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BIOL 1030 – TOPIC 7 LECTURE NOTES Topic 7: Angiosperms, Flowers and Pollination Syndromes – Phylum Anthophyta (Chs. 30, 38)  Phylum Anthophyta – flowering plants (antho – flower) A. also known as angiosperms (angeion – vessel or enclosure; sperma – seed) B. ovules enclosed within carpel (parent diploid sporophytic tissue) at pollination 1. the “vessel” is the carpel, which is a modified leaf 2. carpels, especially their enlarged basal portion (the ovary), usually develop into fruit, which is unique to angiosperms C. about 250,000 known living species (dominant photosynthetic organisms on land) D. predominant source of human food E. most widespread and diverse plant phylum 1. range from microscopic to plants with leaves over 6 m long 2. flowers show incredible variety from species to species 3. variety of lifestyles includes parasites (ex.: mistletoe, dodder, beechdrops); mycotrophs (derive nutrients from fungi; ex.: Indian Pipe, others); epiphytes (ex.: some orchids); “carnivorous” (ex. pitcher plants, sundews, Venus flytrap) F. monophyletic group with seeds, refined xylem, double fertilization, and these synapomorphic characteristics: 1. seed contains endosperm 2. presence of flowers (modified stems and leaves) 3. true fruits G. evolutionary history 1. monophyletic group 2. origin about 140 MYA H. phylogeny 1. 2. historically divided into two classes, dicots and monocots  recent genetic analysis has shown that the traditional dicots are a paraphyletic group  thus, the old classification scheme is being replaced no conclusive cladogram has been produced for angiosperms  studies are ongoing  most modern cladograms have Amborella and water lilies as a sister group (or groups) to the rest of the angiosperms  cladogram below from http://tolweb.org/tree?group=Angiosperms&contgroup=Spermatopsida  various class-level groupings have been proposed, the overall naming and formal classification within Phylum Anthophyta is still in a state of flux 19 BIOL 1030 – TOPIC 7 LECTURE NOTES  nevertheless, by far most of the living angiosperm species are found within two monophyletic groups, eudicots and monocots 3. 4. eudicots  most have embryos have two cotyledons (seed leaves)  leaves have netlike veins  flower part typically in multiples of 4 or 5  groups of vascular tissues in a ring  pollen grains mostly with 3 or more apertures  endosperm mostly used up in mature eudicot seeds  about 175,000 living species; includes nearly all flowering trees and shrubs  about a sixth are annuals (entire growth cycle in one year or less) monocots  embryos have one cotyledon  leaves have essentially parallel veins  flower part typically in multiples of 3  groups of vascular tissues scattered  pollen grains mostly with one aperture  endosperm typically present in mature monocot seeds  about 65,000 living species; no true wood, few annuals  Why were (and are) angiosperms successful? A. 130 MYA two major continental masses 1. Laurasia = North America, Europe, Asia 2. Gondwanaland = South America, Africa, Australia, Antarctica, India, New Zealand) B. angiosperms first appeared in Gondwanaland, in what was likely a drier interior region C. advantages of flowering plants 1. transfer of pollen over great distances promotes outcrossing 2. efficient seed dispersal via fruit 3. endosperm gives seedlings a fast start 4. leaves appropriate for fast growth in hot, dry environment D. coevolution with insects 1. dominant by ~80 MYA, second half of Cretaceous Period 2. all present angiosperm families represented by that time 3. many insect orders appeared or became more abundant at that time  Flowers A. modified stems with modified leaves B. develop as primordium bud at end of stalk called pedicel C. pedicel widens at tip to form receptacle 29 BIOL 1030 – TOPIC 7 LECTURE NOTES D. other flower parts attached to receptacle in four whorls; from outside in: 1. calyx – sepals; usually green, leaf-like, and protect immature flower 2. corolla – petals; usually colorful, attract pollinators; together with calyx called perianth 3. androecium – stamens; male reproductive structures 4.  