BIOL 1030 Topic Four Notes
BIOL 1030 Topic Four Notes BIOL 1031 - 001
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Date Created: 02/15/16
BIOL1030 Topic Four Notes Most of the plants discussed so far do not have specialized gametophytes They are homosporous plants: make one kind of meiospore Principally homosporous groups o Bryophyta o Hepatophyta o Psilophyta o Arthrophyta Heterosporous plants: make two types of meiospores o One becomes male gametophyte and makes sperm o One becomes female gametophyte and makes eggs Thus, make specialized spores and specialized gametophytes o Specialization occurs in the spore to gametophyte movement on the haploid side Heterosporous groups: o One member of Arthrophyta o Some Lycophyta o Some Pterophyta o All seed plant phyla HETEROSPORY Note deadbeat gametophytes o Not free-living. Dependent on sporophyte o Not (very photosynthetic) o Small: microgametophyte reduced to one antheridium, if that. Megagametophyte develops mostly within megaspore Spores specialized o Megaspore makes megagametophyte, which makes archegonia with eggs o Microspore makes microgametophyte, which makes sperm (sometimes in an antheridium) Valuable ideas for life on the land: o Gametophytes small and protected by spore walls o Deadbeat gametophytes fed by big, strong sporophyte (gametophytes can focus on reproduction) o Megaspore: keep it after fertilization and feed embryo. Embryo starts development inside megaspore and protected by its walls Lets new embryo “eat” its mother (megagametophyte) Keeps energy used to make megagametophyte from being wasted o Microspore: disperse it through air (or other ways we will discuss later), instead of using delicate swimming sperm PLANTS Nonvascular plants o Covered, spores used to make free-living gametophytes. Sporophytes dependent on gametophytes Seedless vascular plants o Covered, spores used to make free-living gametophytes. Sporophytes and gametophytes both can live independently Seed plants o Rest of plant kingdom; gametophytes dependent on sporophyte (non free-living) SEED PLANTS All are heterosporous vascular plants Three major reproductive adaptations o Gametophyte reduced to dependence on sporophyte; retained in moist reproductive tissue o Seed-“baby plant in a lockbox with its lunch”; highly resistant structures that allow for a dormant phase in the life cycle to wait out poor environmental conditions o Evolution of pollen as male gametophyte—many seed plants are no longer tied to external water for fertilization Common ancestor with seeds gave rise to all seed plants (gymnosperms and angiosperms) Together, gymnosperms and angiosperms are a monophyletic group Extant gymnosperms form a clade Group is a grade if you include extinct forms Fossil evidence indicates origins in progymnosperm group about 360 MYA Gametophytes are completely dependent on parent sporophytes for nutrition and are composed of only a few cells Male gametophytes develop from microspores o Become pollen grains o Entire male gametophyte moved to the female as pollen grains o Cannot perform photosynthesis, depends on nutrients that came from the parent sporophyte Means of transporting sperm to egg varies, but typically uses a growing pollen tube that does not require outside water Moving pollen to vicinity of ovule called pollination; agents include wind, animals Female gametophytes develop from megaspores within ovules o Ovule contains female gametophyte surrounded by nucellus (megasporangium) o Integuments o Micropyle o Cannot perform photosynthesis, depends on nutrients from the parent sporophyte Seed develops from ovule o Seeds are highly resistant structures that allow for a dormant phase in the life cycle to wait out poor environmental conditions Embryo protected by a seed coat, an extra layer of hardened tissue derived from sporophyte tissue in the ovule (sporophyte tissue from parent, not from embryo) Seed coat o Enhanced protection from drought, cold, heat o Some protection from pathogens and predators o External water only needed at germination o Initial food supply for germinating plant is enclosed Seeds replace spores as means of dispersal; can enhance means of dispersal Seed plants together are a monophyletic group Divided into two “groups” based on whether or not ovule is completely enclosed by sporophyte tissue at time of pollination o Gymnosperms—“naked seed” o Angiosperms—“covered seed” GYMNOSPERMS There are almost certainly some extinct groups that would be classified as gymnosperms, and some of those were almost