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Flowers, the reproductive structures of flowering plants, typically consist of four circular levels: modified leaves, petals to attract pollinators, male parts bearing pollen, and female parts receiving pollen. Flowers enable sexual reproduction through pollination, either by cross-pollination or self-pollination, facilitated by animals or wind. After fertilisation, seeds develop within fruits, which protect seeds and aid dispersal by animals or natural forces. Evolving between 150 and 190 million years ago in the Jurassic, flowers revolutionized plant reproduction and have played vital roles in ecosystems and human culture, symbolizing beauty, medicine, and art.

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Etymology

In botany, flowers are defined as the reproductive structures of angiosperms (flowering plants),1 while cones are regarded as the gymnosperm equivalent.23 Bloom is similarly defined, but may also be used to describe the collective of flowers on a plant, as in the phrase: covered with bloom.4 Flower is also commonly used to describe the whole of a plant that produces flowers.5

Flower is from the Middle English word flour, which referred to both the ground grain and the reproductive structure in plants, before diverging in the 17th century.6 It comes originally from the Proto-Italic *flōs ('flower'; cf. Latin flōs, flōris).7 The Old English word for flower was blossom,8 which is still used today, but refers especially to the flowers of edible fruit trees, and not to the whole flowering plant.9 Flower, bloom, and blossom are all cognates and are derived from the Proto-Indo-European word *bʰleh₃ōs ('blossoming').10 Both bloom and blossom refer to flowers as well as the state of flowering; as in the phrases: in bloom or in blossom.11

Function

The main purpose of a flower is reproduction of the individual,12 aiding in the survival of the species.13 Flowers not only produce spores, which become gametophytes that produce sex cells, leading to fertilised cells, but also develop and help disseminate seeds.14 Sexual reproduction between plants results in evolutionary adaptation, which improves species survival. Plants favour cross-pollination because it promotes the joining of sex cells from genetically distinct plants of the same species, thereby increasing genetic diversity. Facilitating this process is a key function of flowers and is often reflected in their form and structure.15 Features designed to attract pollinators are among the most common adaptations.16

Structure

Main article: Floral morphology

The structure of a flower, termed its morphology,17 can be considered in two parts: the vegetative part, consisting of non-reproductive structures such as petals; and the reproductive or sexual parts. A stereotypical, or complete,18 flower is made up of four kinds of structures arranged in sets called whorls. They grow around the tip of a short stalk or axis, called a receptacle.19 The four main whorls (starting from the base of the flower and working upwards) are the calyx, petals, androecium, and gynoecium.20

Vegetative

Main article: Perianth

The non-reproductive or vegetative part of the flower, known collectively as the perianth, consists of calyx (the modified outer leaves), and the petals. The receptacle is the thickened part of the flower stalk, called the pedicel, which supports all of the other flower structures.2122

Calyx

The sepals, collectively called the calyx, are modified leaves that occur on the outermost whorl of the flower. They are leaf-like,23 in that they have a broad base, pores, green pigment, and may have analogous outgrowths from the stem. Sepals are often waxy, tough, and grow quickly to protect the flower as it develops.2425 Although they sometimes fall off at maturity, sepals more commonly persist to protect the fruit and aid in its dispersal.26 The sepals in some flowers may be partially or completely fused together.2728

Petals

The petals, collectively called the corolla,29 are almost or completely fibreless leaf-like structures that form the innermost whorl of the perianth. They are often delicate and thin and are usually coloured, shaped, or scented, to encourage and facilitate pollination.30 The petals may be fused together.31 Petals also tend to have patterns only visible under ultraviolet light, which is visible to pollinators but not to humans.32 In some flowers, petals and sepals are indistinguishable from one another.33

Reproductive

Main article: Plant reproductive morphology

All flowering plants are heterosporous, that is, every individual plant produces two types of spores. Spores are formed from mature plants, which contain two sets of chromosomes, and are divided into microspores and megaspores—the precursors to pollen and embryo sacs respectively. Pollen and embryo sacs are the male and female gametophytes, sex cell-producing structures, and contain just one set of chromosomes. Microspores are produced by meiosis inside anthers, the male part of flowers,34 and megaspores are produced inside ovules contained within the ovary.3536 As with all heterosporous plants, the gametophytes also develop inside the spores.37