filament + anther  microspores produced within anther, shed as pollen gynoecium – female reproductive structure  center location is most protected  formed from leaf-like structure with ovules along margin  edges fold inwards around ovules, forming carpels    primitive: many separate carpels  advanced: carpels fused (called pistil) carpel/pistil segments  ovary – swollen base with 1 to hundreds of ovules; develops into fruit  stigma – tip; sticky and/or feathery to catch pollen  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 E. know the structures of a flower [Figure 38.2] and their functions  typical Angiosperm life cycle A. female gametophyte 1. single diploid megaspore mother cell in ovule undergoes meiosis while flower develops 2. of 4 haploid megaspores produced, usually 3 break down 3. remaining megaspore expands and replicates and divides until there are 8 haploid nuclei in two groups of 4 39 BIOL 1030 – TOPIC 7 LECTURE NOTES 4. one nuclei from each group migrates toward center; these are polar nuclei 5. polar nuclei usually fuse to make a diploid nucleus, but may remain separate – in either case, they wind up in a single cell 6. cell walls form around other nuclei, creating the 7-celled, 8-nucleate embryo sac or megagametophyte (female gametophyte) 7. meanwhile, two layers (integuments) of ovule develop into seed coat with micropyle (small opening) 8. in the megagametophyte, one of the cells closest to the micropyle becomes the egg; the other two there are synergids 9. the three cells on the other end (the antipodals) eventually break down B. male gametophyte 1. anthers with patches of tissue that become chambers lined with nutritive cells 2. each patch has many diploid microspore mother cells 3. microspore mother cell undergoes meiosis, making 4 haploid microspores that typically remain grouped in a tetrad 4. each microspore nucleus replicates and divides once (via mitosis) without cytokinesis (meaning they remain as one cell with two nuclei, a binucleate microspore) 5. usually, tetrad then breaks up 6. two-layered wall develops around each binucleate microspore, now called a pollen grain  outer wall – sculptured, appearance usually species-specific, often has chemicals that can react with an appropriate stigma to stimulate pollen tube formation  apertures in outer wall – where pollen tube may grow out; eudicots – usually 3; monocots – usually 1 C. pollination – transfer of pollen to a stigma 1. usually between flowers of separate plants 2. agents include wind, water, gravity, mammals, birds, insects 3. various reward systems for animal agents (pollen, nectar, etc.) 4. evolution of floral characteristics associated with pollination 5. some plants self-pollinate (inbreeding) – pollen to same plant 6. pollination followed by fertilization only if chemical signals are right D. fertilization 1. pollen grain cytoplasm absorbs substances from stigma 2. bulge forms through an aperture in pollen grain; becomes pollen tube 3. pollen tube follows chemical gradient through style to micropyle  chemicals diffuse from embryo sac  micropyle usually reached within a few days (up to a year in some species) 4. pollen grain has two nuclei; one, the generative nucleus, lags behind 5. generative nucleus undergoes mitosis to make two non-flagellated sperm; this may occur in pollen grain or in pollen tube (male gametophyte now mature) 6. pollen tube enters embryo sac, destroying a synergid 7. 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 49 BIOL 1030 – TOPIC 7 LECTURE NOTES  endosperm provides nutrients for embryo; in many seeds, it is gone by the time the seed is mature  seed coat hardens  remaining haploid cells degenerate  now have seed with 2N embryo, 3N endosperm, and 2N seed coat (seed coat from parent female tissue) 59 BIOL 1030 – TOPIC 7 LECTURE NOTES  Seeds A. embryo – quickly forms all systems, then growth arrested (dormancy) – mature seed about 10% water, very low metabolic activity B. typically, dormancy occurs just after first leaves (cotyledons, or seed leaves) form C. stored food (in angiosperms, 3N endosperm and/or cotyledons) D. seed coat – tough, relatively impermeable 1. protection from predators, pathogens 2. protection from desiccation, harsh conditions (crucial on land) 3. may allow seed to last hundreds of years E. dormancy broken only when conditions are right (seed bank in soil) F. germination = breaking dormancy = resuming metabolic activity, growing out of seed coat; occurs after water penetrates seed coat to embryo, bringing oxygen  Fruits – mature ovaries A. fleshy – pomes (apples), drupes (peaches), true berries (blueberries, peppers), hesperidiums (oranges), pepos (melons, gourds), aggregate fruits (strawberries, raspberries), multiple fruits (pineapple, fig) B. 