certainly more closely related to angiosperms than to extant gymnosperms Thus, gymnosperms would be a grade if several groups had not died of The gymnosperm group is composed of four phyla with living members o Essentially, all seed plants that are not angiosperms o All lack flowers and fruits that are found in angiosperms o Ovule not completely enclosed by sporophyte tissue at time of pollination o Instead, ovule sits exposed on a scale (a modified leaf) The Gymnosperm Revolution and the Permian Period Permian Period (290-248 MYA) followed Carboniferous: was relatively cool and dry. Most coal swamps dried up Many SVPs (especially tree species) went extinct; gymnosperms prospered Why? Pollen and seeds adapted to dry conditions Four Phyla: o Phylum Coniferophyta (the conifers) o Phylum Cycadophyta (the cycads) o Phylum Ginkgophyta (Ginkgo) o Phylum Gnetophyta (the gnetophytes) CONIFEROPHYTA (the conifers) Monophyletic group ~600 living species; worldwide distribution, more common in cold or dry regions Pines, spruces, firs, cedars, junipers, hemlocks, yews, larches, cypresses, redwoods Nearly all are evergreen Many have needle-shaped leaves adapted to dry conditions (resistant to water loss) o Thick cuticle o Stomata in pits Importance: o Conifer forest biome (covers large part of Earth’s landmass) o Some communities in southeast dominated by conifers o Currently dominant in parts of North (and South) Temperate Zones Superlatives: o Biological records (tallest living tree) Coast redwood (Sequoia sempervirens) Grow on coast of California/Oregon 96% have been logged Tallest coast redwood tree is 368 feet tall o Biological records (smallest trees?) Pygmy forest in Mendocino County, CA o Biological records (largest tree) Sequoiadendron gigantean (big tree) In Sequoia National Park, CA Base: 102 feet circumference Weight: 2,145 tons Volume: 52,500 cubic feet o Biological records (oldest tree) o Bristlecone Pine (Pinus longaeva). Found in high mountains of deserts of California, Nevada, Utah, Colorado, etc Oldest tree was 4,950 years. Cut down in 1964 by graduate student who got corer stuck in tree (USFS gave permission) Now oldest known bristlecone (named Methuselah) is 4,840 years old, Location kept secret from public Sources of timber, paper, resin, cancer drug taxol, etc. “Soft” wood (unlike angiosperm trees, no vessels or fibers in xylem) Economic impact: o Harvest for pulp and timber o Can cause controversy, especially when old-growth and clear-cuts are involved o Importance in Alabama: forestry #1 legal agricultural crop o Many pine plantations in the southeast o Medicines: example, taxol o Antitumor agent. First extracted from bark of Pacific Yew (Taxus) tree o Now made synthetically Importance: o Threatened by humans (acid rain and forest decline) o Acid rain areas (below) include some conifer forests o Acid rain and other stresses cause “forest decline”, weakness and death of conifer trees o Seen in N. Europe and N. America Pines as a representative group o Over 100 species o Native to Northern hemisphere o Typically thick bark (survive fires, drought) o Secrete resin from leaves and bark Response to wounding Deters fungal and insect attacks Source of turpentine (volatile liquid, organic solvent) and solid rosin Pine tree is sporophyte, with sporangia located on cones Gametophyte generations reduced; retained within sporangia o Male gametophyte is pollen grain (no antheridium) o Female gametophyte produces archegonia within ovule Heterosporous: separate male and female cones Male cones (pollen cones) o Clusters of 30-70, usually at tips of lower branches o 1-4 cm long: papery scales in spirals or whorls o Pair of microsporangia sacs within each scale o Microspore mother cells in microsporangia form haploid microspores o Each microspore becomes 4-celled pollen grain (via 2 rounds of mitosis) o Pollen grain carried by wind (pair of air sacs provides buoyancy) o One cluster of pollen cones can yield over 1 million pollen grains Female cone (ovulate cones): o Cone scales with ovules on them o Typically on upper branches of same tree with pollen cones o Larger than pollen cones; scales become woody (highly lignified) o Pair of ovules develop at base of each scale Megasporangium called nucellus embedded in each ovule Nucellus is nutritive tissue surrounded by thick integument (covering) with hole (micropyle) near one end One layer of integument later becomes seed coat o Single megaspore mother cell in each megasporangium o Produces 4 haploid megaspores; 3 break down o