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Flowers and Their Evolution
Spring 2014
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Flower = a short, determinate shoot bearing highly
modified leaves, some of which are fertile (i.e., bearing either microsporangia or megasporangia), with the microsporangia in stamens and the megasporangia in carpels.
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Flower REPRODUCTIVE STRUCTURE – Evolutionary requirement to reproduce by sexual means. Pollen transfer and seed dispersal needed. MODIFIED FOLIAR APPENDAGES – all function together to form the reproductive organ known as the FLOWER. MODIFICATIONS OF LEAVES – All floral organs are modified LEAVES. Four terminal WHORLS of modified leaves: - Two outermost whorls (sepals, petals) are sterile (non- meiotic tissues) - Two innermost whorls (sporophylls) are “fertile” with tissues capable of undergoing meiosis SPOROPHYLLS – those modified leaves with meiotic capacity. - Microsporophylls – stamens – produce pollen in anthers - Megasporophylls – carpels – produce eggs in ovules
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Fig. 6.2 from Simpson
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Floral Whorls Attached to RECEPTACLE Sepals (collectively the Calyx)
Petals (collectively the Corolla) Stamens (anthers + filaments) collectively the Androecium (andros = male; -oecium = house) “Pistil” – carpel(s) [fused or not] collectively the Gynoecium (gynos = female; -oecium = house)
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Floral Parts: Major whorls
pistil (1-many carpels) - gynoecium stamens - androecium petals - corolla sepals - calyx receptacle
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Sepals and petals are relatively leaf-like. sepal young leaves XS of
stamen pistil Sepals and petals are relatively leaf-like. sepal young leaves XS of flower bud petal
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“ABC” Model of Floral development Fig. 6.5 from Simpson
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Floral Anatomy
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Evolution of the Androecium
DERIVED FROM MODIFIED LEAVES - Microsporangia (meiosis microspores pollen grains) on lamina originally INCREASING LEVELS OF REDUCTION - Lamina becomes filament - Sporangial tissue becomes anther wall - Provides for release of pollen CAN BE IN A SINGLE WHORL OR MULTIPLE WHORLS - Tremendous variation in flowering plants. - Often associated with specific type of pollinator.
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Stamen evolution microsporangia laminar stamens
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Fig. 9.26 Fig. 9.25
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Floral Anatomy
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Evolution of the Carpel
MODIFICATION OF MEGASPOROPHYLL - Evolution of megasporophyll structure traced back to seed ferns – 200 to 300 mybp LEAF WITH MARGINAL MEIOTIC ZONES FOLDS - Ovules located at margins of sporophylls - Lamina curves inward (toward the floral axis - adaxially) - Carpel is formed by folding – conduplicate - Margins fuse, enclosing ovules - Carpel(s) = gynoecium FUSION OF CARPELS - Unfused (separate) carpels - apocarpous - Fused (united) carpels - syncarpous POSITION OF THE GYNOECIUM relative to other floral whorls is important in describing floral structures. PLACEMENT OF OVULES (placentation) within the gynoecium is also important; shows evolutionary origins of the carpel.
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integumented megasporangium
The Ovule = integumented megasporangium sporangium female gametophyte (derived from a single spore) integuments micropyle
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Carpel evolution (Ovules) (megasporophyll)
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Folding of one megasporophyll S = suture; formed by fusion of leaf
Folding of megasporophyll to form simple carpel Folding of one megasporophyll S = suture; formed by fusion of leaf margins; receptive to pollen receptacle
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Carpel evolution stigma stigmatic crest
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3 pistils 1 pistil Fig from Simpson
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Simple Carpel – One Pistil
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Apocarpy – Separate Carpels = 5 pistils in this example
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Apocarpous gynoecium – Ranunculus sp.
with many pistils elongated receptacle
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Liriodendron Magnolia
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Simple vs. compound ovary
Fig from Simpson
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Syncarpous gynoecium – One pistil, 3 carpels
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Various gynoecia – Apocarpous vs. Syncarpous
(Hint: stigma number usually = carpel number)
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Syncarpy – How many carpels? Locules?
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Adnation: Fusion of different whorls
Stamens (filaments) adnate to petals, petals adnate to sepals
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Connation: Fusion of parts from the same whorl
Fusion of filaments into a staminal tube
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Ovary position relative to other parts
The hypanthium (floral cup) requires both connation and adnation. Fig. 9.32 from Simpson
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Ovary superior Parts hypogynous Citrus sp.
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Ovary superior, parts perigynous
(floral cup or tube = hypanthium present) Rosa sp.
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Ovary inferior, parts epiperigynous (hypanthium present) Fuchsia sp.
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Ovary inferior, parts epigynous Vaccinium sp.
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Ovules and Placentation
OVULES CONTAIN THE MEGAGAMETOPHYTE - Provides for fertilization of egg cell in megagametophyte and protection during development. - Ovule matures into the SEED. ATTACHMENT OF THE OVULES VIA FUNICULUS - Analogous to the mammalian “umbilical cord” - Point of attachment on inner ovary wall is the PLACENTA - Can vary depending on type of flower. PLACENTATION IS OFTEN DIAGNOSTIC - Plant families typically have one placentation type. - Often best seen with cross section through ovary. PLACENTATION REFLECTS EVOLUTIONARY DEVELOPMENT - Fusion of carpels, presence of vascular bundles, etc. can support hypotheses about evolution of particular flower structures.
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Fig. 9.33 from Simpson
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Fig. 9.33, Part A only
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Placentation Axile Parietal
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Floral Symmetry Radial Bilateral Actinomorphic Zygomorphic
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Merosity = basic number of parts
in each whorl -3 sepals, 3 petals, 6 stamens, 3 carpels = 3-merous (or trimerous) -4 sepals, 4 petals, 6 or 8 stamens, 2 or 4 carpels = 4-merous (or tetramerous) -5 sepals, 5 petals, 5 or 10 stamens, 3 or 5 carpels = 5-merous (or pentamerous)
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Interpretation of Floral Structures
OBSERVE STRUCTURES IN EACH WHORL - How many whorls are there? - How many parts are present in the calyx? Corolla? - Describe the androecium, then the gynoecium. DETERMINE POSITION OF THE FLOWER PARTS RELATIVE TO THE OVARY - Hypogynous? Perigynous? Epigynous? Epiperigynous? GYNOECIUM - Apocarpous? Syncarpous? If so, how many carpels? - Position? Superior or inferior or half-inferior? - Placentation? ADNATION or CONNATION? - Fusion of floral parts can sometimes be diagnostic. UNUSUAL OR REMARKABLE FLORAL STRUCTURES? - Specializations for pollination?
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