Male

The androecium is the whorl of male parts called stamens, which produce pollen. Stamens consist typically of an anther, made up of four pollen sacs arranged in two sheaths called thecae, connected to a filament, or stalk.3839 The anther contains microspores which become pollen, the male gametophyte, after undergoing meiosis.40 Although they exhibit the widest variation among floral organs,41 the androecium is usually confined just to one whorl and to two whorls only in rare cases.42

Female

The gynoecium, consisting of one or more carpels, is the female part of the flower and found on the innermost whorl.43 Each carpel consists of: a stigma, which receives pollen; a style, the stalk; and an ovary, which contains the ovules, and the female gametophytes by extension. Carpels may be fused together and are often described collectively as a pistil. Inside the ovary, the ovules are attached to the placenta by structures called funiculi.4445

Variation

Although most plants have flowers with four whorls—protective leaves, petals, male parts, and female parts—and their typical sub-structures, they vary greatly between flowering plants.4647 This variation encompasses all aspects of flowers, including size, shape, and colour.48 Flowers range in size from 0.1 mm (1⁄250 in) (duckweed) to 1 m (3.3 ft) in diameter (corpse flower).49 Additionally, the four main parts of a flower are generally defined by their positions and not by their function. Many flowers lack some parts, have parts that are modified for other functions, or contain parts that look like what is typically another part.505152 In some flowers, organs such as stamens, stigmas, and sepals are modified to resemble petals. This is most common in cultivation (such as of roses), where flowers with many additional "petals" are found to be more attractive.5354

Most flowers have symmetry.55 When the flower is bisected through the central axis from any point and symmetrical halves are produced,56 the flower is said to be regular (as in sedges). This is an example of radial symmetry. If there is only one plane of symmetry (as in orchids),57 the flower is said to be irregular. If, in very rare cases, they have no symmetry at all they are called asymmetric.5859 Floral symmetry is a key driver of diversity in flower morphology, because it is one of the main features derived through flower-plant coevolution. Irregular flowers often coevolve with specific pollinators, while radially symmetric flowers tend to attract a wider range of pollinators.6061

In the majority of species, individual flowers have both female parts and male parts. These flowers are synonymously described as being perfect, bisexual, or hermaphrodite. In some species of plants, the flowers are imperfect or unisexual: having only either male or female parts. If unisexual male and female flowers appear on the same plant, the species is called monoecious. However, if an individual plant is either female or male, the species is called dioecious.62 Many flowers have nectaries, which are glands that produce nectar: a sugary fluid used to attract pollinators. Their shape varies between different plants,63 are they not considered as an organ on their own.64

Some flowers are lacking or have only a highly reduced stalk, and so are attached directly to the plant.65 There are several structures, found in some plants, that resemble flowers or floral organs. These include: coronas, crown-like outgrowths;66 and pseudonectaries, that look like nectaries but do not contain nectar.67 In plants where disease has taken hold, phyllody—leafy flower parts—may occur.68

Inflorescence

Main article: Inflorescence

In plants that have more than one flower on an axis, the collective cluster of flowers is called an inflorescence.69 Some inflorescences are composed of many small flowers arranged in a formation that resembles a single flower. These are known as pseudanthia.70 A single daisy or sunflower, for example, is not a flower but an inflorescence composed of numerous florets, or tiny flowers.71 An inflorescence may include specialised stems and modified leaves known called bracts, as well as smaller bracteoles.72

Floral diagrams and formulae

Main articles: Floral formula and Floral diagram

A floral formula is a way to represent the structure of a flower using letters, numbers, and symbols in a compact way. It can represent both group of species or a particular species, and usually gives ranges for the numbers of different organs. The format of floral formulae differs in different parts of the world, but the formulae all convey the same information.7374

