Bryophytes include the various mosses and liverworts that are found commonly growing in moist shaded areas in the hills.
Bryophytes are also called amphibians of the plant kingdom because these plants can live in soil but are dependent on water for sexual reproduction. They usually occur in damp, humid and shaded localities. They play an important role in plant succession on bare rocks/soil. The plant body of bryophytes is more differentiated than that of algae. It is thallus-like and prostrate or erect, and attached to the substratum by unicellular or multicellular rhizoids. They lack true roots, stem or leaves. They may possess root-like, leaf-like or stem-like structures. The main plant body of the bryophyte is haploid. It produces gametes, hence is called a gametophyte. The sex organs in bryophytes are multicellular. The male sex organ is calledantheridium. They produce biflagellate
antherozoids. The female sex organ called archegoniumis flask-shaped and produces a single egg. The antherozoids are released into water where they come in contact witharchegonium. An antherozoid fuses with the egg to produce the zygote. Zygotes do not undergo reduction division immediately. They produce a multicellular body called a sporophyte. The sporophyte is not free-living but attached to the photosynthetic gametophyte and derives nourishment from it. Some cells of the sporophyte undergo reduction division (meiosis) to produce haploid spores. These spores germinate to produce gametophyte. Bryophytes in general are of little economic importance but some mosses provide food for herbaceous mammals, birds and other animals. Species of Sphagnum, a moss, provide peat that have long been used as fuel, and as packing material for trans-shipment of living material because of their capacity to hold water. Mosses along with lichens are the first organisms to colonise rocks and hence, are of great ecological importance. They decompose rocks making the substrate suitable for the growth of higher plants. Since mosses form dense mats on the soil, they reduce the impact of falling rain and prevent soil erosion. The bryophytes are divided into liverworts and mosses.
The liverworts grow usually in moist, shady habitats such as banks of streams, marshy ground, damp soil, bark of trees and deep in the woods. The plant body of a liverwort is thalloid, e.g., Marchantia. The thallus is dorsiventral and closely appressed to the substrate. The leafy members have tiny leaf-like appendages in two rows on the stem-like structures. Asexual reproduction in liverworts takes place by fragmentation of thalli, or by the formation of specialised structures called gemmae(sing. gemma). Gemmae are green, multicellular, asexual buds, which develop in small receptacles calledgemma cups located on the thalli. The gemmae become detached from the parent body and germinate to form new individuals. During sexual reproduction, male and female sexorgans are produced either on the same or on different thalli. The sporophyte is differentiated into a foot, seta and capsule. After meiosis, spores are produced within the capsule. These spores germinate to form free-living gametophytes.
The predominant stage of the life cycle of a moss is the gametophyte which consists of two stages. The first stage is the protonema stage, which develops directly from a spore. It is a creeping, green, branched and frequently filamentous stage. The second stage is the leafy stage, which develops from the secondary protonema as a lateral bud. They consist of upright, slender axes bearing spirally arranged leaves. They are attached to the soil through multicellular and branched rhizoids. This stage bears the sex organs. Vegetative reproduction in mosses is by fragmentation and budding in the secondary protonema. In sexual reproduction, the sex organs antheridia and archegonia are produced at the apex of the leafy shoots. After fertilisation, the zygote develops into a sporophyte, consisting of a foot, seta and capsule. The sporophyte in mosses is more elaborate than that in liverworts. The capsule contains spores. Spores are formed after meiosis. The mosses have an elaborate mechanism of spore dispersal. Common examples of mosses are Funaria, Polytrichumand Sphagnum
The Pteridophytes include horsetails and ferns. Pteridophytesare used for medicinal purposes and as soil-binders. They are also frequently grown as ornamentals. Evolutionarily, they are the first terrestrial plants to possess vascular tissues – xylem and phloem. You shall study more about these tissues in Chapter 6. The pteridophytesare found in cool, damp, shady places though some may flourish well in sandy-soil conditions. You may recall that in bryophytes the dominant phase in the life cycle is the gametophytic plant body. However, in pteridophytes, the main plant body is a sporophyte which is differentiated into true root, stem and leaves (Figure 3.3). These organs possess well-differentiated vascular tissues. The leaves in pteridophyta are small (microphylls) as in Selaginellaor large (macrophylls) as in ferns. The sporophytes bear sporangia that are subtended by leaf-like appendages called
sporophylls. In some casessporophylls may form distinct compact structures called strobili or cones (Selaginella, Equisetum). The sporangia produce spores by meiosis in spore mother cells. The spores germinate to give rise to inconspicuous, small but multicellular,free-living, mostly photosynthetic thalloid gametophytes called
prothallus. These gametophytes require cool, damp, shady places to grow. Because of this specific restricted requirement and the need for water for fertilisation, the spread of living pteridophytes is limited and restricted to narrow geographical regions. The gametophytes bear male and female sex organs called antheridia and archegonia,respectively. Water is required for transfer of antherozoids – the male gametes released from the antheridia, to the mouth of archegonium. Fusion of male gamete with the egg present in the archegonium result in the formation of zygote. Zygote thereafter produces a multicellular well-differentiated sporophyte which is the dominant phase of the pteridophytes. In majority of the pteridophytes all the spores are of similar kinds; such plants are called
homosporous. Genera like Selaginellaand Salviniawhich produce two kinds of spores, macro (large) and micro (small) spores, are known as heterosporous. The megaspores and microspores germinate and give rise to female and male gametophytes, respectively. The female gametophytes in these plants are retained on the parent sporophytes for variable periods. The development of the zygotes into young embryos take place within the female gametophytes. This event is a precursor to the seed habit considered an important step in evolution. The pteridophytesare further classified into four classes: Psilopsida
(Psilotum); Lycopsida(Selaginella, Lycopodium), Sphenopsida(Equisetum)
andPteropsida(Dryopteris, Pteris, Adiantum).
