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TEXT Bryophyta is a group of the simplest and primitive plants of class Embryophyta. These plants occupy a position between the green thallophytes (algae) and pteridphytes. Bryophytes are those embryo-bearing land plants that are non-vascular. They have tissue differentiation and enclosed reproductive system but they lack vascular tissues. They have neither flower nor produce seeds and reproduce through spores. They are also called the amphibians of the plant world due to their preference for moist habitat and dependence on water for fertilization, despite being land plants. Bryophytes are generally small, but form a significant component of vegetation in cooler and moist northern and southern latitudes, and extremely humid climates of both temperate and tropical regions. Almost majority of the bryophytes are moisture-loving, mostly terrestrial, growing in shaded places, moist rocks, bases of trees and similar moist places; or a few inhabit aquatic habitats, e.g., Riccia fluitans, Ricciacarpus Fontinalis antypyretica and Riella sp. natans, A number of bryophytes grow in bogs, such as species of Sphagnum, Leucobryam, Sphalozea etc. Others grow in exposed sites, and can endure drought, e.g., Polytrichum juniperinum and Tortula viralis. General Characters of Bryophytes 1. Bryophytes are generally small-statured land-plants, possessing chlorophyll-a and -b, starch, cellulosic walls, and sometimes possess a cuticle. 2. These are mostly autotrophic; a few are saprophytes, e. g., Cryptothalus merabilis is a saprophyte which grows in swamps. It is completely devoid of chlorophyll and lives at the expense of its mycorrhizal fungus. 3. The main plant body of bryophytes is gametophyte. It is more conspicuous, long- lived, independent, green and freely branched. On the other hand, the sporophyte is short-lived and completely dependent on the gametophyte. 4. Gametophytic and sporophytic phases are present in the life cycle of bryophytes. Both these phases are morphologically distinct (i. e., heteromorphic). 5. In the liverworts such as Riccia and Marchantia, the gametophyte is prostrate and thalloid, but in the mosses, the gametophytic plant body is erect and differentiated into stem and lateral appendages (Fig.) 6. The true roots are absent in bryophytes and the function of anchorage and absorption is performed by filamentous structures known as rhizoids. The rhizoids may be unicellular and unbranched (liverworts and hornworts), or multicellular and branched (mosses). Rhizoids are analogous to the roots of higher plants. 7. The plant body is composed of parenchymatous cells and lacks differentiated xylem and phloem characteristic of true vascular plants. 8. The plan body of bryophytes is devoid of filaments as seen in many algae, but rather of true parenchyma derived by three dimensional growth, usually from apical meristem. 9. Being embryophytes, they have multicellular sporangia and gametangia: reproductive cells are always surrounded by one or several layers of sterile cells. 10. Bryophytes, being non-vascular plants, can never grow too large; their small and simple structure provides great selective advantage in certain habitats. The tiny parenchymatous bodies of mosses and liverworts permit them to thrive in microhabitats, such as stone-walls, fences and bear rock, microhabitats that have too little water or soil for larger vascular plants. 11. The bryophytes reproduce by vegetative, asexual and sexual methods. 12. Vegetative multiplication in bryophytes takes place by: a. Fragmentation: Progressive death and decay of older portions of thallus at the posterior end reaching a dichotomy, and younger branches at anterior end by apical growth separating into new individuals. In this way, an increasing number of plants are produced. b. Adventitious branches: In thallose liverworts and hornworts, often there is formation of adventitious branches from the underside midrib which on breaking form new plants. c. Innovations: These are few axillary branches which grow vigorously and after separation from mother plant grow into new individuals, e.g. Sphagnum. d. Gemmae: These are special vegetative organs, and in a more restricted sense refer to a propagative organ of definite form. The gemmae characteristically originate from single cell, but may be unicellular, bicellular or multicellular at the time of its release. 13. Typical spore formation in bryophytes is lacking. But they show the formation of gametophyte directly from the cells of sporophyte, other than a spore, a phenomenon called as apospory. The mosses have great power of regeneration and the wounding of unspecialized cells of various parts of sporophyte induces the production of green filament, the latter bears a new crop of leafy gametophores. 