Floral diagrams are schematic diagrams that can be used to show important features of flowers, including the relative positions of the various organs, the presence of organ fusion and symmetry, and structural details.75

Colour

In contrast to the mostly green vegetative parts of plants, flowers are often colourful. This includes the petals and, in some plants, the stamens, anthers, stigmas, ovaries, pollen, styles, and even nectar.76 These colours are produced mainly by biological pigments, which are molecules that can absorb and retain energy from light.7778 Specific pigments, and so colours, provide different benefits to the plant. These benefits include protecting the plant against degradation and guiding pollinators—both general and specific—to the plant.7980

Colour, or colour effects, may also be produced by structural coloration, in which colour is produced by tiny surface structures interfering with waves of light.81 This includes iridescence (as in some tulips) and photonic crystals (as in edelweiss), which diffract light using tiny grooves.8283 The colour of flowers can also change; sometimes this acts as a signal to pollinators (as in Viola cornuta). Change may also occur as a result of temperature; pH, as in the anthoxanthins found in Hydrangea; metals; sugars; and cell shape.84

Development

Further information: ABC model of flower development

Floral development begins with the transformation of vegetative growth into floral growth.85 This is regulated by both genetic and environmental factors.86 The eventual formation of a flower starts with a shoot apical meristem (SAM): a group of dividing cells responsible for leaves and buds. The organs which make up a flower—in most cases the sepals, petals, male parts, and female parts—grow out of a growth-limited floral meristem, which a SAM creates.87 The ABC model of flower development can be used, for many plants, to describe how groups of genes come together to induce each organ being produced.88 In general, all aspects of floral development are controlled by a gene regulatory network of specialised MADS-box genes—which includes the ABC genes—and associated proteins.8990 For plants, the transition into flowering is a major change and must occur at the right time so as to ensure reproductive success. Plants determine this time by interpreting both internal and environmental cues, such as day length.91

The ABC model was the first unifying principle in the development of flowers, and its major tenets have been found to hold in most flowering plants.92 It describes how three groups of genes—A, B, and C—are responsible for the development of flowers. These three gene groups' activities interact together to determine the developmental identities of the primordia organ within the floral apical meristem. Alone, A genes produce sepals in the first whorl. Together, A and B produce the petals in the second whorl. C genes alone produce carpels in the centre of the flower. C and B together produce the stamens in the third whorl.93 This can also be extended to the more complex ABCDE model, which adds an additional two gene groups to explain the development of structures like ovules.94

The transition to flowering is one of the major phase changes that a plant makes during its life cycle.95 The transition must take place at a time that is favourable for fertilisation and the formation of seeds, hence ensuring maximal reproductive success. To meet these needs a plant can interpret important internal and environmental cues such as: changes in levels of plant hormones (such as gibberellins),96 seasonable temperature, and day length changes.97 Many plants, including many of those that have more than two-year lifespans and just two-year lifespans, require cold exposure to flower.9899100 These cues are interpreted molecularly through a complex signal called florigen, which involves a variety of genes. Florigen is produced in the leaves in reproductively favourable conditions and acts in stem tips to force switching from developing leaves to flowers.101 Once developed, flowers may selectively open and close their flowers at different times of day; usually around dusk and dawn.102 They may also track the path of the sun to remain warm—potentially both for their own benefit and to attract pollinators. Both of these mechanisms are controlled by a plant's circadian rhythm and in response to environmental changes.103

Pollination

Main article: Pollination

Since the flowers are the reproductive organs of the plant, they mediate the joining of the sperm, contained within pollen, to the eggs in the ovules—contained in the ovary.104 Pollination is this movement of pollen from the male parts to the female parts.105 It occurs either between flowers (or from one part of a flower to another) of the same plant, as in self-pollination, or between flowers of different plants, as in cross-pollination. Cross-pollination is more common in flowering plants as it increases genetic variation.106107 Pollination typically only takes place when the flower is fully expanded and functional.108