The gymnosperms (gymnos: naked, sperma: seeds) are plants in which the ovules are not enclosed by any ovary wall and remain exposed, both before and after fertilisation. The seeds that develop post-fertilisation, are not covered, i.e., are naked. Gymnosperms include medium-sized trees or tall trees and shrubs (Figure 3.4). One of the gymnosperms, the giant redwood tree Sequoia is one of the tallest tree species. The roots are generally tap roots. Roots in some genera have fungal association in the form of mycorrhiza(Pinus), while in some others (Cycas)small specialised roots called coralloid roots are associated with N
– fixing cyanobacteria. The stems are unbranched (Cycas) or branched (Pinus, Cedrus). The leaves may be simple or compound. In Cycasthe pinnate leaves persist for a few years. The leaves in gymnosperms are well-adapted to withstand extremes of temperature, humidity and wind. In conifers, the needle-like leaves reduce the surface area. Their thick cuticle and sunken stomata also help to reduce water loss.
The gymnosperms are heterosporous; they produce haploid microspores and megaspores. The two kinds of spores are produced within sporangia that are borne on sporophylls which are arranged spirally along an axis to form lax or compact strobili or cones. The strobili bearing
microsporophylls and microsporangia are called microsporangiate or male strobili. The microspores develop into a male gametophyticgeneration which is highly reduced and is confined to only a limited number of cells. This reduced gametophyte is called a pollen grain. The development of pollen grains take place within the microsporangia. The cones bearing megasporophylls with ovules or megasporangiaare calledmacrosporangiate or female strobili. The male or female cones or strobili may be borne on the same tree (Pinus). However, in cycas male cones and megasporophylls are borne on different trees. The megaspore mother cell is differentiated from one of the cells of the nucellus. The nucellusis protected by envelopes and the composite structure is called an ovule. The ovules are borne on megasporophylls which may be clustered to form the female cones. The megaspore mother cell divides meiotically to form four megaspores. One of the megaspores enclosed within the megasporangium
(nucellus) develops into a multicellular female gametophyte that bears two or more archegonia or female sex organs. The multicellular female gametophyte is also retained within megasporangium. Unlike bryophytes and pteridophytes, in gymnosperms the male and the female gametophytes do not have an independent free-living existence. They remain within the sporangia retained on the sporophytes. The pollen grain is released from the microsporangium. They are carried in air currents and come in contact with the opening of the ovules borne on megasporophylls. The pollen tube carrying the male gametes grows towards archegonia in the ovules and discharge their contents near the mouth of the archegonia. Following fertilisation, zygote develops into an embryo and the ovules into seeds. These seeds are not covered.
Unlike the gymnosperms where the ovules are naked, in the angiosperms or flowering plants, the pollen grains and ovules are developed in specialised structures called flowers. In angiosperms, the seeds are enclosed by fruits. The angiosperms are an exceptionally large group of plants occurring in wide range of habitats. They range in size from tiny, almost microscopic
Wolfia to tall trees of Eucalyptus (over 100 metres). They provide us with food, fodder, fuel, medicines and several other commercially important products. They are divided into two classes : the dicotyledons and the
monocotyledons(Figure 3.5). The dicotyledons are characterised by having two cotyledons in their seeds while the monocolyledons have only one. The male sex organ in a flower is the stamen. Each stamen consists of a slender filament with an anther at the tip. The anthers, following meiosis, produce pollen grains. The female sex organ in a flower is the pistil or the carpel. Pistil consists of an ovary enclosing one to many ovules. Within ovules are present highly reduced female gametophytes termed embryo- sacs. The embryo-sac formation is preceded by meiosis. Hence, each of the cells of an embryo-sac is haploid. Each embryo-sac has a three-celled egg apparatus – one egg cell and two synergids, three antipodal cells and two polar nuclei. The polar nuclei eventually fuse to produce a diploid secondary nucleus. Pollen grain, after dispersal from the anthers, are carried by wind or various other agencies to the stigma of a pistil. This is termed aspollination. The pollen grains germinate on the stigma and the resulting pollen tubes grow through the tissues of stigma and style and reach the ovule. The pollen tubes enter the embryo-sac where two male gametes are discharged. One of the male gametes fuses with the egg cell to form a zygote (syngamy). The other male gamete fuses with the diploid secondary nucleus to produce the triploid primary endosperm nucleus (PEN). Because of the involvement of two fusions, this event is termed as double fertilisation, an event unique to angiosperms. The zygote develops into an embryo (with one or two cotyledons) and the PEN develops into endosperm which provides nourishment to the developing embryo. The synergids and antipodals degenerate after fertilisation. During these events the ovules develop into seeds and the ovaries develop into fruit. The life cycle of an angiosperm.
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