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Lecture 3 – Floral Morphology 2 major phylogenetic/ developmental theories on origin of flower • Euanthial hypothesis (Arber & Parkin 1907): Flower is derived from a uniaxial cone (cone with one axis) i.e., a central axis with spirally attached fertile & sterile modified leaves (sporophylls & bracts Ø Angiosperm flower is equivalent to simple gymnosperm strobilus (or cone), as in Cycads and the extinct Bennettitales Ø superficially strobilar flowers (e.g. Magnoliales) viewed as primitive – Large, radially symmetrical, numerous free & spirally arranged flower parts of foliar origin (incl. laminar stamens, multiovular leaf-like carpels), large seeds • Pseudanthial hypothesis (Von Wettstein 1901; 1907): Flower derived from pluriaxial cone (cone with multiple axes) i.e., a central axis with numerous secondary axes, each bearing fertile or sterile parts Ø Angiosperm flower is equivalent to cones of conifers, Gnetales & the extinct Cordaitales Ø Current uniaxial angiosperm flower derived by aggregation and condensation of a gymnosperm “inflorescence” Ø Primitive flower is a compilation of small, simple, unisexual units on several orders of axes, with a whorl of bract-like structures (“perianth”) at base. Bracts on other axes now lost. Small simple carpels with one to a few ovules Ø Piperales and “amentiferous” taxa (incl. Fagales) would thus be primitive. Ø Latter is contradicted by fossil record REVIEW OF BASIC TERMINOLOGY – FLORAL WHORLS – Sepals (outer perianth whorl) à Petals (inner perianth whorl) à …. Tepals (perianth parts look alike) Stamens Carpels Collective names CALYX COROLLA ANDROECIUM GYNOECIUM Perianth = Floral envelope (all sepals, petals or tepals) Carpel: a simple pistil formed by one modified leaf (megasporophyll) … or that part of a compound pistil formed by one modified leaf. PISTIL = stigma, style plus ovary… megasporphyll STAMEN = Filament plus anthers… microsporophyll Locule: The chamber or cavity of an ovary containing the ovules Nectary: glandular tissue on the flower (or elsewhere) producing nectary Pedicel: Stalk of a single flower in an inflorescence Peduncle: Stalk of a solitary flower of the entire inflorescence Receptacle: Portion of the pedicel on which the flower parts are borne [In composites (Asteraceae) refers to the part of the peduncle where the flowers of the head are borne] ________________ Floral Diagram (e.g. Judd et al. p. 193) -- see handout Floral Formula – see Judd et al. page 57 NUMBERS OF FLORAL PARTS COMPLETE – With all the parts belonging to a typical flower (petals, sepals – or tepals – stamens, pistil) INCOMPLETE – Lacking one of the whorls ASEPALOUS – lacking sepals APETALOUS – lacking petals IMPERFECT – Unisexual flowers (pistillate or staminate) Imperfect flowers are also incomplete, by definition! PERFECT flowers are bisexual à Other aspects of plant “gender” • A large and complex vocabulary has grown up around the expression and timing of plant sexuality (gender). • A plant can be hermaphroditic in many ways, including possessing only perfect flowers Gender terms to know: MONOECIOUS (noun: monoecy) – “One house”. Hermaphroditic plants with imperfect flowers, staminate and pistillate, on the same individual DIOECIOUS (noun: dioecy) – “Two houses”. Each individual possesses only one kind of imperfect flower, staminate or pistillate. Humans and most animals are “dioecious”. Most angiosperms are not! HERKOGAMY: “Male” and “female” sex organs are spatially separated in a bisexual flower. Ø Heterostylous species have a special form of this, “reciprocal herkogamy” DICHOGAMY: “Male” and “female” organs functional at different points in time: Ø Protandry (anthers open first) Ø Protogyny (stigma receptive first) • Dioecy, herkogamy & dichogamy