Bryophytes Mosses 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. Vascular plants Liverworts lignin Gametophyte: the 1n gamete producing generation at least 475 million years ago Hornworts Gametangium: a general term for a structure where gametes are produced Sporophyte: the 2n generation in which meiosis occurs and meiospores are produced cuticle Sporangium: a general term for an enclosure where spores are made multicellular gametangia sporophyte Charophytes Chara Coleochaete 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. Other charophytes Other green algae sporangium Three lineages of bryophytes: 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. 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 leafy gametophyte The dominant generation in bryophytes is the gametophyte generation. neck canal neck Gemmae are an asexual reproductive structure in bryophytes. sterile jacket fertile cells Gemmae are multicellular bodies that give rise to new gametophytes. egg cell archegonium sporophyte seta rhizoids cover cells venter calyptra antheridium gametophyte thallus gemmae cups 1 pore and air chamber Bryophytes do not have true leaves but there is some cellular differentiation. 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. epidermis chlorenchyma parenchyma thallus antheridiophore The life cycle of the liverwort Marchantia thallus archegoniophore gemma cup Liverworts in Glendalough, Ireland Liverworts, they are primitive but diverse, there are 9000 species. Liverworts are the most primitive (extant) plants. Liverwort sporangia 2 Liverworts, undetermined species from Puerto Rico 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 Liverworts, undetermined species from Dominican Republic columella Hornwort lifecycle Hornwort sporophyte Note, hornworts have stomata on their sporophytes. All other plants have stomata except liverworts. Gametophytes can be unisexual or bisexual. pseudoelaters for spore ejection spore tetrads sporocytes meiosis sporangium (capsule) meristematic region gametophyte tissue foot 3 Life cycle of a moss paraphyses There are 9500 species of moss. antheridia Moss gametophytes: three phases, protonema, bud and leafy phases. Bud formation is induced by cytokinin. neck spore protonema bud archegonium neck canal venter egg cell rhizoids stalk Fig. 22-15c, p. 396 4 Moss stem showing large epidermal cells specialized for storing water. 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 large epidermal cells outside center operculum Asexual reproduction in mosses: - protonema continue to produce buds - leafy tissue can produce protonema - rhizoids sometimes can also produce buds - gemmae can be produced 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 peristome teeth capsule capsule 5 Tracheophytes, early seedless vascular plants (Ch 21 in Mauseth) peristome teeth * 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. sporangia True ferns Horsetails heterospory (in some) Dicotomous branching, no leaves, no roots. They had aerial stems, rhizomes and rhizoids. Ophioglossalean ferns leptosporangia Whisk ferns heterospory (in some) Rhynia sporophyte (from fossils). Few examples of gametophytes are known. Marattialean ferns megaphylls Isoetes Selaginella (Zosterophyllophyta) (Lepidodendrids) heterospory heterospory Seed plants chloroplast DNA inversion Lycopodium group upright stems reduced gametophyte Lycophytes (Other Rhyniophytes) (Aglaophyton) (Rhyniophytes) lignin (Rhynia) rhizome strongly lignified vascular tissue (xylem and phloem) branched, independent sporophyte Bryophytes Rhynia stem cross section rhizoids Why did the sporophyte generation become dominant in land plants? epidermis cortex phloem 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. xylem 6 Lepidodendron (extinct) 35 m in height, 1m diameter stem 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. True stems, true roots, true leaves. pith primary xylem secondary xylem Lycopodium is homosporous and has subterranean gametophytes and laterally attached sporangia. cambium phloem cortex cork Selaginella is heterosporous. Isoetes is heterosporous and has secondary growth. The club mosses, Lycopodium This is Huperzia, another club moss. The sporangia are in the axils of the microphylls. sporangium microphylls epidermis air spaces The club moss stem, note central vascular area and lignified cells (sclerenchyma). Cross section of a microphyll of Huperzia, note single unbranched vein and stomata. xylem phloem sclerenchyma parenchyma cortex epidermis parenchyma cortex air spaces epidermis sclerenchyma phloem mesophyll cells stomate vein xylem 7 Lifecycle of a club moss. Next time, finish Tracheophytes 8
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