Topic 1 Flowers Botany Reference Guide CII 106746951 (3 March 2016) Page 1 1.1 Flower shapes and inflorescences This section deals with the various shapes of flowers and the way these flowers can be arranged as a group to form an inflorescence. When a flower shape is discussed, it is the shape of the petals and/or sepals (the perianth) which constitutes a flower shape. However, because there are also many flowers which do not fit exactly into any one of the flower shapes, it is quite acceptable to combine more than one shape name to describe a specific flower shape. If the flower does not have distinct sepals and petals (perhaps only one whorl) then it is the shape of that petal like structure that is described. Figure 1 - Principal organs of a ‘typical’ flower. Flower shapes As you are now aware, the arrangement and shape of the perianth segments (usually the petals) within the one flower structure gives rise to a range of different shapes. Knowing the flower shapes makes identification of some plant species easier and can be used to classify plants into plant families. For example, all flowers which are in family Fabaceae, sub - fam. Faboideae, have a papilionate flower shape. Again, labiate flowers are usually in family Lamiaceae. The stalk of a single flower is known as the pedicel. Botany Reference Guide CII 106746951 (3 March 2016) Page 2 Tubulate Shaped like a tube, the sides are parallel or nearly so, eg Cuphea ignea (cigar flower). Salverform Having a slender cylindrical lower part and opening out flat at the mouth, eg Phlox spp, Plumbago spp. The sides of the lower part of floral tube are nearly parallel. Funnelform Funnel - shaped; petal lobes may turn back at the top, but not as flat as salverform flowers. The sides of the floral tubes are not parallel gradually widening from the base to the top of the flower, eg Petunia x hybrida, Hibiscus rosa - sinensis. Campanulate Bell - shaped; base of flower is more rounded than funnelform flowers. Flowers normally hang down, eg Abutilon x hybridum (Chinese lantern). Botany Reference Guide CII 106746951 (3 March 2016) Page 3 Rotate Wheel - shaped with a short tube at base and spreading at the mouth; petals free from each other, eg Hibbertia spp. (guinea flower), Tibouchina spp, Lagerstroemia indica. Stellate Star - shaped, petals usually free from each other, usually 4, 5 or 6 petals, eg Hemerocallis spp. (day lily), Trachelospermum jasminoides. Labiate Having one or more petals forming a lip, eg Prostanthera spp, Salvia splendens. Also bilabiate – having two lips, eg Westringia fruticosa. Flowers with labiate or bilabiate shape are typical of members of family Lamiaceae. Saccate Irregularly shaped flower with lowest petal spurred or modified into a rounded sac. Corolla tube often with a sac - like bulge on one side, eg Grevillea spp., Nemanthus spp. Calceolate Shaped like the toe of a slipper, eg Paphiopedilum spp. (slipper orchid), Calceolaria. Flower with large bulge on one side. Botany Reference Guide CII 106746951 (3 March 2016) Page 4 Papilionate Shaped like the wings of a butterfly, eg Lathyrus (sweet pea), Wisteria; made up of three distinct petal types: 1. standard 2. wings 3. keel. Typical of members of family Fabaceae, sub - fam. Faboideae. Crateriform Flowers shaped like a shallow open bowl, saucer, or crater of an extinct volcano, eg Papaver spp. (poppy), roses (single). Cyathiform Cup - shaped, and open at the top; more curved at the bottom than funnelform and deeper than crateriform; petals not curving out as in campanulate flowers, eg Cobaea scandens (cup and saucer vine), Tulipa spp. Urceolate Urn - shaped, narrow at the mouth and wider below, similar to cyathiform but nipped in towards the top; these flowers are often quite small, eg Erica carnea (spring heath), Epacaris spp, Arbutus unedo, Pieris japonica spp. Radiate Petals spreading like many rays from a ‘centre’ of reproductive structures; more petals present than rotate or stellate flowers (eg Mesembryanthemum (pigface) Lampranthus). This term is often incorrectly used to describe the inflorescence of members of family Asteraceae. Botany Reference Guide CII 106746951 (3 March 2016) Page 5 Having showy stamens With these types of flowers, we are describing what we see (which are the stamens) rather than the shape of the perianth. It does not mean that these flowers do not have all the parts of a ‘typical’ flower, it is just that it is the stamens which are obvious and showy. There are two types of flowers with showy stamens. Conspicuously staminate Flwers with many conspicuous, often brightly coloured and showy stamens; sepals and petals present but may be small and inconspicuous (eg Melaleuca, Callistemon). Apetalous staminate Fowers with many conspicuous, often brightly coloured and showy stamens; flowers have no petals when the stamens are mature (eg Eucalyptus). The petals have been modified into a protective structure which is shed as the stamens expand and the flower matures. Inflorescences