14. The sexual reproduction in bryophytes is of oogamous type. The sex organs are multicellular and considerably more complicated than those of thallophytes. The male sex organs are known as antheridia; the latter are stalked, globose or somewhat elliptic. They have an outer sterile one-cell thick jacket, which surrounds a solid mass of fertile cells, the androcytes. Each androcyte eventually metamorphosis into a motile, biflaggelate antherozoid. 15. The female sex organ known as archegonium is a flask-shaped structure having basal swollen ventor and somewhat more slender and elongated upper part, the neck. The ventor and neck are surrounded by a jacket of sterile cells. Four cover cells are located at the top of the neck. Each archegonium contains a single egg cell which is located in the ventor. A short stalk attaches the archegonium to gametophyte. 16. Water is necessary for fertilization. Shortly before egg cell is mature, the cover cells separate. At the same time, the cells in the centre of neck dissolve so that an open canal connects the ventor with moisture outside the archegonium. Free swimming sperms substances formed move by the towards chemical archegonium. This response to chemical stimuli is known as chemotactic response. Several sperms may enter the archegonium but only one fertilizes the egg to form diploid zygote within ventor. The zygote does not have any resting period and it divides immediately after fertilization. The first division of zygote is always transverse and outer cell gives rise to embryo. Thus, they show exoscopic mode of embryo development. 17. The embryo develops within the ventor of the archegonium and gives rise to sporophyte or sporogonium. The sporophyte is a simple structure without rhizoids, stem or leaves. It is completely dependent on gametophyte. 18. The sporophyte is a projecting structure differentiating into foot, seta and capsule. 19. The sporogenous cells present in capsule form haploid spores (meiospores) after reduction division. All spores are similar in shape and size. The spores are non-motile and disseminate exclusively by wind. Under favorable conditions, spores either form filamentous germ tube which divides to give rise to young gametophyte (Riccia, Marchantia) or form a protonema, as in mosses. The protonema bears many buds which develop into erect gametophores. 20. In bryophytes, growth of gametophore is by means of a single apical cell rather than by a meristematic tissue. Regardless of where bryophyte grows, all bryophytes require water for fertilization and dispersal. Importance of Bryophytes Bryophytes form an important part of vegetation in cold temperate regions of world, including Himalayas. Different species of bryophytes are widely used as medicine, construction material, insect repellant, packing materials and smoke filters. Some are also used in horticulture because of their high water holding capacity. Gardeners use moss- wrapped baskets and pots for cultivating flowers. Mosses are used as soil-additives. Some insects and birds use mosses for building shelter. Many bryophytes are first to colonize open and often nutrient poor sites where no other plant is capable to grow; for instance, they grow on bare rocks and on recently deposited volcanic ash. Thus, they act as pioneers of vegetation. Certain mosses have preference for special kind of substrate for their growth. Therefore, some of them may serve as the indicators of mineral deposits, pH condition of substrate and also of particular seed plant community and pollution levels. Classification of Bryophytes The term bryophytes was first time used by R. Braun (1864), who included algae, fungi, lichens and mosses in this group. In later systems of classification, however, algae, fungi and lichens were placed in a separate division thallophyta, and liverworts and mosses together in bryophytes. Eichler, way back in 1813, had recognized two classes in division bryophytes; i. Hepaticae (liverworts) ii. Musci (mosses) Takhtajan (1953) recognized three classes in bryophytes: i. Hepaticae (liverworts) ii. Anthocerotae (hornworts) iii. Musci (mosses) Rothmaler (1957) changed nomenclature of three classes of bryophytes as under: i. Hepaticae Hepatocopsida ii. Anthocerotae Anthocerotopsida iii. Musci Bryopsida At present, majority of the workers recognize three divisions for those plants previously grouped together as bryophytes under the Kingdom Plantae. These are: Division: Hepatophyta (liverworts) Division: Anthocerotophyta (hornworts) Division: Bryophyta (mosses) Diagnostic features of the Divisions I. Hepatophyta (liverworts) The name Hepatophyta has been derived from Latin word ‘hepato’, which means liver. Hence, members of this class are commonly known as liverworts. The name liverwort is very old having been used in the 9th century. It was probably applied to these plants because of their resemblances to liver and the belief that plant resembling human organs would cure diseases of organs they resemble. The diagnostic features of Hepatophyta include: 1. The gametophyte which forms main plant body is independent, dorsi-ventral and thalloid or foliose. 