The flower and flowering “The flowers the organ of sexual reproduction, and is usually an hermaphrodite structure, but it may be unisexual with separate male an female flowers on the same plant (monoecious, e.g. Corvus avellana (Hazel), or with separate male and female plants (dioecious, e.g. Mercurialis perennis (Dog's Mercury). ‘The flower develops atthe tip ofa shoot, andthe parts of the flower may be seen as modified leaves. But we are ignorant of the evo~ Iutionary history ofthe flower (lowers are ephemeral structures, arly preserved inthe fossil record) and it sa matter of conject tie whether comparison can be made between the flower of modem angiosperms andthe spore-producing structures of other tracheo= piytes (meaning plats with vascular tissue, e.g cones of Selagnella spp (Club Mos), cones of Pins sylvestris (Scots Pine) see P68) Tlowover the point of transition from vegetative growth to flowering in plant development has been investigated. The external emb= ‘Goture and the chang in day length have proved to be environmental factors of key importance. After a certain stage in vegetative frowth (Le. when a certain numberof nodes has been produced) a condition of ‘ripeness to flower’is achieved, Day-neutal varieties ail then flower, e Lycopersicum esculentum (Tomato). Such plants often continue vegetative and reproductive growth simultan- ously, Other plants then require exposure 1 days of a particular length before they wll ower (photoperiodism), For example, many plants of temperate regions will flower only inthe ong days of summer and are called long-day plants, e.g. Avena sativa (Oats) Other puhtsrequite long dark periods and this group includes many plants indigenous to regions of low latitude north and south of the equator where day length never exceeds more than 14 hour at any seaton ofthe year. An example of «short-day plant i Glycine soja (Soya Bean) Flower may occur solitary ona stem, orn groups (an inflorescence) onthe same mainstem. The parts ofeach flower are bome in ‘whorls on the expanded tip (receptacle) ofthe flower stalk (pedicel) ‘The sepals (collectively called the calyx) are usually green and small. They enclose the flower in the bud. ‘The peal (collectively called the corolla) are often coloured and conspicuous, and may attract insects to the flower. The stamens fandroecium ~ derived from the Greek for ‘man’ and “house’) are the male parts ofthe flower, consisting of anthers (Containing pollen grains) and a flament (stalk) ‘The carpets (qvnoecitum ~ derived from the Greek for ‘woman’ and house") may be one or many, separate or joined, Each consists of anovary (containing ovule/s), stigma (receptive surface for pollen) anda style (connecting ovary and stigma). Figute 4.5) LS of a flower bud of Ranunculus sp. (Buttercup) (x 25) Figure 454i) Haltlower of Ranunculus repens (Buttercup) = ea nie joe Lower down the flower stalk pedicel) i attached & Featlite bract (shown inthe floral agram below) Figute 4.) Floral diagram and floral fo enunculus sp. filament [A androscium {G= pynoecium @KSCSAm Ge, “The tral diagram is rally a plo of tho flower, with the parts ep ‘xonted gs seg in tanevorse section. Ifthe flower parts ae joined opetter thes shown By brackets and lines. Together with the floral ‘Shmule ts a rmeana of showing the structure of the Flower et (th oectary pedis! 54Figure 4.10) Halt flower and tora formula Figure 4.6) TS of flower bud of Litium sp. (x 40) anthers with —— Bolan sacs Lcaoatory, totapat the Berlanth tube ee Fuad ino 8 tube the tower ‘There is similar basic structure to most flowers, but there is enormous variation too. Many dicotyledons have flower parts in fives but the Cruciferae (e.g. Sinapis arvensis (Charlock)) have their parts in fours, and the Poppy family in twos. Ranunculus sp. (Butter. cup) has a very large number of stamens and carpels, but many plants have small and regular numbers of both stamens and carpels, Flower parts are not always symmetrically arranged about any axis as in regular or actinomorphie flowers. The Leguminosae (2 Lupinus polyphyttus (Garden Lupin)) and the Labiatee (e.g. Lamium album (White Deadnettle)) are examples with zygomorphic flowers, having only one plane of symmetry. Also the flower parts may be joined into tubes, as is both calyx and corolla of White Deadnettle. Ifthe sepals and petals ae indistinguishable they are called the perianth, The perianth may be petaloid asin Lilium spp. (Lilies) This is the case in many families of monocotyledons, where flower parts occur in threes. Many flowers have a conical re ceptacle like that of Ranunculus spp.; in others the receptacle is saucer- or flask-shaped, and for some the ovary is ‘inferior’ bing completely embedded in receptacle tissue. The various ways in which flowers are grouped together give rise to different forms of inflorescence, all classified according to whether the youngest flower occurs at the apex or at the base. Probably the least wasteful and therefore most efficient inflorescence is as in the head or eapitulum shown by Compositae, where flowers are clustered tightly together. Figure 4.611) LS of eapitlum of Taraxscum sp. Figure 4.6\iv TS of captulum of Scobios sp. (Seablous) (x50) (Dandelion) 8) coraa tube corolla as tube surrounding Stigma style tnd stamens four anthers with pollen rains Erelosed seraceous teeth frmodited calyx) inferior ovary stom expanded Into platform one ofthe bracts surroundingThe gynoecium ‘The gynoecium (female part of the flower) consists of a carpel or carpels which may be free-standing or fused together in various ‘ways. Most carpels sit upon the receptacle tisue (ovary superior), but in some cases the ovary is embedded in the receptacle (ovary inferior). The evolutionary origin (phylogeny) of the carpel may be from a fertile leaf, the margins of which bore ovules. Figure 4.7) Diagrammatic representation of probable evolutionary development of present-day Lilium (Lily) carpets from a fertile laf of <> 1 Fertile oat seen in sation 2 Folded a lemictiy. 