function to prevent self-fertilization and may promote cross-pollination Back to basic morphology… FUSION CONNATE (adj.) – FUSION OF LIKE PARTS e.g. the fusion of staminal filaments into a tube SYNANDROUS SYNCARPELOUS SYNSEPALOUS SYMPETALOUS (SYNTEPALOUS) ADNATE (adj.) – FUSION OF UNLIKE PARTS i.e. fusion involving parts from different whorls. Stamens “adnate” to the corolla are EPIPETALOUS. Various terms exist for the fusion structures (will cover later in term) FLORAL SYMMETRY ACTINOMORPHIC – Radially symmetrical. Multiple lines can be drawn through the middle of the flower to yield a mirror image on either side ZYGOMORPHIC – Bilaterally symmetrical. Only one possible line can be drawn through the middle of the flower to produce a mirror image on either side IRREGULAR – No axis of symmetry STAMEN TERMS STAMINODE – sterile organ derived from a stamen ANTHER ARRANGEMENT – Antepetalous: Stamens described as being “opposite” the petals, meaning each stamen sits right over the middle of a petal. Antesepalous: stamens “opposite” the sepals. Diplostemonous: 2 series of stamens, the outer opposite the sepals, the inner opposite the petals Will talk about other types of stamen arrangement when we cover the families ANTHER ATTACHMENT – Basifixed – anther attached at the base Dorsifixed – anther attached at the back ANTHER DEHISCENCE – Longitudinal – along axis of the anther Transverse – at right angles to long axis of the anther Poricidal – through a pore Valvular – through flap-covered pores CARPAL FUSION APOCARPOUS – a flower with carpels forming separate pistils SYNCARPOUS – with ovaries united to each other OVARY POSITION SUPERIOR – Ovary is not united to any other flower part INFERIOR – Ovary is united with the hypanthium, appears to be sunken “below” the other floral organs Hypanthium – fusion product of the calyx, corolla and androecium (may also include receptacle material). Forms a “floral cup” that may or may not be fused to the ovary FLOWER TYPES Three basic flower types are recognized, depending on: (1) (2) Whether the ovary is superior or inferior, If a hypanthium occurs, whether it has become evolutionarily fused to the carpal à HYPOGYNOUS – superior ovary, no hypanthium. The other flower parts arise below (“hypo”) the ovary PERIGYNOUS – superior ovary, hypanthium present and unfused The other flower parts appear to arise around (“peri”) the ovary on the rim of a hypanthium. Hypanthium may be tubular, bowl or saucer-shaped EPIGYNOUS – inferior ovary. The hypanthium may not be readily recognizable because it is fused to the ovary. The other flower parts appear to arise from or on (“epi”) the top of the ovary OVULE PLACENTATION The arrangement or configuration of the placentas (the part that attaches each ovule to the carpal wall) Axile Parietal Free-central Basal Apical TYPES OF BLOSSOM Several ways of classifying pollination units (flowers or aggregated groups of flowers) (1) Mode of pollination Ø BIOTIC, e.g., insect-pollinated (entomophilous), Ø ABIOTIC: wind or water (2) Functional structure (see Faegri and van der Pijl 1979) (a) (b) Ø Ø Ø Ø Flowers inconspicuous, no optical attraction Flowers conspicuous Dish/ bowl e.g., Magnolia, Caltha, Rosa. “Mess-and-spoil pollination” Bell/ funnel e.g., Campanula Brush Gullet. e.g. many Lamiaceae and Scropulariaceae. Flower has upper and lower division. Sex organs on upper side, pollen deposited on insect’s back Ø Flag. e.g. many Fabaceae. Flower has upper and lower division. Sex organs on lower side, pollen deposited on insect’s ventral side Ø Tube e.g., Delphinium, Aconitum. Nectar is only accessible by insertion of insect’s mouth parts (tube may be the entire corolla or a “spur” containing nectar NOTE: Not all flowers fit easily into the functional system! (many orchids, gingers)