An inflorescence is defined as a group of flowers borne on one stalk or stem. This inflorescence stalk is called the peduncle. The arrangement of individual flowers on a peduncle gives rise to several different inflorescences. Some inflorescences can be very difficult to determine—in fact, some may be a combination of two inflorescence types. For example, the inflorescence of Lantana species is an umbellate spike. You may find it easier to identify some inflorescences by studying them before all the flowers are opened, or if the flowers have opened, shake the inflorescence gently and turn it upside down. You must always locate the flowers that have opened first; that is, the oldest flowers in the inflorescence. This will help to determine the inflorescence type. You will note that the individual flowers on some inflorescence diagrams are numbered. Number 1 = oldest flower, or flower formed and opened first, 2 = second formed and opening flower, 3 = third formed and opening flower—and so on to the youngest flower on the inflorescence. Botany Reference Guide CII 106746951 (3 March 2016) Page 6 Types of inflorescence There are three main types of inflorescence: solitary raceme - type, or indeterminate cyme - type, or determinate. Solitary Solitary flower borne singly on a pedicel. They may be formed as a terminal flower or singly in the leaf axils up a stem, eg Papaver nudicaule (poppy), Westringa fruticosa. Raceme type or indeterminate inflorescences These are all characterised by having the oldest flower opening at the base of the inflorescence while growth is still continuing longitudinally. Raceme A group of flowers attached to the peduncle with pedicels of about the same length; the oldest flower is at the bottom of the inflorescence and the youngest at the top, eg Salvia splendens. Spike A racemose inflorescence with sessile flowers (without a pedicel), eg Callistemon spp. (bottlebrush). Botany Reference Guide CII 106746951 (3 March 2016) Page 7 Panicle A branched racemose inflorescence with each whole branch being a raceme. The side branches are smaller racemes. A panicle is a compound raceme, eg Yucca spp. Corymb A racemose inflorescence in which all the flowers are ultimately borne at the same level because the pedicels are of uneven lengths. The flowers towards the bottom of the peduncle (that is, the oldest flowers) have the longest pedicels, eg Iberis spp. (candytuft), Spiraea spp. Simple umbel A racemose inflorescence in which all the pedicels are equal in length and arise at one point on the peduncle. The flowers are at the same level, eg Hippeastrum. Compound umbel An inflorescence made up of small umbels arranged in a larger umbel, the peduncles are equal in length, eg fennel, parsley. Botany Reference Guide CII 106746951 (3 March 2016) Page 8 Catkin A pendulous racemose inflorescence modified for wind pollination. It is a loose spike make up of numerous sessile, usually unisexual flowers. Female catkins usually have long hairy styles and stigmas to enhance pollen interception. Capitulum (Head) A racemose inflorescence with sessile flowers on a flattened and expanded peduncle, eg Zinnia elegans. Each inflorescence consists of two floret types ray florets: a ligulate floret around the edge of a capitulum, has a conspicuous ‘strap’; disc florets: tubular florets in the centre of a capitulum. Botany Reference Guide CII 106746951 (3 March 2016) Page 9 Cyme type or determinate inflorescences These are characterised by having an inflorescence in which the terminal flower terminates the growth of that inflorescence and other flowers arise laterally below it. The oldest flowers are nearest the apex, eg Ranunculus spp. (buttercup), Tibouchina spp. When the flower arise both sides below the terminal flower sides the inflorescence is a dichasial cyme. When the flowers arise from one side only below the terminal flower, the inflorescence is a monochasialcyme. Botany Reference Guide CII 106746951 (3 March 2016) Page 10 1.2 The ‘typical’ flower Flowers come in all shapes, sizes and colours, yet at the same time they have some features in common. Figure 2 - A ‘typical’ flower Arrangement of floral parts Floral parts are inserted on the receptacle of each flower. They can either be arranged in whorls or spirally on the receptacle. Whorls In most flowers the calyx, corolla, androecium and gynoecium are arranged in circles or whorls on the axis of the flower, eg rose, Petunia spp, Hibiscus spp. Spirals Some flowers have the calyx, corolla, androecium and gynoecium attached at different levels in an ascending spiral, eg Magnolia spp, Ranunculus spp. Botany Reference Guide CII 106746951 (3 March 2016) Page 11 Functions of floral parts If the flower part is not directly involved with the reproduction process then it is referred to as a non - essential part of the flower. The essential parts of a flower are those concerned directly with the reproduction process. They include both the essential male parts