2 Three free-standing carpe 4 Fusion of carpet walls ‘The ovule consists of the mucellus (parenchyma cells) which is surrounded by integuments (when these develop) and attached to the placenta of the ovary by a stalk (the funiculus). Within the nucellus isthe megaspore-mother cel. Figure 4.7) Stage inthe ontogeny ofthe ove ss f ‘ ation of / : ot * ae | } ) x | \ — ae aa ‘of integument parenchyma cells ‘of nucelue cavity of ovary @ ica! 3 ges oe e e ragion of development. OF an integument 56‘The female gamete isthe egg cel, which develops within the embryo sac. The embryo sac develops from the megaspore-mother-cll Numerous recent studies have discovered surprising diversity in the details ofthis process within different angiosperm species. In Fritllaria spp. and Lilium spp. megasporogenesis and megagametogenesis are as shown in the diagram below: Figure 4,8) LS of ovule of Fritileria ep (Friary) showing the mature embryo be (e130) lovle embryosec —nucallus | Ox¢ woos com the developing saeesco OW one haploid z 5 tenis 5 sete Sn) 8 g ae) See =e mea integument \4 5 cal nd four hao ae octet tal — (Sn fra in cal a The original observation of megasporogeness and megagametogenesis in Polygonum spp. (Knotgrass), made by Strasburger in 1879, revealed a syne (n} proces which i outlined in the drawings below. This has often beet e- ported asthe ‘normal type of gamete development inthe gynoecium of Theiosperm. Infact the process is a much more diverse one. For any past- ieular species there may exist important departures from the two examples a aiven here. aig \EGASPOROGENESIS MEGAGAMETOGENESIS ipo mappa 4 5 6 developing 7 ® soy \ “ides 1 2 2 famed © | ° = = and * a e apt @ MEIOSIS e > wirosis MITOSIS. «—_«MITOSIS. se ie) 37‘The androecium and the male gamete ‘The androecium (male part of the flower) consists of, stamens, which are upgrowths from the receptacle. Each stamen consists of an anther, containing four pollen sacs (microsporangia), and a fllament, ‘Anther contains four pollen sos. filament: Each pollen sac has a wall several cell layers thick; the innermost layer, (tapetum) provides food for the developing pollen ‘grains (miérospores). The microspore-mother-cells ‘undergo meiosis, o that each cell produces four pollen grains ina tetrad, Pollen grain formation goes on in a nutritive liquid secreted by the vapetum. Pollen grains ‘may be arranged in various ways in the tetrads, eg: Figure 4.91) TS of part ofthe anther of Lifiumy ep Lily] showing high-power detail ‘of part of a pollen sac microsporangiam) (x 250 calls of wall ayers epidermis of anther ‘ierospore-motner-cl vapetum i) TS through part of Lilium sp. pollen sae, showing @ 1 talophese (x 750) 7As the anther matures a fibrous layer of calls (the endothecium) appears below the epidermis, with U-shaped thickenings in- _epiéermis, gonneetve, with volving all but the outer facing wall of each feos we NC Vee ore 101) TS through the rie anther of Liu gp. (Lily! near the base, ata point where the "nt fused with the connective bx 110) cell. As these cells dry out, the tension created ruptures the thin-walled cells (the stomium) at the junction of the pollen sacs. ‘A longitudinal split exposes the pollen grains, (The majority of anthers dehisce in this way.) ‘The mature pollen grain has « haploid nucleus which divides to form a generative cell and a tube nucleus (see p.60). The pollen grain js surrounded by a thin cellulose wall, the intine, and an outer, usually sculptured wall, the exine. The pollen grain wall has thin areas (apertures) where growth of the pollen tube can occur. Pollen from monocotyledons is often oval shaped with one aperture; that from dicotyledons usually has three apertures. ‘Analysis of the different sculpturings on the surface of pollen grains is of great importance to: 1) understanding the history of floras all over the world ~ by means of quantitative pollen analysis of recent geological layers it is possible to determine the species of plants that existed when no other relic of them has been preserved; (2) the study of allergic diseases — pollen grains are one of the most important causes of allergic disease of the respiratory tract (3) research on honey plants — the pollen collected by bees can be identified. Pollen shows a chemical composition of the following order: proteins, 726%: carbohydrates (starch, later converted to sugar), 24-48%; fats, 114% ash (ie, minerals), 1—S7%; and water 716%, Figure 4.10 (i Pollen gran of Lilium sp. (x 1500) Figure 4.10(i Pollen grain of Gossypium sp. (Cotton) sen by sean slectron microscope (x 1400) ener tube 9