Flowering plants usually face evolutionary pressure to optimise the transfer of their pollen, and this is typically reflected in the morphology of their flowers and their reproductive strategies.109110 Agents that transport pollen between plants are called vectors. Around 80% of flowering plants make use of biotic or living vectors. Others use abiotic or non-living vectors, or some combination of the two.111112

Biotic pollination

Further information: Pollination syndrome

Flowers that use biotic vectors attract and use animals to transfer pollen from one flower to the next. Often they are shaped and designed to both attract pollinators and ensure pollen is transferred effectively.113114 Flowers most commonly employ insects,115116 but also: birds, bats, lizards,117 other mammals,118 snails and slugs,119 and, in rare cases, crustaceans and worms.120 Rewards given to pollinators by flowers to encourage pollination include: food (such as pollen, starch, or nectar), mates, shelter, a place to raise their young, and pseudocopulation (sexual deception).121 In the latter, the flower is scented or shaped so as to encourage sexual arousal and pollination from the subsequent intercourse.122 They may also be attracted by various stimuli such as size and scent (as in carrion flowers). Colour is also a factor, and includes nectar guides, which show pollinators where to look for nectar; they may be visible only under ultraviolet light.123124125

Many flowers have close relationships with just one or a few specific pollinators. They may be structured to allow or encourage pollination from these organisms. This increases efficiency, because there is a higher chance pollination comes from pollen of the same species of plant.126 This close relationship is an example of coevolution, as the plant and pollinator have developed together over a long period to match each other's needs.127

Abiotic pollination

Main articles: Anemophily and Hydrophily

Flowers that use abiotic, or non-living, vectors use the wind or, much less commonly, water, to move pollen from one flower to the next.128 Wind-dispersed species do not need to attract pollinators and therefore tend not to grow large, showy, or colourful flowers, and do not have nectaries, nor a noticeable scent.129 Whereas the pollen of insect-pollinated flowers is usually large, sticky, and rich in protein to act as a "reward", wind-pollinated flowers' pollen is typically small, very light, smooth, and of little nutritional value.130131

Fertilisation and seed development

Fertilisation is the fusion of the male and female sex cells to produce a zygote, from which the new organism develops.132 In humans, sexual intercourse results in the depositing of sperm cells into the vagina. Although not all survive, they travel until one reaches the egg in the fallopian tube, where the male and female sex cells fuse in the process of fertilisation.133

In flowering plants, fertilisation is preceded by pollination, which is the movement of pollen from the stamen to the carpel. It encompasses both plasmogamy, the fusion of the protoplasts (cell without cell wall), and karyogamy, the fusion of the nuclei. When pollen lands on the stigma of the flower it begins creating a pollen tube, which runs down through the style and into the ovary. After penetrating the centre-most part of the ovary it enters the egg apparatus and is guided by a specialised cell.134

Next, the end of the pollen tube bursts and releases the two sperm cells, one of which makes its way to an egg, while also losing its cell membrane and much of the jelly-like substance that fills its cells. The sperm's nucleus then fuses with the egg's nucleus, resulting in the formation of a zygote; a diploid cell, containing two copies of each chromosome.135136 Flowering plants undergo double fertilisation, which involves both karyogamy and plasmogamy. In double fertilisation the second sperm cell subsequently also fuses with the two polar nuclei of the central cell. Since all three nuclei are haploid, they result in a large nutrient tissue nucleus which is triploid.137

Seed and fruit development

Main articles: Seed development and Fruit § Development

Following its formation, the zygote begins to grow through nuclear and cellular divisions, called mitosis, eventually becoming a small group of cells. One section of it becomes the embryo,138 while the other becomes the suspensor; a structure which forces the embryo into the endosperm and is later undetectable. Two small groups of cells also form at this time, which later become the cotyledon, or initial leaf, which is used as an energy store. The next stage involves the growth of several key structures, including: the embryotic root, the embryotic stem, and the root or shoot junction itself. In the final step, vascular tissue develops around the seed.139