of these flower and the essential female parts of the flower. Non - essential flower parts Calyx If you examine a flower from the base, the first part you see is a group of small leaf - like structures arranged in a circle. Each one of these is called a sepal and the whole group of sepals is called the calyx. Before a flower bud opens, it is enclosed and protected by the calyx. The calyx then opens out to let the other parts of the flower emerge. Corolla The next whorl in from the calyx you can see is a group of (usually) coloured petals. Each petal may have different shapes. A group of petals is known as the corolla. Perianth This word can be used in the following ways: as a collective term for the calyx and corolla; if either the calyx or corolla is absent, the remaining whorl is called the perianth; if there are two whorls present but both whorls are similar in shape, size, texture and/or colour they are both referred to as the perianth. Botany Reference Guide CII 106746951 (3 March 2016) Page 12 Receptacle (or torus) The tip of a stalk from which the flower parts arise, the shape may vary: from convex, to flattened or concave. Where the receptacle is concave, the ‘arms’ which extend out from the receptacle and hold the sepals and petals is called the hypanthium. Pedicel This supports the flower and is attached to the stem. Peduncle This is the stalk of a whole inflorescence. Botany Reference Guide CII 106746951 (3 March 2016) Page 13 Essential flower parts - male The collective term for the total of all stamens in the one flower is the Androecium. Stamen This consists of the: anther, which produces pollen grains containing the male reproductive cells; filament, the stalk which support the anther. This figure shows the possible types of stamen structures and variations: (a) stamen structure; (b–c) anther dehiscence; (d) staminodes. Botany Reference Guide CII 106746951 (3 March 2016) Page 14 Essential flower parts - female This is the collective term for the female parts of the flower is the Gynoecium. It may be composed of one or more pistils. Where there is one loculus only and therefore one carpel, the whole structure (ie stigma, style and ovary) may be referred to as a carpel. Pistil This is a structure composed of a stigma, style and ovary. Stigma: the receptive tip of the pistil which receives pollen and on which the pollen grain germinates. Style: the stalk - like structure which supports the stigma. The germinating pollen grain tube which develops and grows down through the style to the ovary. Ovary: the swollen base of the pistil which contains the ovules. Inside the ovary are found: ovules: the structure containing the female reproductive cell, called the egg or ovum; loculus: the cavity or chamber within the ovary where the ovules are located; placenta: the tissue within the ovary to which the ovules are attached; carpel: the region of the ovary including the carpel wall, loculus, placenta(s) and ovule(s). An ovary may be composed of a single carpel or two or more fused carpels. Botany Reference Guide CII 106746951 (3 March 2016) Page 15 1.3 How do flowers differ from each other If you look at the many different kinds of flowers, you will notice that their structure varies. The sepals, petals, stamens and pistils may or may not be present, and may be arranged in a variety of different ways, or the flowers themselves may be arranged in different positions on the plant. Presence or absence of flower parts Complete flower This is a flower with all parts present, that is the calyx, corolla, androecium and gynoecium present (eg Petunia). Incomplete flower This is a flower in which one or more of the parts is missing (eg spinach has a flower with no corolla). Perfect flower This is a flower with both male and female parts present (eg Petunia); also called bisexual. Imperfect flower This is a flower with only female or male parts present. Imperfect flowers are divided into two categories: Staminate flowers have only the stamens present (eg male pawpaw flowers). Botany Reference Guide CII 106746951 (3 March 2016) Page 16 Pistillate or carpellate flowers have only the pistils present (eg female pawpaw flower). Monoecious plant This has both staminate and pistillate flowers on the same plant (eg maize, cucumber). Dioecious plant This has staminate and pistillate flowers on separate plants (eg avocado, chinese gooseberry). Union of flower parts In a flower, members of one whorl may be fused to each other or to members of another whorl. The following terms describe some of the conditions which can occur. Apopetalous The petals are free from each other and other parts of the flower (eg Boronia). Connate Members of a single whorl are joined or fused together (eg connate petals in Petunia). Botany Reference Guide CII 106746951 (3 March 2016) Page 17 Sympetalous Sympetaly is a special case of connation where the petals are attached to one another (eg Narcissus). Synstemonous Synstemony is a special case of connation