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) C. dispersal 1. by wind  wings – maples  parachutes – dandelions, milkweeds  dust-like seeds – orchids 2. by water – coconuts 3. by vertebrates 4.  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.) by explosive dehiscence (jewelweed, others) 69 BIOL 1030 – TOPIC 7 LECTURE NOTES  many angiosperms also have asexual (or vegetative) reproduction A. stolons – runners – long slender stems that grow along soil (ex.: strawberry) B. rhizomes – underground stems – common in grasses; bulbs and tubers are rhizomes specialized for storage (ex.: potato) C. suckers – roots produce sprouts that grow into new plants (ex.: apple, raspberry, banana) D. adventitious leaves – numerous plantlets develop from tissue in notches along leaves E. apomixes – embryos in seeds may be produced asexually F. artificial: cuttings (for some species, can get roots to grow with appropriate environment)  Floral Evolution A. first flowers 1. numerous spirally arranged sepals, petals, stamens, and carpels 2. petals and sepals similar in color and form 3. all parts free (not fused) B. parts 1. complete – calyx + corolla + androecium + gynoecium 2. incomplete – one or more whorls absent 3. perfect – has both androecium and gynoecium 4. imperfect – missing either androecium or gynoecium 5. complete flowers are always perfect; incomplete flowers can be either perfect or imperfect C. trends 1. separate floral parts grouped together or fused  connation – fusing within a whorl  adnation – fusing between whorls (for example, sepals and petals fused together) 2. reduction or loss of floral parts 3. bilateral symmetry instead of radial symmetry  ancestral type: radial symmetry; example: buttercups  derived type: bilateral symmetry; examples: snapdragons, orchids  bilateral symmetry in some cases has arisen independently in different groups  Pollination mechanisms (pollination syndromes) A. wind – passive, primitive (oaks, cottonwoods, birches, grasses) 1. copious amounts of pollen 2. most pollen travels no more than 100 m 3. flowers small, greenish, odorless 4. corollas reduced or absent 5. often grouped in large numbers, may hang down with tassels that wave in wind and shed pollen freely 6. male and female parts often well-separated on plant to reduce chance of self-pollination 7. often flower before leaves grow – keeps leaves out of the way B. animals – some cycads and gnetophytes also have this, so symplesiomorphic trait 79 BIOL 1030 – TOPIC 7 LECTURE NOTES 1. 2. 3. 4. 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 to indicate nectaries (nectar guides)  nectar offered as food for bees (pollen also)  often close association between a bee species and a plant species  flower only open when bees are active  pollen collecting apparatus specific for particular plant 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  large amounts of nectar  red – bees can’t see red, less likely to feed on the copious nectar  usually odorless – birds have a poor sense of smell  often in long, thick tube mammals (bats especially) – uncommon, but for some species is the only means of pollination; variety of appearances C. self-pollination 1. small, inconspicuous flowers 2. shed pollen directly onto stigma (or falls there by gravity); often before bud opens 3. advantageous occasionally because no other plant is needed and no vector is needed – good when pollinators aren’t around (Artic, mountains) 4. if you are well-adapted, might as well produce clones 5. disadvantage of genetic load of bad mutations  Promoting outcrossing A. staminate and pistillate flowers B. dioecious – separate sexes C. monoecious 1. dichogamous – stamens and pistils reach maturity at different times 2. stigma and stamens don’t touch (includes heterostyle) 3. genetic self-incompatibility – pollen tube arrested or never germinates 89