Bryophytes: land plants that have a dominant gametophyte generation and lack lignified (true) vascular tissue. Their unbranched sporophytes produce a single sporangium. They do not have true leaves, roots or stems. Gametophyte: the 1n gamete producing generation Gametangium: a general term for a structure where gametes are produced Sporophyte: the 2n generation in which meiosis occurs and meiospores are produced Sporangium: a general term for an enclosure where spores are made Bryophytes Mosses Vascular plants Liverworts lignin at least 475 million years ago Hornworts cuticle sporophyte multicellular gametangia Charophytes Chara Coleochaete Other charophytes Other green algae Evidence that plants are descended from green alga: Green algae have chlorophyll a and b and carotenoids and store starch. All land plants and some green algae have cellulose cell walls and phragmoplasts. All plants are oogamous and they have embryos that depend on the female gametophyte during development. Three lineages of bryophytes: Liverworts (Marchantiophyta) Hornworts (Anthercerotophyta) Mosses (Bryophyta) First appeared 475 million years ago. (mammals appeared 200 M years ago) The ancestors of bryophytes are the green algae. Innovations of the bryophytes: -multicellular gametangia called antheridia and archegonia -multicellular sporophyte that is dependent on the gametophyte -cuticle -stomata -do not have “true” lignified vascular tissue sporangium All bryophytes have the same overall structure. The dominant generation is the gametophyte. The antheridia and archegonia occur on the gametophyte. Fertilization occurs in the archegonium. The sporophyte grows out of the gametophyte. Meiosis occurs in the sporophyte and meiospores are released. Meiospores germinate to become gametophytes. calyptra seta leafy gametophyte rhizoids sporophyte cover cells neck canal neck sterile jacket fertile cells egg cell venter archegonium antheridium The dominant generation in bryophytes is the gametophyte generation. Gemmae are an asexual reproductive structure in bryophytes. Gemmae are multicellular bodies that give rise to new gametophytes. gametophyte thallus gemmae cups pore and air chamber Bryophytes do not have true leaves but there is some cellular differentiation. epidermis chlorenchyma parenchyma Some liverwort gametophytes make special structures to house their gametangia. The male structure is called the antheridiophore and the female structure is called the archegoniophore. thallus archegoniophore gemma cup antheridiophore thallus Liverworts in Glendalough, Ireland The life cycle of the liverwort Marchantia Liverworts, they are primitive but diverse, there are 9000 species. Liverworts are the most primitive (extant) plants. Liverwort sporangia Liverworts, undetermined species from Puerto Rico Liverworts, undetermined species from Dominican Republic Liverworts, undetermined species from Dominican Republic Hornworts - the gametophyte forms symbiotic relationships with cyanobacteria (Nostoc) that fix nitrogen. There are around 100 species of hornworts. Each cell has one large chloroplast (similar to algae) rather than many. columella sporophyte splitting capsule (sporangium) gametophyte Hornwort lifecycle Note, hornworts have stomata on their sporophytes. All other plants have stomata except liverworts. Gametophytes can be unisexual or bisexual. columella Hornwort sporophyte pseudoelaters for spore ejection spore tetrads sporocytes meiosis sporangium (capsule) meristematic region gametophyte tissue foot Life cycle of a moss There are 9500 species of moss. paraphyses antheridia neck archegonium neck canal venter egg cell stalk Fig. 22-15c, p. 396 Moss gametophytes: three phases, protonema, bud and leafy phases. Bud formation is induced by cytokinin. spore protonema bud rhizoids Moss stem showing large epidermal cells specialized for storing water. large epidermal cells leptoids Mosses have some primitive vascular tissue. Hydroids are dead empty cells for water conduction, similar to xylem but not lignified. Leptoids are living and similar to phloem in that sugar can be transported in leptoids. hydroid outside center Asexual reproduction in mosses: - protonema continue to produce buds - leafy tissue can produce protonema - rhizoids sometimes can also produce buds - gemmae can be produced operculum Moss sporangia are complex. They have a cuticle, epidermis with stomata, a cortex, central conducting tissue. The cap is called the operculum, with triangular peristome teeth below it. The teeth move in response to atmospheric humidity to disperse spores. peristome teeth spores Spores can remain dormant for years. columella seta (stalk) tip or operculum capsule peristome teeth capsule peristome teeth Tracheophytes, early seedless vascular plants (Ch 21 in Mauseth) * They have lignified vascular tissue. * Branched sporophytes are independent of gametophyte Water is still required for movement of sperm from the antheridia to archegonia. Monilophytes Earliest trachiophytes are the Rhyniophytes. Only known from fossils, named after Rhynie Scotland. True ferns Horsetails heterospory (in some) Dicotomous branching, no leaves, no roots. They had aerial stems, rhizomes and rhizoids. Ophioglossalean ferns leptosporangia heterospory (in some) Whisk ferns Marattialean ferns megaphylls Isoetes Selaginella (Zosterophyllophyta) (Lepidodendrids) heterospory heterospory Seed plants chloroplast DNA inversion Lycopodium group reduced gametophyte Lycophytes (Other Rhyniophytes) (Aglaophyton) (Rhyniophytes) lignin (Rhynia) strongly lignified vascular tissue (xylem and phloem) branched, independent sporophyte Bryophytes sporangia Rhynia sporophyte (from fossils). Few examples of gametophytes are known. upright stems rhizome rhizoids Rhynia stem cross section epidermis cortex phloem xylem Why did the sporophyte generation become dominant in land plants? There are theories: 1. Diploid cells are protected from mutation by the presence of two copies of each gene? (Increased exposure to solar radiation causes more mutations.) 2. Since sperm required water for dispersal, maybe gametopyte size was more restricted than sporophyte size. And size was important to produce more spores, and later to compete for sunlight. The Lycophytes: True leaves, roots and stems. The bryophytes do not have true leaves, roots or stems. The rhyniophytes are the first to have have true stems with central conducting tissue. Lycophytes are the first to have true leaves, roots, and stems. Lycopodium is homosporous and has subterranean gametophytes and laterally attached sporangia. Selaginella is heterosporous. Isoetes is heterosporous and has secondary growth. Lepidodendron (extinct) 35 m in height, 1m diameter stem True stems, true roots, true leaves. pith primary xylem secondary xylem cambium phloem cortex cork The club mosses, Lycopodium This is Huperzia, another club moss. The sporangia are in the axils of the microphylls. sporangium microphylls Cross section of a microphyll of Huperzia, note single unbranched vein and stomata. air spaces epidermis mesophyll cells stomate vein epidermis air spaces The club moss stem, note central vascular area and lignified cells (sclerenchyma). xylem phloem sclerenchyma parenchyma cortex epidermis parenchyma cortex sclerenchyma phloem xylem Lifecycle of a club moss. Next time, finish Tracheophytes
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