Variegation

This article is about variegation in botany. For histology, see Variegation (histology).

Variegation of fruits and wood in Ficus carica Panascè, a bicolor (yellow-green) common fig cultivar. This Italian cultivar is a chimera

Variegation is the appearance of differently coloured zones in the leaves, and sometimes the stems, of plants. This may be due to a number of causes. Some variegation is attractive and ornamental and gardeners tend to preserve these. The term is also sometimes used to refer to colour zonation in flowers, minerals, and the skin, fur, feathers or scales of animals.

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, 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.

Variegation in holly leaves

Reflective variegation in Pilea cadierei.

Some variegation is due to visual effects caused by reflection of light from the leaf surface. This can happen when an air layer is located just under the epidermis resulting in a white or silvery reflection. It is sometimes called blister variegation.

Pilea (aluminum plant) is an example of a house plant that shows this effect. Leaves of most Cyclamen 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 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 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

Leaves of Red Clover (Trifolium pratense) have a typical v-shaped variegation.

A common cause of variegation is the masking of green pigment by other pigments, such as anthocyanins. 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 clovers, bromeliads, certain Pelargonium and Oxalis species. On others, such as the commonly grown forms of Coleus, the variegation can vary widely within a population.

In Nymphaea lotus, the tiger lotus, leaf variegations appear under intense illumination.

Pathological

Virus 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 (CVV). Hosta Virus X (HVX) has caused such dramatic and attractive mosaic-type variegations that, when the disease was new to the United States, affected plants were mistakenly thought to be new mutations and were registered as new cultivars.

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 varieties.

Nutrient deficiency symptoms may cause a temporary or variable yellowing in specific zones on the leaf. Iron and magnesium deficiencies are common causes of this.

Transposons ("jumping genes") can also cause colour variegation2. Barbara McClintock won a Nobel Prize for figuring out how transposons caused colour variegation in maize1. References 1.http://www.nature.com/scitable/topicpage/barbara-mcclintock-and-the-discovery-of-jumping-34083# 2. Suzuki et al. 2014. Transposon-mediated mutation of CYP76AD3 affects betalain synthesis and produces variegated flowers in four o'clock (Mirabilis jalapa).J Plant Physiol. 2014 Nov 1;171(17):1586-90. doi: 10.1016/j.jplph.2014.07.010. Epub 2014 Jul 25.

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 an insect leaf miner 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 name, indicates a species found in the wild with variegation (Aloe variegata). The much more common, non-italicised, inclusion of 'Variegata' as the third element of a name indicates a variegated cultivar of an unvariegated parent (Aucuba japonica 'Variegata'). However, not all variegated plants have this Latin tag, for instance many cultivars of Pelargonium have some zonal variegation in their leaves. Other types of variegation may be indicated, e.g. Daphne odora 'Aureomarginata' has yellow edging on its leaves.

Garden plants

Variegated foliage used in an English garden. Plants shown include Aralia elata 'Aureovariegata' and Carex ornithopoda 'Variegata'

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's Variegated Plant Special Interest Group in the UK. Several gardening books which deal exclusively with variegated plants are available.

The rare rainforest plant, Cryptocarya williwilliana showing leaf venation and variegated leaves

References

• Variegated Plants - article from Union County College, New Jersey.
• BBC: Earth News - article suggesting that some variegation is to feign disease.