The ovary, inside which the seed is forming from the ovule, grows into a fruit. All the other main floral parts wither and die during this development, including: the style, stigma, stamens, petals, and sepals. This process is called floral senescence; it is often accelerated or initiated by the completion of pollination. Death is preferred because flowers are costly to the plant; nevertheless, flowers can last for between a few hours and several months.140141 The fruit contains three main structures: the outer layer of peel; the fleshy part; and the stone, or innermost layer. The pericarp, which may include one or more of these structures, represents collectively the fruit wall—everything but the seed. The size, shape, toughness, and thickness of the pericarp varies among different dry and fleshy fruits. These traits are directly connected to the plant's method of seed dispersal, since the purpose of fruit is to encourage or enable the seed's dispersal and protect the seed while doing so.142143

Seed dispersal

Main articles: Biological dispersal and Seed dispersal

Following the pollination of a flower, fertilisation, and finally the development of a seed and fruit, a mechanism, or vector, is typically used to disperse the fruit away from the plant.144 In flowering plants, seeds are dispersed away from the plant so as to not force competition between the mother and daughter plants,145 as well as to enable the colonisation of new areas. Vectors can generally be divided into two categories: external vectors and internal vectors.146147 External vectors include living things like birds or bats, or non-living things such as water and wind.148149 Internal vectors, which are derived from the plant itself,150151 include, for example, the fruit exploding to release the seeds, as in dwarf mistletoes.152

Evolution

Further information: Evolutionary history of plants § Flowers, and Floral biology

Flowers originated between 150 and 190 million years ago, during the Jurassic.153154 Although molecular analyses indicate this early appearance of angiosperms—flowering plants, the earliest definitive evidence from the fossil record comes from between 125 and 130 years ago, during the Early Cretaceous.155156157158 The exact time at which angiosperms diverged from other seed plants is a classic open question in evolutionary biology.159160161 Prior to the advent of flowers, plants reproduced using cones (as in gymnosperms),162 and spores (as in pteridophytes).163 The transformation of spore-producing leaves into structures like stamens and carpels, is the most clear milestone in the complex evolution of flowers.164 There is debate both over whether these and other changes happened gradually or as sudden shifts like homeotic mutations, and which aspect of flower morphology came first.165166

The flower was the angiosperms' most significant evolutionary innovation,167 granting the ability to effectively take advantage of animal pollinators.168 Other evolutionary advantages included: being able to have both male and female parts on the same axis; and on this axis have carpels, to protect the ovules; stamens, to present the pollen; and the perianth, to provide protection. In addition, they pioneered double fertilisation, which allows energy investment (into endosperm) to be prolonged until after pollination. The gametophytes, which lead to sex cells, were very reduced, which allowed for greater protection of this critical process.169 The net effect of these features was greater reproductive security and efficiency.170 This allowed the angiosperms to replace many other seed plants—such as Pinales, cycads, Gnetophyta and Ginkgoales—in the majority of ecosystems.171

A key driving force in the evolution of flowers is coevolution, where pollinator and flower evolve with one another,172 often to their mutual benefit. This is particularly prominent in insect species such as bees, but is also found in flower-pollinator relationships with birds and bats. Many flowers have evolved in such a way so as to make pollination by specific species easier, thus providing greater efficiency and also ensuring higher rates of pollination. This is because they receive less pollen from other plant species.173174 However, this close interdependence increases the risk of extinction, since the extinction of either member almost certainly means the extinction of the other member as well.175 Modern-day flowers exhibit a variety of features derived through coevolution including: shape, size, symmetry, timing of flower opening, colour, scent, and pollinator rewards (including pollen, nectar, and oils).176177 For example, Japanese honeysuckle flowers strategically open during the night to attract nocturnal moths, which are more efficient pollinators than diurnal bees.178 With the innovation of the flower—and other adaptations—angiosperms rapidly diversified.179 Approximately 90% of all living land plant species are angiosperms.180 This is attributed, in part, to coevolution, which caused specialisation and so speciation; where populations diverge into separate species.181 Both the strength of close pollinator-flower relationships and the survival of either species are effected by climate change. Reducing numbers of pollinators have led to the extinction of many flowering plants.182