where the stamens are joined together (eg Hibiscus). Adnate Members of two different whorls are attached to each other. Most often adnation occurs between stamen and petal or petals and sepals (eg in the snapdragon flower, the filament of each stamen is attached to the corolla rather than to the receptacle). Epipetalous Epipetaly is a special case of adnation where the stamens are attached to the corolla (eg Petunia). Symmetry of flowers Zygomorphic (irregular flower) A flower which is irregular (or zygomorphic) is one in which the perianth can be divided into two similar halves through one plane only (eg sweet pea). Botany Reference Guide CII 106746951 (3 March 2016) Page 18 Actinomorphic (regular flower) In a flower which is described as being regular (or actinomorphic) the perianth can be divided into two similar halves through a number of planes. This characteristic is also described as radial symmetry (eg Petunia). Position of the ovary The position of the ovary is an important point to consider if you are trying to identify a particular plant. There are three main ovary positions, although within these there are numerous variations within each group. These main positions are: Superior with floral parts either hypogynous or perigynous. Botany Reference Guide CII 106746951 (3 March 2016) Page 19 Inferior with floral parts epigynous. Half inferior with floral parts perigynous. Superior ovary with hypogynous floral parts This is where the point of attachment of the ovary to the receptacle is at the base of the ovary only. The sepals, petals and stamens arise from (or are attached) beneath the ovary on the receptacle. The flowers parts, other than the pistil, when arranged in this manner are referred to as being hypogynous. Superior ovary with perigynous floral parts In some flowers the sepals, petals and stamens are attached to the margin of a cup - shaped structure which surrounds the ovary. The ‘cup - like structure’ may be either an extension of the receptacle, or hypanthium or ‘floral tube’ which is made up of the fusion of the lower portions of the sepals, petals and stamens. The hypanthium is not fused at any point to the ovary, therefore, the ovary is still a superior ovary. The flower parts other than the pistil are referred to as being perigynous. Inferior ovary with epigynous floral parts This is where the ovary is fused to an extension of the receptacle or the hypanthium below the point of attachment to the sepals, petals and stamens. The ovary tissue is fused around the base of the pistil (that is fused to the ovary tissue). Botany Reference Guide CII 106746951 (3 March 2016) Page 20 The flower parts other than the pistil are referred to as being epigynous. Half - inferior ovary with perigynous floral parts This is where the ovary is partly fused to an extension of the receptacle or the hypanthium so that a portion of the ovary is visibly free from it. The flower parts other than the pistil referred to as being perigynous. Botany Reference Guide CII 106746951 (3 March 2016) Page 21 1.4 Reproductive growth In the life cycle of a ‘typical’ flowering plant, a seed is planted, it germinates and grows into a plant which produces flowers. These flowers usually contain stamens and pistils. Pollen is formed within the anthers of the stamens and is transferred by the process of pollination to the stigma of the pistil. Pollen tubes grow down the style and eventually penetrate the embryo sacs. Here the male gamete (contained in the pollen) and the female gamete (the egg which is formed within the embryo sac) join to form a zygote which develops into the embryo. The male and female gametes (or the sex cells) are formed by a type of cell division called meiosis. This process halves their chromosome number so that, when fertilisation occurs and the male and female gametes unite, the gene - carrying chromosome sets combine to form a zygote. This is called fertilisation. These two processes of meiosis and fertilisation form the basis of the sexual reproductive cycle. Growth from the zygote state into the mature plant is by cell division called mitosis. Pollination Pollination is the transfer of pollen from the anther to the stigma of a flower. Self - pollination is pollination of a stigma by pollen from the same flower or another flower of the same plant. Cross - pollination is the transfer of pollen from the anther of a flower of one plant to the stigma of a flower of another plant of the same or related species. Figure 3 (a) Self - pollination Botany Reference Guide CII 106746951 (3 March 2016) (b) Cross – pollination Page 22 Cross-pollination results in increased variability of the offspring, because two individuals contribute to the genetic material of the resulting individual. Increased variability allows for greater adaptability to new environments and so affords an evolutionary advantage to a species. Cross-pollination in many species results in the production of more seeds or greater vigour of the