Surviving megaspore develops over about one year into female gametophyte with sometimes thousands of cells o Female gametophyte has 2-6 archegonia, at micropylar end o Each archegonium has one large egg (visible without a microscope) o Female cones take two or more seasons to mature Female gametophyte has several arhegonia, each with an egg Reproduction o Scales of ovulate cone open o Pollen lands near micropyle, caught by sticky fluid o Evaporating fluid pulls pollen through micropyle to ovule o Scales close (female gametophyte not mature) o Pollen grain germinates, forming pollen tube that digests through nucellus (takes about 15 months to reach archegonium) o One of the four pollen grain cells (the generative cell) undergoes mitosis; one of the products divides again, making two sperm cells o Mature male gametophyte is germinated pollen grain with pollen tube and sperm o When archegonium is reached, one sperm fertilizes egg Zygote develops into embryo as surrounding ovule becomes seed o Usually only one successful zygote per ovule o Embryo (new sporophyte, 2N) has rudimentary root, several embryonic leaves (cotyledons) o Food source in seed drives from rest of female gametophyte (1N) (eats its mother) o Integument (2N tissue from parent sporophyte) becomes hard protective seed Scales of cone open and separate; winged seeds disperse From initial ovulate cone formation to final seed production 3 years or more Pollen grain is immature male gametophyte (microgametophyte) Male gametophyte matures in ovule, when pollen tube and sperm cell are made. Entire male gametophyte is just a few cells Ovule contains megasporangium. Meiosis produces megagametophyte, which makes archegonia, still kept in megasporangium Fertilization takes place and embryo digests female gametophyte Seed is baby plant (embryo), in box (seed coat from parent sporophyte tissue), with its lunch (female gametophyte) Male gametophyte (immature) small, protected by microspore wall, flies through air to female gametophyte No swimming sperm; male gametophyte delivers sperm to female gametophyte protected by ovule in female cone Embryo nourished b megametophyte and protected by cone of sporophyte Seed has hard protective coat, supply of food (megametophyte); can be dispersed to a new location Embryo can be dormant: allows new sporophyte to wait for good conditions to germinate CYCADOPHYTA (the cycads): Monophyletic group ~200 living species, tropical and subtropical; many in danger of extinction Along with conifers, dominated Mesozoic era (245 MYA-65 MYA) Slow-growing; some grow >15 m tall Most resemble palm trees, but produce cones (female cones up to 45 kgs, or 100 lbs) Have life cycle similar to pines Unusual sperm o Have thousands of flagella arranged in spirals o Swim within ovule to archegonium o Largest sperm known GINKGOPHYTA (Ginkgo): Monophyletic 1 living species, Ginkgo biloba (also known as the maidenhair tree) Exists only in cultivation (no natural native populations remain); first cultivated in Japan and China Deciduous—lose leaves annually Flagellated sperm (similar to cycads) Dioecious Stinky seed coverings (produce butyric acid, making the smell of rancid butter) Males often planted on city streets (do not stink like females; resistant to air pollution) GNETOPHYTA (the gnetophytes): Modern genetic studies support this as a monophyletic group ~70 living species in 3 genera: Welwitschia, Ephedra, and Gnetum Based on physiology was once considered paraphyletic with angiosperms as closer relatives to Gnetum than the rest of the group o Vessels in xylem of Gnetum (common in angiosperms, found only in these gymnosperms) o Members of Gnetum have broad leaves similar to angiosperm leaves Welwitschia—bizarre plants of southwest African deserts o Stem is shallow cup that tapers into a taproot o Two leathery leaves (often split) grow continuously from base o Conelike reproductive structures at leaf base o Dioecious—separate male and female plants Ephedra common in Mexico an southwestern US, but found on most continents o Shrubby, stems resemble horsetails (jointed, with tiny scale-like leaved at each node) o Some species monoecious (male and female parts on same plant), some dioecious o Drug Ephedrine, historically extracted mainly from a Chinese species of Ephedra KINGDOM PLANTAE: ANGIOSPERMS ANTHROPHYTA (flowering plants): Anthro- flower Also known as angiosperms (angeion- vessel or enclosure; sperma- seed) Ovules enclosed within carpel (parent diploid sporophytic tissue) at pollination o The “vessel” is