Variegated Lemon Thyme Thymus Citriodorus Variegata Leaves 3264px
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Variegation is the appearance of differently coloured zones in the leaves and sometimes the stems and fruit of plants. Species with variegated individuals are sometimes found in the
understory In forestry and ecology, understory (American English), or understorey (Commonwealth English), also known as underbrush or undergrowth, includes plant life growing beneath the forest canopy without penetrating it to any great extent, but abov ...
of
tropical rainforest Tropical rainforests are rainforests that occur in areas of tropical rainforest climate in which there is no dry season – all months have an average precipitation of at least 60 mm – and may also be referred to as ''lowland equatori ...
s, and this habitat is the source of a number of variegated houseplants. Variegation is caused by mutations that affect chlorophyll production or by viruses, such as mosaic viruses, which have been studied by scientists. The striking look of variegated plants is desired by many gardeners, and some have deliberately tried to induce it for aesthetic purposes. There are a number of gardening books about variegated plants, and some gardening societies specialize in them. The term is also sometimes used to refer to colour zonation in
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,
mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. ( ...
s, and the skin, fur, feathers or scales of animals.


Causes


Chimeral

Because the variegation is due to the presence of two kinds of plant tissue, propagating the plant must be by a vegetative method of propagation that preserves both types of tissue in relation to each other. Typically, stem cuttings, bud and stem
grafting Grafting or graftage is a horticultural technique whereby tissues of plants are joined so as to continue their growth together. The upper part of the combined plant is called the scion () while the lower part is called the rootstock. The succ ...
, and other propagation methods that results in growth from leaf axil buds will preserve variegation. Cuttings with complete variegation may be difficult, if not impossible, to propagate. Root cuttings will not usually preserve variegation, since the new stem tissue is derived from a particular tissue type within the root. Some variegation is due to visual effects caused by
reflection Reflection or reflexion may refer to: Science and technology * Reflection (physics), a common wave phenomenon ** Specular reflection, reflection from a smooth surface *** Mirror image, a reflection in a mirror or in water ** Signal reflection, in ...
of light from the leaf surface. This can happen when an air layer is located just under the
epidermis The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermis layer provides a barrier to infection from environmental pathogens and regulates the amount of water rele ...
resulting in a white or silvery reflection. It is sometimes called blister variegation. ''
Pilea ''Pilea'', with 600–715 species, is the largest genus of flowering plants in the nettle family Urticaceae. It is distributed throughout the tropics, subtropics, and warm temperate regions (with the exception of Australia and New Zealand). Des ...
'' cadierei (aluminum plant) is an example of a house plant that shows this effect. Leaves of most ''
Cyclamen ''Cyclamen'' ( or ) is a genus of 23 species of perennial flowering plants in the family Primulaceae. ''Cyclamen'' species are native to Europe and the Mediterranean Basin east to the Caucasus and Iran, with one species in Somalia. They grow ...
'' species show such patterned variegation, varying between plants, but consistent within each plant. Another type of reflective variegation is caused by hairs on parts of the leaf, which may be coloured differently from the leaf. This is found in various ''
Begonia ''Begonia'' is a genus of perennial flowering plants in the family Begoniaceae. The genus contains more than 2,000 different plant species. The Begonias are native to moist subtropical and tropical climates. Some species are commonly grown ind ...
'' species and garden hybrids. Sometimes venal variegation occurs – the veins of the leaf are picked out in white or yellow. This is due to lack of green tissue above the veins. It can be seen in some
aroids The Araceae are a family of monocotyledonous flowering plants in which flowers are borne on a type of inflorescence called a spadix. The spadix is usually accompanied by, and sometimes partially enclosed in, a spathe (or leaf-like bract). Also k ...
. The blessed milk thistle, ''Silybum marianum'', is a plant in which another type of venal variegation occurs, but in this case it is due to a blister variegation occurring along the veins.


Pigmentary

A common cause of variegation is the masking of green
pigment A pigment is a colored material that is completely or nearly insoluble in water. In contrast, dyes are typically soluble, at least at some stage in their use. Generally dyes are often organic compounds whereas pigments are often inorganic compo ...
by other pigments, such as
anthocyanin Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical compo ...
s. This often extends to the whole leaf, causing it to be reddish or purplish. On some plants however, consistent zonal markings occur; such as on some
clover Clover or trefoil are common names for plants of the genus ''Trifolium'' (from Latin ''tres'' 'three' + ''folium'' 'leaf'), consisting of about 300 species of flowering plants in the legume or pea family Fabaceae originating in Europe. The genus ...
s,
bromeliads The Bromeliaceae (the bromeliads) are a family of monocot flowering plants of about 80 genera and 3700 known species, native mainly to the tropical Americas, with several species found in the American subtropics and one in tropical west Africa, ...
, certain ''
Pelargonium ''Pelargonium'' () is a genus of flowering plants that includes about 280 species of perennials, succulents, and shrubs, commonly called geraniums, pelargoniums, or storksbills. '' Geranium'' is also the botanical name and common name of a separ ...
'' and ''
Oxalis ''Oxalis'' ( (American English) or (British English)) is a large genus of flowering plants in the wood-sorrel family Oxalidaceae, comprising over 550 species. The genus occurs throughout most of the world, except for the Polar region, polar are ...
'' species. On others, such as the commonly grown forms of
Coleus ''Coleus'' is a genus of annual or perennial herbs or shrubs, sometimes succulent, sometimes with a fleshy or tuberous rootstock, found in the Old World tropics and subtropics. The relationship among the genera ''Coleus'', '' Solenostemon'' and ...
, the variegation can vary widely within a population. In ''
Nymphaea lotus ''Nymphaea lotus'', the white Egyptian lotus, tiger lotus, white lotus or Egyptian white water-lily, is a flowering plant of the family Nymphaeaceae. Distribution It grows in various parts of East Africa and Southeast Asia. The ''Nymphaea lot ...
'', the tiger lotus, leaf variegations appear under intense illumination.