Taxonomy

Further information: Plant taxonomy and Linnaean taxonomy

In plant taxonomy, which is the study of plant classification and identification, the morphology of plants' flowers is used extensively—and has been since at least classical Greece.183184 Despite earlier works, Carl Linnaeus's 1753 book Species Plantarum, in which he laid out his system of classification, is regarded as the first taxonomic work to recognise the significance of flowers.185186 He identified 24 classes of flowering plants, based mainly on the number, length, and union of the stamens.187188189 Subsequent systems in the 18th and 19th centuries focused more on natural characteristics. This included taking into account the rest of the plant, so that diverse plants weren't put into the same groups, as often happened in Linnaeus's system.190191192

In 1963, the biologists Robert Sokal and Peter Sneath created the method of numerical taxonomy, which differentiates taxa based on their tabulated morphological characteristics; such as their flowers. This was an effort to make plant taxonomy more objective, but it remained inconsiderate of evolution, and so not useful in that context.193 While this and earlier methods, such as Linnaeus's, used morphological features, many botanists today employ genetic sequencing, the study of cells, and the study of pollen. These come as a result of advancements in DNA-related science.194 Despite this, morphological characteristics such as the nature of the flower and inflorescence still make up the bedrock of plant taxonomy.195196197

Uses

Further information: Flowering plant § Human uses, and Human uses of plants

Humans have used flowers globally for millennia for many purposes, including decoration, medicine, drugs,198 food, spices,199 perfumes,200 and essential oils. Many flowers are edible and are often used in drinks and dishes, such as salads, for taste, scent, and decoration.201 Inflorescences and the bracts or stems of some flowers are commonly described as vegetables. These include: broccoli, cauliflower, and artichoke. Flowers may be eaten freshly after being picked, or dried and eaten later.202 Floristry is the production and sale of flowers, and involves preparing freshly cut flowers and arranging them—in a bouquet, for example—to the client's liking.203

Most crop plants are have flowers,204 and they produce much of the most common crop products—such as seeds and fruits;205 around half of all cropland is used to grow three flowering plants: rice, wheat, and corn.206 Flowers are steeped to make teas, either alone, as in herbal teas, or in combination with the tea plant.207208 Essential oils and other flower extracts are widely used in herbal medicines and decoctions because they contain phytochemicals and may have anti-microbial effects.209210 Flowers from many plants are also used in the production of drugs, such as cannabis, bush lily, and Madagascar periwinkle.211 Some flowers are used in cooking as spices, these include saffron and cloves; derived from Crocus and Syzygium aromaticum respectively.212

In culture

Further information: Human uses of plants § Symbolic uses

Flowers are the subject of much symbolism, and feature often in art, ritual, religious practices, and festivals. Plants have been cultivated in gardens for their flowers for around ten thousand years.213214 Flowers are associated with burial in many cultures, and are often placed by headstones to pay respect.215216 They are also placed by statues or temples of religious or other figures—sometimes formed into floral wreaths.217218 In some places, the dead are buried covered in flowers or on a bed of flowers.219 They are also associated with love and celebration, and given to others in many places for this reason.220221 Economic demand has led to the cultivation of flowers that are longer-lasting, more colourful, and visually appealing.222

Flowers feature extensively in art across a variety of mediums, and different flowers are ascribed symbolic meanings.223224 For example, violets may represent modesty, virtue, or affection.225 In addition to hidden meanings, flowers are used in flags, emblems, and seals. In this way, they represent countries or places. Some countries have national flowers; for example, Hibiscus × rosa-sinensis is the national flower of Malaysia.226 In literature, flowers feature in imagery of places and as metaphors for pleasure, beauty, and life.227

Notes

Bibliography

  • The dictionary definition of flower at Wiktionary
  • Media related to Flowers at Wikimedia Commons
  • Quotations related to Flowers at Wikiquote

