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"Double-flowered" describes varieties of
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
s with extra
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s, often containing flowers within flowers. The double-flowered trait is often noted alongside the
scientific name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...
with the abbreviation ''fl. pl.'' (''flore pleno'', a
Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...
ablative In grammar, the ablative case (pronounced ; sometimes abbreviated ) is a grammatical case for nouns, pronouns, and adjectives in the grammars of various languages; it is sometimes used to express motion away from something, among other uses. T ...
form meaning "with full flower"). The first abnormality to be documented in flowers, double flowers are popular varieties of many commercial flower types, including
rose A rose is either a woody perennial flowering plant of the genus ''Rosa'' (), in the family Rosaceae (), or the flower it bears. There are over three hundred species and tens of thousands of cultivars. They form a group of plants that can be ...
s,
camellia ''Camellia'' (pronounced or ) is a genus of flowering plants in the family Theaceae. They are found in eastern and southern Asia, from the Himalayas east to Japan and Indonesia. There are more than 220 described species, with some controversy ...
s and
carnation ''Dianthus caryophyllus'' (), commonly known as the carnation or clove pink, is a species of ''Dianthus''. It is likely native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years.Med ...
s. In some double-flowered varieties all of the reproductive organs are converted to petals — as a result, they are sexually sterile and must be propagated through cuttings. Many double-flowered plants have little wildlife value as access to the nectaries is typically blocked by the mutation.


History

Double flowers are the earliest documented form of floral abnormality, first recognized more than two thousand years ago.
Theophrastus Theophrastus (; grc-gre, Θεόφραστος ; c. 371c. 287 BC), a Greek philosopher and the successor to Aristotle in the Peripatetic school. He was a native of Eresos in Lesbos.Gavin Hardy and Laurence Totelin, ''Ancient Botany'', Routledge ...
mentioned double roses in his ''Enquiry into Plants'', written before 286 BC.
Pliny Pliny may refer to: People * Pliny the Elder (23–79 CE), ancient Roman nobleman, scientist, historian, and author of ''Naturalis Historia'' (''Pliny's Natural History'') * Pliny the Younger (died 113), ancient Roman statesman, orator, w ...
also described double roses in 1st century BC. In China, double
peonies The peony or paeony is a flowering plant in the genus ''Paeonia'' , the only genus in the family Paeoniaceae . Peonies are native to Asia, Europe and Western North America. Scientists differ on the number of species that can be distinguished, ...
were known and selected by around 750 AD, and around 1000 AD double varieties of roses were cultivated to form the China rose (one of the ancestors of modern
hybrid tea rose Hybrid tea is an informal horticultural classification for a group of garden roses. The first hybrid tea roses were created in France in the mid-1800s, by cross-breeding the large, floriferous Hybrid Perpetuals with the tall, elegant Tea roses. ...
s). Today, most cultivated rose varieties bear this double-flower trait. Herbalists of the Renaissance recognized double flowers and began to cultivate them in their gardens—
Rembert Dodoens Rembert Dodoens (born Rembert Van Joenckema, 29 June 1517 – 10 March 1585) was a Flemish physician and botanist, also known under his Latinized name Rembertus Dodonaeus. He has been called the father of botany. Life Dodoens was born Rember ...
published a description of double flowers in 1568, and
John Gerard John Gerard (also John Gerarde, c. 1545–1612) was an English herbalist with a large garden in Holborn, now part of London. His 1,484-page illustrated ''Herball, or Generall Historie of Plantes'', first published in 1597, became a popular gard ...
created illustrations of many double flowers beside their wild-type counterparts in 1597. A double-flowered variety of
Marsh Marigold ''Caltha palustris'', known as marsh-marigold and kingcup, is a small to medium size perennial herbaceous plant of the buttercup family, native to marshes, fens, ditches and wet woodland in temperate regions of the Northern Hemisphere. It flower ...
was discovered and cultivated in Austria in the late 16th century, becoming a valued garden plant. The first documented double-flowered mutant of ''
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organi ...
'', a model organism for plant development and genetics, was recorded in 1873. The mutated gene likely responsible for the phenotype, AGAMOUS, was cloned and characterized in 1990 in
Elliot Meyerowitz Elliot Meyerowitz (born May 22, 1951) is an American biologist. Career Meyerowitz did his undergraduate work at Columbia University (A.B. in biology, 1973), where he worked part-time in the laboratory of Cyrus Levinthal on combined microscopic a ...
's lab as part of his study of molecular mechanisms of pattern formation in flowers.