Pathological

Virus A virus is a submicroscopic infectious agent that replicates only inside the living cells of an organism. Viruses infect all life forms, from animals and plants to microorganisms, including bacteria and archaea. Since Dmitri Ivanovsky's 1 ...
infections may cause patterning to appear on the leaf surface. The patterning is often characteristic of the infection. Examples are the ''mosaic viruses'', which produce a mosaic-type effect on the leaf surface or the
citrus variegation virus Citrus variegation virus (CVV) is a plant pathogenic virus, a member of subgroup 2 of ilarviruses in the family ''Bromoviridae'', is the causal agent of infectious variegation, a disease occurring all over the world, causing problems for productio ...
(CVV). Recently a virus disease, Hosta Virus X (HVX) has been identified that causes mottled leaf coloring in
hostas ''Hosta'' (, syn. ''Funkia'') is a genus of plants commonly known as hostas, plantain lilies and occasionally by the Japanese name gibōshi. Hostas are widely cultivated as shade-tolerant foliage plants. The genus is currently placed in the fa ...
. At first, diseased plants were propagated and grown for their mottled foliage, at the risk of infecting other healthy hostas. While these diseases are usually serious enough that the gardener would not grow affected plants, there are a few affected plants that can survive indefinitely, and are attractive enough to be grown for ornament; e.g. some variegated
Abutilon ''Abutilon'' is a large genus of flowering plants in the mallow family, Malvaceae. It is distributed throughout the tropics and subtropicsNutrient A nutrient is a substance used by an organism to survive, grow, and reproduce. The requirement for dietary nutrient intake applies to animals, plants, fungi, and protists. Nutrients can be incorporated into cells for metabolic purposes or excret ...
deficiency symptoms may cause a temporary or variable yellowing in specific zones on the leaf.
Iron Iron () is a chemical element with symbol Fe (from la, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 of the periodic table. It is, by mass, the most common element on Earth, right in f ...
and
magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...
deficiencies are common causes of this.
Transposable element A transposable element (TE, transposon, or jumping gene) is a nucleic acid sequence in DNA that can change its position within a genome, sometimes creating or reversing mutations and altering the cell's genetic identity and genome size. Transp ...
s can cause colour variegation.


Defensive masquerade

It has been suggested that some patterns of leaf variegation may be part of a defensive "masquerade strategy." In this, leaf variegation may appear to a leaf mining insect that the leaf is already infested, and this may reduce parasitization of the leaf by leaf miners.


Nomenclature

By convention, the italicised term ''variegata'' as the second part of the Latin
binomial Binomial may refer to: In mathematics *Binomial (polynomial), a polynomial with two terms * Binomial coefficient, numbers appearing in the expansions of powers of binomials *Binomial QMF, a perfect-reconstruction orthogonal wavelet decomposition ...
name, indicates a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
found in the wild with variegation (''
Aloe variegata ''Gonialoe variegata'' ( syn. ''Aloe variegata''), also known as tiger aloe and partridge-breasted aloe, is a species of flowering plant in the family Asphodelaceae. It is an evergreen succulent perennial indigenous to South Africa and Namibia. ...
''). The much more common, non-italicised, inclusion of 'Variegata' as the third element of a name indicates a variegated
cultivar A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture, ...
of an unvariegated parent (''
Aucuba japonica ''Aucuba japonica'', commonly called spotted laurel, Japanese laurel, Japanese aucuba or gold dust plant (U.S.), is a shrub () native to rich forest soils of moist valleys, thickets, by streams and near shaded moist rocks in China, Korea, and Jap ...
'' 'Variegata'). However, not all variegated plants have this Latin tag, for instance many cultivars of ''
Pelargonium ''Pelargonium'' () is a genus of flowering plants that includes about 280 species of perennials, succulents, and shrubs, commonly called geraniums, pelargoniums, or storksbills. '' Geranium'' is also the botanical name and common name of a separ ...
'' have some zonal variegation in their leaves. Other types of variegation may be indicated, e.g. ''
Daphne odora ''Daphne odora'', winter daphne, is a species of flowering plant in the family Thymelaeaceae, native to China, later spread to Japan and Korea. It is an evergreen shrub, grown for its very fragrant, fleshy, pale-pink, tubular flowers, each with f ...
'' 'Aureomarginata' has yellow edging on its leaves.


Garden plants

Variegated plants have long been valued by gardeners, as the usually lighter-coloured variegation can 'lift' what would otherwise be blocks of solid green foliage. Many gardening societies have specialist variegated plants groups, such as the
Hardy Plant Society The Hardy Plant Society is a British charity that promotes the cultivation of hardy herbaceous plants. The society was founded in 1957 by a group of gardeners and nurserymen. It has approximately 7,500 members and provides information about fa ...
's Variegated Plant Special Interest Group in the UK. Several gardening books which deal exclusively with variegated plants are available.


Variegated plants at auction

In August 2020 a variegated ''
Rhaphidophora tetrasperma ''Rhaphidophora tetrasperma'', the mini monstera, is a species of flowering plant in the family Araceae, genus '' Rhaphidophora''. It is native to Southern Thailand and to Malaysia.Rhaphidophora tetrasperma ''Rhaphidophora tetrasperma'', the mini monstera, is a species of flowering plant in the family Araceae, genus '' Rhaphidophora''. It is native to Southern Thailand and to Malaysia.center, 350px, The rare rainforest plant, '' Cryptocarya williwilliana'', showing leaf venation and variegated leaves


References

{{reflist Plant physiology Chimerism