References

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  2. Mauseth 2016, p. 221. - Mauseth, James D. (2016). Botany: an introduction to plant biology (6th ed.). Jones & Bartlett Learning. ISBN 9781284077537. ↩

  3. There are some gymnosperm cones which resemble flowers. The cones of Ginkgo biloba, for example, are mostly considered to be simple strobili, and not flowers.[3] /wiki/Gymnosperm ↩

  4. Sinclair 1998, p. 169. - Sinclair, John M. (1998). Collins English dictionary (4th Revised ed.). HarperCollins. ISBN 9780004704531 – via Internet Archive. https://archive.org/details/collinsenglishdi0000unse_k7q4 ↩

  5. Sinclair 1998, p. 169. - Sinclair, John M. (1998). Collins English dictionary (4th Revised ed.). HarperCollins. ISBN 9780004704531 – via Internet Archive. https://archive.org/details/collinsenglishdi0000unse_k7q4 ↩

  6. Cresswell 2010, p. 172. - Cresswell, Julia (2010). Oxford dictionary of word origins. Internet Archive. Oxford University Press. ISBN 9780199547937. http://archive.org/details/oxforddictionary0000unse_p6k3 ↩

  7. de Vaan 2008, pp. 227–228. - de Vaan, Michiel (2008). Etymological dictionary of Latin and the other Italic languages. Brill. ISBN 9789004167971. https://books.google.com/books?id=ecZ1DwAAQBAJ ↩

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  9. Sinclair 1998, p. 169. - Sinclair, John M. (1998). Collins English dictionary (4th Revised ed.). HarperCollins. ISBN 9780004704531 – via Internet Archive. https://archive.org/details/collinsenglishdi0000unse_k7q4 ↩

  10. de Vaan 2008, pp. 227–228. - de Vaan, Michiel (2008). Etymological dictionary of Latin and the other Italic languages. Brill. ISBN 9789004167971. https://books.google.com/books?id=ecZ1DwAAQBAJ ↩

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  12. Beekman et al. 2016, p. 5. - Beekman, Madeleine; Nieuwenhuis, Bart; Ortiz-Barrientos, Daniel; Evans, Jonathan P. (2016). "Sexual selection in hermaphrodites, sperm and broadcast spawners, plants and fungi". Philosophical Transactions: Biological Sciences. 371 (1706): 1–13. doi:10.1098/rstb.2015.0541. ISSN 0962-8436. JSTOR 26143395. PMC 5031625. PMID 27619704. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5031625 ↩

  13. Pandey 2023, p. 7. - Pandey, A. K. (2023). Reproductive biology of angiosperms (1st ed.). CRC Press. ISBN 978-1-032-19620-6. ↩

  14. Mauseth 2016, p. 238. - Mauseth, James D. (2016). Botany: an introduction to plant biology (6th ed.). Jones & Bartlett Learning. ISBN 9781284077537. ↩

  15. Mauseth 2016, p. 238. - Mauseth, James D. (2016). Botany: an introduction to plant biology (6th ed.). Jones & Bartlett Learning. ISBN 9781284077537. ↩

  16. Mauseth 2016, pp. 239–240. - Mauseth, James D. (2016). Botany: an introduction to plant biology (6th ed.). Jones & Bartlett Learning. ISBN 9781284077537. ↩

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  18. Pandey 2023, p. 15. - Pandey, A. K. (2023). Reproductive biology of angiosperms (1st ed.). CRC Press. ISBN 978-1-032-19620-6. ↩

  19. De Craene 2010, p. 4. - De Craene, Louis P. Ronse (2010). Floral diagrams. Cambridge University Press. doi:10.1017/cbo9780511806711. ISBN 9780511806711. https://dx.doi.org/10.1017/cbo9780511806711 ↩

  20. De Craene 2010, p. 3. - De Craene, Louis P. Ronse (2010). Floral diagrams. Cambridge University Press. doi:10.1017/cbo9780511806711. ISBN 9780511806711. https://dx.doi.org/10.1017/cbo9780511806711 ↩

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