Bibcode:2006PNAS..10315511Q. doi:10.1073/pnas.0603335103. PMC 1622854 . PMID 17030812.  ^ a b Cox, Cymon J.; Li, Blaise; Foster, Peter G.; Embley, T. Martin & Civáň, Peter (2014). "Conflicting Phylogenies for Early Land Plants are Caused by Composition Biases among Synonymous Substitutions". Systematic Biology. 63 (2): 272–279. doi:10.1093/sysbio/syt109. PMC 3926305 . PMID 24399481.  ^ Kenrick, Paul & Crane, Peter R. (1997a). "The Origin and Early Diversification of Land Plants: A Cladistic Study". Washington, D.C.: Smithsonian Institution Press. ISBN 978-1-56098-730-7.  ^ Kenrick, P. & Crane, P.R. (1997b). "The origin and early evolution of plants on land". Nature. 389 (6646): 33–39. Bibcode:1997Natur.389...33K. doi:10.1038/37918.  ^ Bell, N. E. & Hyvönen, J. (2010). "Phylogeny of the moss class Polytrichopsida (BRYOPHYTA): Generic-level structure and incongruent gene trees". Molecular Phylogenetics and Evolution. 55 (2): 381–398. doi:10.1016/j.ympev.2010.02.004. PMID 20152915.  ^ Crane, P. R.; Herendeen, P. & Friis, E. M. (2004). "Fossils and plant phylogeny". American Journal of Botany. 91 (10): 1683–99. doi:10.3732/ajb.91.10.1683. PMID 21652317.  ^ Cox et al. 2014, p. 274. ^ Karol, Kenneth G.; Arumuganathan, Kathiravetpillai; Boore, Jeffrey L.; Duffy, Aaron M.; Everett, Karin DE; Hall, John D.; Hansen, S.K.; Kuehl, Jennifer V.; Mandoli, Dina F.; Mishler, Brent D.; Olmstead, Richard G.; Renzaglia, Karen S. & Wolf, Paul G. (2010). "Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages". BMC Evolutionary Biology. 10 (1): 321. doi:10.1186/1471-2148-10-321. PMID 20969798.  ^ Konrat, M.; Shaw, A.J.; Renzaglia, K.S. (2010). "A special issue of Phytotaxa dedicated to Bryophytes: The closest living relatives of early land plants". Phytotaxa. 9: 5–10.  ^ Karol, Kenneth G.; Arumuganathan, Kathiravetpillai; Boore, Jeffrey L.; Duffy, Aaron M.; Everett, Karin DE; Hall, John D.; Hansen, S.K.; Kuehl, Jennifer V.; Mandoli, Dina F.; Mishler, Brent D.; Olmstead, Richard G.; Renzaglia, Karen S. & Wolf, Paul G. (2010). "Complete plastome sequences of Equisetum arvense and Isoetes flaccida: implications for phylogeny and plastid genome evolution of early land plant lineages". BMC Evolutionary Biology. 10 (1): 321. doi:10.1186/1471-2148-10-321. PMID 20969798.  ^ Shaw, A. Jonathan; Szövényi, Péter; Shaw, Blanka (2011-03-01). "Bryophyte diversity and evolution: Windows into the early evolution of land plants". American Journal of Botany. 98 (3): 352–369. doi:10.3732/ajb.1000316. ISSN 0002-9122. PMID 21613131.  ^ a b Purcell, Adam. "Bryophytes". Basic Biology.  ^ a b c d Everet, Ray; Eichhorn, Susan (2013). 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Bibliography[edit] Lesica, P.; McCune, B.; Cooper, S. V.; Hong, W. S. (1991). "Differences in lichen and bryophyte communities between old-growth and managed second-growth forests in the Swan Valley, Montana". Canadian Journal of Botany. 69: 1745–1755. doi:10.1139/b91-222. 