offspring. Many angiosperms have adaptations that prevent or reduce self - pollination. Examples of these include: stamens and pistils of the same flower maturing at different times; stigma unreceptive to the flower’s own pollen; stigma held up above the anthers in the flower so the pollen is placed on the stigma by pollinating animals which have collected pollen from a different plant; individuals which are dioecious or monoecious. Pollination agencies The transfer of pollen from anther to stigma (pollination) may be carried out by: insects such as bees, wasps, butterflies, moths, beetles and flies birds small mammals such as bats. Animals pollinate flowers whilst feeding on pollen, nectar secreted by nectaries or on the flower parts themselves. Some pollen from the anther sticks to the animal and is removed when it comes into contact with the sticky surface of the style. Animals may be attracted to certain flowers by the scent, colour or pattern of the flowers, which they learn are a source of food. wind pollinates flowers that are usually small and inconspicuous, and without bright colours or nectar. The perianth is often reduced or lacking and stamens may protrude from the flower, while the stigmas are often large and branched. The pollen grains are small, light and dry and are commonly produced in larger quantities than in animal pollinated flowers. All these features serve to help pollination by wind. Pollen germination Once pollen grains are in contact with the stigma, they germinate to produce a pollen tube which grows downwards through the style to the ovule. If the generative cell has not already divided, it soon does, forming the two sperm Botany Reference Guide CII 106746951 (3 March 2016) Page 23 (the male gametes). Figure 4 - Growth of the pollen tube The tube nucleus plus the two sperm cells move down the pollen tube which passes through the micropyle in the integuments of the ovule and the two sperm are released into the embryo sac. Figure 5 - Release of the sperm into the embryo sac Botany Reference Guide CII 106746951 (3 March 2016) Page 24 Fertilisation Fertilisation is the union of the haploid (n) male and female gametes to form a diploid (2n) cell called the zygote, which develops into a diploid individual as a result of mitotic division. In the embryo sac, one of the two sperm nuclei fuses with the egg nucleus to form the zygote (2n). The second sperm nucleus fuses with both of the polar nuclei of the central cell which forms the triploid (3n) primary endosperm nucleus. Because both sperm nuclei fuse with nuclei of the other cells, this process (which occurs in angiosperms only) is known as double fertilisation. Figure 6 - Fertilisation This occurs in the ovule just after fertilisation. Note that the fertilisation process has to be repeated for every ovule, so for flowers of watermelons and pumpkins (and every other plant which produces masses of seeds) hundreds of separate pollen grains have to germinate and grow down the style, one to each ovule. Seed development Following fertilisation, the zygote produced by fertilisation divides by mitosis to produce the embryo. The primary endosperm nucleus begins to divide and forms a multicellular tissue, the endosperm. The endosperm grows by means of food supplied by the parent plant and it in turn nourishes the embryo. The embryo continues to grow until all the endosperm is absorbed. In some seeds, the nucellus develops into a food - storing tissue known as the perisperm. As the embryo matures it develops one or two cotyledons (seed leaves) which accumulate most of the food from the endosperm. Above the node between the cotyledons is a stem, called the epicotyl, which may bear young Botany Reference Guide CII 106746951 (3 March 2016) Page 25 leaves. These young leaves together with the epicotyl make up the plumule, which will form the shoot of the plant after germination. The basal end of the embryo is called the radicle, which develops into the primary root after germination. The region between the radicle and the cotyledons is the embryonic stem or hypocotyl. The seed is surrounded by the seed coat or testa which develops from the integuments of the ovule. The micropyle is often visible on the seed coat as a small pore and is commonly associated will a scar called the hilum, which remains after the seed separates from the funiculus. As the seed ripens, the embryo gradually passes into a dormant state in which it remains until it germinates. Development of fruit These changes which mark the transition of the flower into a young fruit are called fruit set. The capacity of the flower to set fruit depends in many instances on the receptivity of the female parts to the pollen. The whole process of growth is summarised in the figure below which shows the basic life cycle of an angiosperm. Botany Reference Guide CII 106746951 (3 March 2016) Page 26 Figure 7 - Basic life cycle of an angiosperm Botany Reference Guide CII 106746951 (3 March 2016) Page 27