the carpel, which is a modified leaf o Carpels, especially their enlarged basal portion (the ovary), usually develop into fruit, which is unique to angiosperms About 250,000 known living species (dominant photosynthetic organism on land) Predominant source of human food Most widespread and diverse plant phylum o Range from microscopic to plants with leaves over 6 m long o Flowers show incredible variety from species to species o Variety of lifestyles Monophyletic group with seeds, refined xylem, double fertilization, and these synapomorphic characteristics: o Seed contains endosperm o Presence of flowers (modified stems and leaves) o True fruits Evolutionary history o Monophyletic group o Origin about 140 MYA Historically divided into two classes, dicots and monocots o Recent genetic analysis has shown that the traditional dicots are a paraphyletic group o Thus, the old classification scheme is being replace No conclusive cladogram has been produced for angiosperms o Studies are ongoing o Most modern cladograms have Amborella as a sister group to the rest of the angiosperms o Within the rest, water lilies are a sister group to the others Various class-level groupings have been proposed o The overall naming and formal classification within Phylum Anthophyta is still in a state of flux o Nevertheless, by far most of the living angiosperms species are found within two monophyletic groups, eudicots and monocots Eudicots o Most have embryo which have two cotyledons (seed leaves) o Leaves have netlike veins o Flower part typically in multiples of four or five o Groups of vascular tissue in a ring o Pollen grains mostly with 3 or more apertures o Endosperm mostly used up in mature eudicot seeds o About 175,000 living species; includes nearly all flowering trees and shrubs o About a sixth are annuals (entire growth cycle in one year or less) Monocots o Embryos have one cotyledon o Leaves have essentially parallel veins o Flower part typically in multiples of 3 o Groups of vascular tissues scattered o Pollen grains mostly in one aperture o Endosperm typically present in mature monocot seeds o About 65,000 living species o No true wood (bamboo is not true wood) o Few annuals o Examples: grasses (including many crop plants such as corn, wheat, etc.); lilies, irises, orchids, onions, and many more 130 MYA two major continental masses o Laursia- North America, Europe, and Asia o Gondwanaland- South America, Africa, Australia, Antarctica, India, and New Zealand The geographic and temporal origin of the angiosperms remains under considerable debate Clear angiosperm fossils date back at least to 125 MYA in Laurasia Advantages of flowering plants o Transfer of pollen over great distances promotes outcrossing o Efficient seed dispersal via fruit o Endosperm gives seedlings a fast start o Leaves appropriate for fast growth in hot, dry environment Coevolution with insects o Dominant by ~80 MYA, second half of Cretaceous Period o All present angiosperm families represented by this time o Many insect orders appeared or became more abundant at this time FLOWERS Modified stems with modified leaves Develop as primordium bud at end of stalk called pedicel Pedicel widens to form receptacle Other flower parts attached to receptacle in four whorls; from outside in: o Calyx—sepals o Corolla—petals o Androecium—stamens o Gynoecium—carpels Calyx—sepals; usually green, leaf-like, and protect immature flower Corolla—petals; usually colorful, attract pollinators, together with calyx called perianth Androecium—stamens, male reproductive structures o Filament and anther o Microspores produced within anther, shed as pollen Gynoecium- female reproductive structure o Center location is most protected o Formed from leaf-like structure with ovules along margin o Edges fold inwards around ovules, forming carpels o Primitive: many separate carpels o Advanced: carpels fused (called pistil) Carpel/pistil segments o Ovary- swollen base with 1 to hundreds of ovules; develops into fruit o Stigma- tip; sticky and/or feathery to catch pollen o Style- usually present; separates stigma from ovary Nectaries may be present at base of pistil; secrete sugar, amino acids, and other compounds to attract pollinators Female gametophyte o Single diploid megaspore mother cell in ovule undergoes meiosis while flower develops o Of four haploid megaspores produced, usually three break down o Remaining megaspores expands and replicates and divides until there are eight haploid nuclei in two groups of four o One nuclei from each