2. In thalloid forms, plant body is prostrate, lobed and dichotomously branched. 3. The ventral surface of thallus bears many unicellular, unbranched, smooth-walled and tuberculate rhizoids and scales. 4. In foliose forms, central axis bears leaves in two or three rows. Leaves are without midrib. 5. Anatomically, thallus is either simple or complex and is devoid of mechanical tissue. 6. Photosynthetic cells contain numerous chloroplasts without pyrenoids. 7. Vegetative multiplication takes place by death and decay of other parts of thallus and by adventitious branches, gemmae or tubers. 8. The members of Hepatophyta are homothallic (monoecious, e.g. Riccia) or heterothallic (dioecious, e.g. Marchantia). 9. The sex organs occupy dorsal or terminal position on thallus. They develop from single superficial initial. 10. Sporophyte is simple, represented by capsule only (e.g. Riccia), or differentiated into foot, seta and capsule (e.g. Marchantia). 11. The sporogenous cells, called as archesporium, develop from the endothecium. The sporogenous tissue either forms only spore mother cells (e.g. Riccia), or becomes differentiated into fertile spore mother cells and sterile elater mother cells (e.g. Marchantia, Pelia). 12. The capsule does not have sterile columella. The only sterile cells are elaters. The spores give rise to gametophyte on germination. II. Anthocerotophyta (hornworts) It is a small group of bryophytes, widely distributed in both temperate and tropical regions of world. The diagnostic features of anthocerotophyta are as under: 1. Gametophyte is flat, dorso-ventrally lobed and thalloid. 2. The thallus is attached to substratum with the help of smooth-walled rhizoids. Tuberculate rhizoids and scales are absent. 3. Air pores and air chambers are absent but intercellular mucilage cavities are present which open on ventral surface by slit-like pores. 4. Sex organs are embedded in thallus. Antheridia develop on dorsal surface from hypodermal cells. They occur singly or in groups in antheridial chambers. 5. Archegonia develop from dorsal epidermal cells. 6. Sporophyte is differentiated into a long cylindrical capsule, small meristematic seta and a bulbous foot. 7. The capsule wall is 4-6 layers and the epidermis has stomata. 8. Elaters do not have thickening bands and are called pseudo-elaters. III. Bryophyta (mosses) The mosses are the largest group of bryophytes, which occur in almost all parts of the world. They are abundant at altitudes ranging from 1200 to 2500 meters, and some species occur up to 6500 meters. The characteristic features of differentiate vegetative them from features of other the the mosses bryophytes are gametophyte, that the early development and structure of mature sex organs and that of sporophyte. These features are explained as under: 1. The gametophyte has two growth stages. a. Protonema stage: It is the juvenile stage represented by prostrate, creeping, green and branched filamentous structure; it develops from the spore and is only a transitory vegetative stage. b. Leafy stage or gametophores: an erect cylindrical shoot with persistent leaves and sex organs. 2. The main axis of gametophore may be, branched or unbranched. The branches always arise below the leaves. 3. The leaves are simple, minute, sessile and usually with a distinct midrib. They are only one cell in thickness. 4. The rhizoids are multicellular, branched and have oblique septa. 5. The axis is differentiated into a central conducting tissue and a peripheral cortex. 6. Vegetative multiplication takes place by fragmentation, stolons, branching of protonema, special leafy shoots, gemmae and persistent apices. 7. The sex organs are borne in groups at the apices of the branches. The plants are are club-shaped, dioecious or monoecious. 8. The antheridia narrow and elongated. The antheridial jacket is single layered. 9. The male gametes are biflagellate and spirally-coiled. 10. The archegonia are stalked with a much elongated neck and a massive ventor. 11. The sporophyte is usually differentiated into foot, seta and capsule. The elongated seta raises the capsule much above the gametophyte to facilitate the dispersal of spores. 12. The capsule usually has a peristome which helps in dispersal of spores. 13. On germination, the spores first form protonema which in turn develops into leafy gametophore.