External links[edit] Wikimedia Commons has media related to Bryophytes. Look up bryophyte in Wiktionary, the free dictionary. 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.[1] 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. v t e Botany History of botany Subdisciplines Plant systematics Ethnobotany Paleobotany Plant anatomy Plant ecology Phytogeography Geobotany Flora Phytochemistry Plant pathology Bryology Phycology Floristics Dendrology Plant groups Algae Archaeplastida Bryophyte Non-vascular plants Vascular plants Spermatophytes Pteridophyte Gymnosperm Angiosperm Plant morphology (glossary) Plant cells Cell wall Phragmoplast Plastid Plasmodesma Vacuole Tissues Meristem Vascular tissue Vascular bundle Ground tissue Mesophyll Cork Wood Storage organs Vegetative Root Rhizoid Bulb Rhizome Shoot Stem Leaf Petiole Cataphyll Bud Sessility Reproductive (Flower) Flower development Inflorescence Umbel Raceme Bract Pedicellate Flower Whorl Floral symmetry Floral diagram Floral formula Receptacle Hypanthium (Floral cup) Perianth Tepal Petal Sepal Sporophyll Gynoecium Ovary Ovule Stigma Archegonium Androecium Stamen Staminode Pollen Tapetum Gynandrium Gametophyte Sporophyte Plant embryo Fruit Fruit anatomy Berry Capsule Seed Seed dispersal Endosperm Surface structures Epicuticular wax Plant cuticle Epidermis Stoma Nectary Trichome Prickle Plant physiology Materials Nutrition Photosynthesis Chlorophyll Plant hormone Transpiration Turgor pressure Bulk flow Aleurone Phytomelanin Sugar Sap Starch Cellulose Plant growth and habit Secondary growth Woody plants Herbaceous plants Habit Vines Lianas Shrubs Subshrubs Trees Succulent plants Reproduction Evolution Ecology Alternation of generations Sporangium Spore Microsporangia Microspore Megasporangium Megaspore Pollination Pollinators Pollen tube Double fertilization Germination Evolutionary development Evolutionary history timeline Hardiness zone Plant taxonomy History of plant systematics Herbarium Biological classification Botanical nomenclature Botanical name Correct name Author citation International Code of Nomenclature for algae, fungi, and plants (ICN) - for Cultivated Plants (ICNCP) Taxonomic rank International Association for Plant Taxonomy (IAPT) Plant taxonomy systems Cultivated plant taxonomy Citrus taxonomy cultigen cultivar Group grex Practice Agronomy Floriculture Forestry Horticulture Lists Related topics Botanical terms Botanists by author abbreviation Botanical expedition Category Portal WikiProject v t e Classification of Archaeplastida / Plantae sensu lato Domain Archaea Bacteria Eukaryota (Supergroup Plant Hacrobia Heterokont Alveolata Rhizaria Excavata Amoebozoa Opisthokonta Animal Fungi) Rhodophyta (red algae) Cyanidiophyceae Porphyridiophyceae Compsopogonophyceae Stylonematophyceae Rhodellophyceae Bangiophyceae Florideophyceae Glaucocystophyta (glaucophytes) Glaucocystophyceae Glaucocystis Cyanophora Gloeochaete Viridiplantae (green algae & land plants) Chlorophyta Palmophyllales Nephroselmidophyceae Prasinophyceae Pseudoscourfieldiales Pyramimonadophyceae Scourfieldiales Pedinophyceae Chlorodendrophyceae UTC clade Ulvophyceae Trebouxiophyceae Chlorophyceae Streptophyta (charophytes, & land plants) Mesostigmatophyceae Chlorokybophyceae Klebsormidiophyceae Phragmo- plastophyta Charophyceae Coleochaetophyceae Zygnematophyceae Embryophyta (land plants) Bryophytes (non-vascular) Marchantiophyta Anthocerotophyta Bryophyta "Moss" †Horneophytopsida Tracheophyta (vascular) Lycopodiophyta (microphylls) †Zosterophyllopsida †Sawdoniales Isoetopsida Lycopodiopsida Euphyllophyta (megaphylls) Moniliformopses (ferns) †Cladoxylopsida †Stauropteridales †Zygopteridales Equisetopsida Psilotopsida Marattiopsida Filicopsida Spermatophyta (seed plants) †Seed ferns Gymnosperms Gnetopsida Pinopsida Cycadopsida Ginkgoopsida Angiosperms or flowering plants Amborellales Nymphaeales Austrobaileyales Magnoliids Monocots Eudicots Other †Trimerophytopsida †Progymnosperm Other †Rhyniopsida † = extinct. See also the list of plant orders. Authority control LCCN: sh85017380 GND: 4040211-3 BNF: cb11964814z (data) NDL: 00566261 Retrieved from "" Categories: BryophytesCryptogamsPlant taxonomyHidden categories: Articles needing cleanup from April 2017All pages needing cleanupArticles with sections that need to be turned into prose from April 2017Wikipedia articles with LCCN identifiersWikipedia articles with GND identifiersWikipedia articles with BNF identifiers

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