Genetics of double-flower mutations

Double-flower forms often arise when some or all of the
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s in a flower are replaced by
petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s. These types of mutations, where one organ in a developing organism is replaced with another, are known as
homeotic In evolutionary developmental biology, homeosis is the transformation of one organ into another, arising from mutation in or misexpression of certain developmentally critical genes, specifically homeotic genes. In animals, these developmental ge ...
mutations. They are usually
recessive In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and t ...
, although the double flower mutation in
carnation ''Dianthus caryophyllus'' (), commonly known as the carnation or clove pink, is a species of ''Dianthus''. It is likely native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years.Med ...
s exhibits
incomplete dominance In genetics, dominance is the phenomenon of one variant (allele) of a gene on a chromosome masking or overriding the effect of a different variant of the same gene on the other copy of the chromosome. The first variant is termed dominant and ...
. In ''
Arabidopsis ''Arabidopsis'' (rockcress) is a genus in the family Brassicaceae. They are small flowering plants related to cabbage and mustard. This genus is of great interest since it contains thale cress (''Arabidopsis thaliana''), one of the model organi ...
'', which has been used as a model for understanding flower development, the double-flower gene AGAMOUS encodes a protein responsible for tissue specification of stamen and
carpel Gynoecium (; ) is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of (one or more) ''pistils'' ...
flower segments. When both copies of the gene are deleted or otherwise damaged, developing flowers lack the signals to form stamen and carpel segments. Regions which would have formed stamens instead default to petals and the carpel region develops into a new flower, resulting in a recursive
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
-petal-petal pattern. Because no stamens and carpels form, the plants have no reproductive organs and are sexually sterile. Mutations affecting flower morphology in ''Arabidopsis'' can be described by the ABC model of flower development. In this model, genes involved in flower formation belong to one of three classes of genes: A class genes which affect sepal and petal formation, B class genes which affect petal and stamen formation, and C class genes which affect stamen and carpel formation. These genes are expressed in certain regions of the developing flower and are responsible for development of organs in those regions. Agamous is a C class gene, a
transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The fu ...
responsible for activating genes involved in stamen and carpel development.


Gallery

Image:Bridal_pink_-_morwell_rose_garden.jpg, Most garden
rose A rose is either a woody perennial flowering plant of the genus ''Rosa'' (), in the family Rosaceae (), or the flower it bears. There are over three hundred species and tens of thousands of cultivars. They form a group of plants that can be ...
s are double-flowered Image:Lilac.head.600pix.jpg, Double-flowered
lilac ''Syringa'' is a genus of 12 currently recognized species of flowering plant, flowering woody plants in the olive family or Oleaceae called lilacs. These lilacs are native to woodland and scrub from southeastern Europe to eastern Asia, and wid ...
, often called "French lilac" Image:Double-flowered_Camellia.jpg, Double-flowered
Camellia ''Camellia'' (pronounced or ) is a genus of flowering plants in the family Theaceae. They are found in eastern and southern Asia, from the Himalayas east to Japan and Indonesia. There are more than 220 described species, with some controversy ...
Image:Gartennelke_1.jpg, Double-flowered
carnation ''Dianthus caryophyllus'' (), commonly known as the carnation or clove pink, is a species of ''Dianthus''. It is likely native to the Mediterranean region but its exact range is unknown due to extensive cultivation for the last 2,000 years.Med ...
s Image:DOUBLE_PINK_PATUNIA.jpg, Double-flowered
petunia ''Petunia'' is genus of 20 species of flowering plants of South American origin. The popular flower of the same name derived its epithet from the French, which took the word ''petun'', meaning "tobacco," from a Tupi–Guarani language. A tende ...
Image:Paeonia_officinalis_officinalis0.jpg, Double-flowered
peony The peony or paeony is a flowering plant in the genus ''Paeonia'' , the only genus in the family Paeoniaceae . Peonies are native to Asia, Europe and Western North America. Scientists differ on the number of species that can be distinguished, ...
Image:Kanzan, Cerasus lannesiana 'Sekiyama' Koidzumi.JPG, Double-flowered ''Prunus'' 'Kanzan'


References

{{DEFAULTSORT:Double-Flowered Flowers Plant morphology