Microgametophyte Example. What Makes Them Different? Megagametophyte. The Seed

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Wrap-up of cryptogams Introduction to Seed Plants How seeds allowed the full colonization of land by plants Free living gametophyte Gametophytes independent of parent sporophytes May be small & inconspicuous, but still free-living Have motile sperm AND require water for fertilization Sperm swim through open environment to reach egg Therefore, not fully terrestrial Vestiges from aquatic heritage Only with development of seeds do plants become fully terrestrial Advantages of Seeds Mechanism for fertilization without water Sperm do not swim through open environment Exploit new habitats (fertilization times, places, etc.) Provide new method for dispersal Move into new areas Open specialized dispersal mechanisms (wind, animals) Dormancy: wait for favorable conditions Unlike (most) cryptogams Provide nutrition to developing sporophytes Increase survivability; dormancy Advantages of Seeds For these reasons, seed plants are now dominant Replaced cryptogams in most areas Often restricted to specialized habitats, now Other adaptations: Axillary branching vs. uneven dichotomous branching Secondary xylem & phloem production Development of bifacial vascular cambium

What Makes Them Different? All are heterosporous Spore & gametophyte types modified Pollen not unlike microgametophyte of Selaginella Sperm do not swim through open environment Entire microgametophyte delivered to area of the egg Sperm move through pollen tube to the egg Pollen = immature microgametophyte Pollen grain with pollen tube = mature microgametophyte Microgametophyte Example Megagametophyte A new outer, protective layer surrounds the megasporangium = integument OVULE = integumented megasporangium Megagametophyte is endosporic, but no longer free living Nutritionally dependent on sporophyte, and not released from sporophyte tissue See Seed Plants handout The Seed A SEED is an embryo (young sporophyte) encased in the megagametophyte (nutritive tissue), all enclosed by seed coat (remnants of nucellus & integument of old sporophyte generation) Contains 3 generations in one: Integument(s) = seed coat - remnants of megasporangium (2n) [grandmother] Megagametophyte = nutritional source for developing embryo (1n) [mother] Embryo = young sporophyte (2n) [daughter]

Major Seed Plant Lineages Gymnosperms naked seeds [Greek, gymnos = naked] Borne in exposed position, e.g., sporophylls, scales, etc Angiosperms seeds inside vessels [Greek, angios = vessel] Enclosed in a carpel Seed Plant Lineages Gymnosperms Pinophyta Gnetophyta Ginkgophyta Cycadophyta Angiosperms Magnoliophyta G. Chandler Oldest, Single, Living Organism Perfect Pinophyta Pinus longaeva (bristle-cone pine)

Tallest Living Organism Eucalyptus regnans Tallest recorded tree Chopped down ~ 135 m high [443 ft.] Cupressaceae Sequoia semperivirens (coastal redwood) Pinophyta Largest division of extant gymnosperms 600 living spp. [spp. = species, plural] Secondary growth common Wood Greatest distribution among extant gymnosperms Especially in N. boreal regions Coniferous forests for a nearly continuous ring around the N. Hemisphere (= boreal coniferous forest Pinophyta Also found in more temperate areas, e.g., "pyro-phytic" pinelands of SE U.S. Fire-maintained ecosystems Where fire is eliminated, replaced by angiosperms Mostly monoecious (some dioecious) Pollen grains have a pair of sac-like appendages (lost in some lineages)

Pinophyta Economics Timber Pinus, Pseudotsuga, Picea, Tsuga, Larix, Abies Pulp (for paper) Chemicals Resins, turpentine, oils; "pitch" from pines used to coat naval ships Important draw for England to SE Va. Capt. John Smith Ornamentals Pine nuts Pinus edulis seeds (NOT nuts!) Pinophyta Taxonomy Taxopsida Taxaceae Pinopsida Pinaceae Cupressaceae Podocarpaceae Araucariaceae * Phyllocladaceae * Cephalotaxaceae * Pinaceae (10 gen., 220 spp.) Pinus (pines) 100 spp. Abies (firs) 40 spp. Picea (spruces) 40 spp. Tsuga (hemlocks) 10 spp. Pseudotsuga (Douglas fir) 5 spp. Larix (larches) 10 spp. Cedrus (true cedars) 4 spp. Vegetative Morphology: Pinus Pinus (& some others) have 2 types of shoots: long & short Resin canals 3 types of leaves Scale (cover long shoots) Sheath (cover short shoots) Needle (photosynthetic) Evergreen Leaves not shed all at once

Long & Short Shoots Pinus Leaf Types G. Chandler Long shoot Needle leaves Short shoot Sheath leaves Scale leaves G. Chandler G. Chandler Pinus Leaf Types Pine Needle Leaves Sheath leaves Needle leaves Scale leaves

Reproduction in Pinaceae Monoecious (male & female on same plant) Microstrobilus = pollen cone Megastrobilus = pine cone Microstrobilus Simple strobilus 1 axis with sporophylls Pollen with bladders Immature microgametophyte Microgametophyte Shed at 4 nucleate stage (immature) Fully mature when pollen tube grows 6 nuclei (including 2 sperm nuclei/cells) Megstrobilus Compound cone 2 axes Main axis bract Lateral axes = branches not leaves (scales) ovule Bracts represent leaves See handout on Pinus gametogenesis scale Immature Mature

Young Ovule Pinaceae Examples Wing of seed derived from cone scale In angiosperms, would be an outgrowth of the fruit Some keep scales E.g., Pinus, Picea Some cone scales deciduous E.g., Cedrus, Abies scale ovule bract Pseudotsuga Douglas Fir Picea Spruce Deciduous Leaves Cedrus True Cedars Tsuga Hemlock Abies Firs Larix Larch

Pinopsida: Cupressaceae Cupressaceae Important woods! Many are rot- and termite-resistant Cupressus (true cedars) Juniperus (junipers, E red cedar) Thuja (arbor vitae, W red cedar, white cedar) Chamaecyparis (white cedar) Taxodium (bald cypress) Sequoia (coastal redwood) Sequoiadendron (mountain redwood) Taxodium Pinopsida: Podocarpaceae Seeds borne on swollen tip of branch: the aril Dispersal of seed by animals NOT the same as a fruit! Seed still naked

Taxopsida: Taxaceae Only family in class Also has aril! Convergence Aril surrounds seed Torreya californica Taxus Taxus (yews) Torreya (stinking cedar; California nutmeg)