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Mosses and Ferns Evolutionary developments necessary for plants to colonize land Stage One: Becoming multicellular Occurred in water Enables specialized tissues to develop Stage Two: Developing sporangia Enables dispersal on land Stage Three: Developing a large sporophyte Confers competitive advantage Provides perennial spore production Stage Four: Removing dependence of fertilization on a film of water Enables survival in dry environments History of evolution of major plant types on land Stage One of adaptation to living on land The land that plants colonized was hostile to life. Soil development was minimal. Land plants required several adaptations to be successful that require multi-cellular tissues : mechanical strength for support, exposed light catching surfaces, anchoring system, conducting system for water, system for obtaining mineral nutrients, a way to restrict water loss in desiccating air, a means of reproducing and dispersing on land Devonian plant community found at Rhynie, in Scotland. A reed-like marsh, 370-380 million years ago. Asteroxylon MAIN FEATURES Simple dichotomous branching Sporangia !5 to 30 cm tall No roots Stomata with guard cells Most had a central vascular strand Cuticle Devonian plant community Asteroxylon had leaves – without a vascular connection Plants living in water release spores and gametes that swim and may be helped to dispersed by water movement To live on land plants faced two challenges for their reproduction: 1. Dispersal 2. Fertilization Dispersal was solved first – through production of sporangia. angeion is Latin for case So a sporangia is a spore case The important feature of sporangia is that they lift spores above the ground so they can be dispersed by the wind Stage Two: Developing sporangia Mosses Retention of the zygote by the female gametophyte Delayed meiosis and growth of the sporophyte by mitosis Developing sporophyte Zygote Gametophyte Archegonium Meiosis in the sporangium producing haploid spores. 1. Dispersal Eight Terms to Learn to understand Alternation of Generations of Land Plants Spores – haploid, single cells produced by meiosis The word “phyte” is Greek for plant Gametes – collective term for sperm and egg Gametophyte – haploid plant that develops from a spore and produces gametes by mitosis Gametangium – a “case” holding gametes Archegonium – flask–shaped container holding the egg cell. (Ancient gonad) The female gametangium. Antheridium – The male gametangium Sporophyte – diploid plant that grows from the zygote and produces spores by meiosis Sporangium – the “case” holding spores Moss life cycle Fig. 25.4, p. 406 Mature sporophyte (sporeproducing structure and stalk), still dependent on gametophyte. Zygote grows, develops into a sporophyte while still attached to zygote gametophyte. Diploid Stage Fertiliztion Sperm reach eggs by moving through rain drops or film of water on the plant surface. Haploid Stage rhizoid sperm-producing structure at shoot tip of male gametophyte. egg-producing structure at shoot tip of female gametophyte. Meiosis Spores form by way of meiosis and are released. Spores germinate. Some grow and develop into male gametophytes. Other germinating spores grow and develop into female gametophytes. Moss sporophyte Top of capsule Developing protonema Moss antheridium and archegonium Important life cycle features of mosses Spores n Development of Mitosis gametophyte thallus Female gamete(?) Gametes remains attached to the haploid thallus Meiosis HAPLOID Fusion (syngamy) DIPLOID Development of sporophyte thallus – remains attached to gametophyte Zygote 2n Mitosis Dessication tolerance in Tortula ruralis RAPID WATER LOSS Constitutive Cellular Induction of Hormone ? Protection Recovery and Repair Mechanisms Hydrated Dry Rehydrated Spagnum – the bog-forming species The species forms clumpsminimizing surface area to volume ratio. Unique leaf cells (hyaline cells) of Spahgnum species enable the plant to absorb up to 20 times its own dry weight of water. Fig. 25.5, p. 407 How can we characterize mosses? 1. Plants accumulate matter and make growth 2. Plant growth is an organized process following rules of anatomy and morphology 3. Plants maintain their heat and water balance 4. Plants have a life cycle with reproduction and dispersal 5. Evolution is a constant process Stage Three: Developing a large sporophyte In ferns the sporophyte is only dependent on the gametophyte for obtaining nutrient, water, and physical support when it is first formed. Sporophyte originally grows from a gametophyte and then develops roots, rhizome, and fronds Ferns Large size enables competition as well as effective spore dispersal. The perennial root stock enables continued frond and spore production from year to year Fronds growing from a rhizome Fern life cycle The sporophyte (still attached to the gametophyte) grows, develops. sorus (one of the spore-producing structures) zygote Diploid Stage fertilization egg sperm Haploid Stage eggproducing Archegonia structure spermproducing structure Antheridia meiosis Spores develop. mature gametophyte (underside) Spores are released Spore germinates, grows into a gametophyte. http://departments.bloomu.edu/biology/chamuris/concepts2/labimg.html Sporangia Polypodium spp sori sporangia A sorus Polypodium spp a g n i f i e d Sporangia Developing spores Polypodium spp Gametophyte Gametophyte Developing sporophyte Arrangement of sporangia on two ferns In lines on a broadleaved type Asplenium At the end of the leaves Adiantum Tree ferns Cibotium menziesii in habitat in Hawaii. Photo courtesy of Peter Richardson. Conducting tissue Cyathea australis with the uncurling croziers visible. Photo courtesy of Scott Ridges How can we characterize ferns? 1. Plants accumulate matter and make growth 2. Plant growth is an organized process following rules of anatomy and morphology 3. Plants maintain their heat and water balance 4. Plants have a life cycle with reproduction and dispersal 5. Evolution is a constant process Coal formation Jungle-like forests of the Carboniferous were dominated by giant ancestors of club mosses, horsetails, ferns, conifers, and cycads. Most of the plant fossils found in the coals and associated sedimentary rocks show no annual growth rings, suggesting rapid growth rates and lack of seasonal variation in the climate (tropical). Anaerobic conditions and periodic inundations of the sea Early Carboniferous Equator Appalachians Britain Ice cap Equator Appalachians Britain Ice cap