group migrates toward center; these are polar nuclei o Polar nuclei usually fuse to make a diploid nucleus, but may remain separate—in either case, they wind up in a single cell o Cell walls form around other nuclei, creating the 7-celled, 8-nucleate embryo sac or megagametophyte (female gametophyte) o Meanwhile, two layers (integuments) of ovule develop into seed coat with micropyle (small opening) o In the megagametophyte, one of the cells closest to the micropyle becomes the egg; the other two there are synergids o The three cells on the other end (the antipodals) eventually break down Male gametophyte o Anthers with patches of tissue that become chambers lined with nutritive cells o Each patch has many diploid microspore mother cells o Microspore mother cell undergoes meiosis, making 4 haploid microspores that typically remain grouped in a tetrad o Each microspore nucleus replicates and divides once (via mitosis) without cytokinesis (meaning they remain as one cell with two nuclei, a binculeate microspore) o Usually, tetrad breaks up o Two-layered wall develops around each binucleate microspore, now called a pollen grain Pollen grain o Outer wall—sculptured, appearance usually species specific, often has chemicals that can react with an appropriate stigma to stimulate pollen tube formation o Apertures in outer wall—where pollen tube may grow out; eudicots—usually 3, monocots—usually one Pollination- transfer of pollen to a stigma o Usually between flowers of separate plants o Agents include wind, water, gravity, mammals, bird, insects o Various reward systems for animal agents (pollen, nectar, etc) o Evolution of floral characteristics associated with pollination o Some plants self-pollinate (inbreeding) - pollen to same plant o Pollination followed by fertilization only if chemical signals are right Fertilization o Pollen grain cytoplasm absorbs substances from stigma o Bulge forms through an aperture in pollen grain; becomes a pollen tube o Pollen tube follows chemical gradient through style to micropyle Chemicals difuse from embryo sac Micropyle usually reached within a few days (up to a year in some species) o Pollen grain has two nuclei; one, the generative nucleus, lags behind o Generative nucleus undergoes mitosis to make two non- flagellated sperm; this may occur in pollen grain or in pollen tube (male gametophyte now mature) o Pollen tube enters embryo sac, destroying a synergid o Double fertilization – essentially unique to angiosperms One sperm unites with egg, forming zygote Other sperm unites with polar nuclei, forming 3N primary endosperm Primary endosperm rapidly undergoes many cycles of mitosis, forming endosperm Endosperm provides nutrients for embryo; in many seeds, it is gone by the time the seed is mature o Seed coat hardens o Remaining haploid cells degenerate o Now have seed with 2N embryo, 3N endosperm, and 2N seed coat (seed coat from parent female tissue) LIFE CYCLE Zygote grows into embryo, endosperm also grows Embryo uses endosperm for nourishment (eats sibling) Seed: baby plant (embryo), in box (seed coat, make from integuments), with its lunch (endosperm) Seed or seeds develop inside of ovary Ovary wall in pistil becomes pericarp in fruit SEEDS Embryo – quickly forms all systems, then growth arrested (dormancy)—mature seed about 10% water, very low metabolic activity Typically, dormancy occurs just after first leaves (cotyledons, or seed leaves) form Stored food (in angiosperms, 3N endosperm and/or cotyledons) Seed coat- tough, relatively impermeable o Protection from predators, pathogens o Protection from desiccation, harsh conditions (crucial on land) o May allow seed to last hundreds of years Dormancy broken only when conditions are right (seed bank in soil) Germination=breaking dormancy=resuming metabolic activity, growing out of seed coat; occurs after water penetrates seed coat to embryo, bringing oxygen FRUITS—Mature Ovaries Terms below are examples only, don’t try to memorize all of these for exams Fleshy—pomes (apples), drupes (peaches), hesperidiums (oranges), pepos (melons, gourds), aggregate fruits (strawberries, raspberries), multiple fruits (pineapple, fig) Dry—follicles (milkweed, magnolia), legumes (peas, beans), siliques and silicles (mustards), capsules (irises, lilies, orchids), caryopses (grasses), nuts (chestnuts, hazelnuts, acorns), achenes (sunflowers), samaras (maples, elms, ashes), schizocarps (parsleys) Dispersal o By wind Wings- maples Parachutes- dandelions, milkweeds Dust-like seeds- orchids o By water- coconuts o By vertebrates Fleshy, edible fruits (blue, black, red)—seeds often deposited in feces Dry, edible- nuts, others—squirrels bury and forget about them Dry, inedible- hooks to grasp hair, feathers (cockleburs, etc.) o By explosive dehiscence (jewelweed, others) Stolons- runners – long slender stems that grow along soil (strawberries) Rhizomes—underground stems—common in grasses; bulbs and tubers are rhizomes specialized for storage (potatoes) Suckers—roots produces sprouts that grow into new plants (apple, raspberry, banana) Adventitious leaves—numerous plantlets develop from tissue in notches along leaves Apomixes—embryos in seeds may be produced sexually Artificial: cuttings (for some species, can get roots to grow with appropriate environment) FLORAL EVOLUTION First flowers o Numerous spirally arranged sepals, petals, stamens, and carpels o Petals and sepals similar in color and form o All parts free (not fused) Parts o Complete- calyx + corolla + androecium + gynoecium o Incomplete- one or more whorls absent o Perfect- has both androecium and gynoecium o Imperfect- missing either androecium or gynoecium o Complete flowers are always perfect; incomplete flowers can be either perfect or imperfect Trends o Separate floral parts grouped together or fused Connation- fusing within a whorl Adnation- fusing between whorls (for example, sepals and petals fused together) o Reduction or loss of floral parts o Bilateral symmetry instead of radial symmetry Ancestral type: radial symmetry; examples: buttercups, geranium, lilies Derived type: bilateral symmetry; examples: snapdragons, orchids Bilateral symmetry in some cases has arisen independently in diferent groups Pollination Mechanisms (pollination syndromes) o Wind—passive, primitive (oaks, cottonwoods, birches, grasses) Copious amounts of pollen Most pollen travels no more than 100 m Flowers small, greenish, odorless Corollas reduced or absent Often grouped in large numbers, may hand gown with tassels that wave in wind and shed pollen freely Male and female parts often well separated on plant to reduce chance of self-pollination Often flower before leaves grow—keeps leaves out of the way o Animals—some cycads and gnetophytes also have this, so symplesiomorphic trait Bees—most numerous of insect-pollinated plants use bees Find via odor Orient via shape, color, and texture Usually blue or yellow flowers, bee sees in ultraviolet Many have stripes or lines of dots (nectar guides) to indicate nectaries Nectar ofered as food for bees (pollen also) Often close association between a bee species and a plant species o Flower only open when bees are active o Pollen collecting apparatus specific for particular plant Sexual mimics Flowers that mimic female bees or wasps Look/smell like females Males attempt to mate, pick up pollen, then fly to another flower and repeat process No reward supplied to male Other insects Butterflies—flower usually has flat landing platform and long, slender floral tubes for long proboscis Moths—flower usually white, yellow, or other pale color, heavily scented, typically need to be found at night Flies—flower usually smells and somewhat appears like feces or rotting meat Beetle—large flowers, copious pollen; beetle may eat other flower parts Birds o Large amounts of nectar o Red—bees can’t see red, less likely to feed on the copious nectar o Usually odorless—birds have a poor sense of smell o Often in long, thick tube Mammals (bats especially)—uncommon, but for some species is the only means of pollination; variety of appearances o In US, saguaro cactus is one of few bat- pollinated species Self-pollination o Small, inconspicuous flowers o Shed pollen directly onto stigma (or falls there by gravity); often before bud opens o Advantageous occasionally because no other plant is needed and no vector is needed—good when pollinators aren’t around (arctic, mountains) o If you are well-adapted, might as well produce clones o Disadvantage of genetic load of bad mutations FLY FLOWER STORIES Ex: Rafflesia of Sumatra Largest single flower on Earth Largest inflorescence is made by fly-pollinated plant, the Corpse- flower PROMOTING OUTCROSSING Staminate and pistallate flowers Dioecious- separate sexes What about plants that are monoecious? –several options still left Dichogamous—stamens and pistils reach maturity at diferent times Stigma and stamens don’t touch (includes heterostyle) Genetic self-incompatibility—pollen tube arrested or never germinates
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