Gall

This article is about the abnormal plant growths. For other uses, see Gall (disambiguation).
Kalanchoë infected with crown gall using Agrobacterium tumefaciens.
A detail photo of a crown gall on a Kalanchoë infected with Agrobacterium tumefaciens.
Gall of Japanagromyza inferna in Centrosema virginianum L.
Galls can also appear on skeletal animals and in the fossil record. Two galls with perforations on a crinoid stem (Apiocrinites negevensis) from the Middle Jurassic of southern Israel.

Galls or cecidia are a kind of swelling growth on the external tissues of plants or animals. Plant galls are abnormal outgrowths[1] of plant tissues, similar to benign tumors or warts in animals. They can be caused by various parasites, from fungi and bacteria, to insects and mites. Plant galls are often highly organized structures and because of this the cause of the gall can often be determined without the actual agent being identified. This applies particularly to some insect and mite plant galls. The study of plant galls is known as cecidology.

In human pathology, a gall is a raised sore on the skin, usually caused by chafing or rubbing.[2]

Causes of plant galls

Insects and mites

Insect galls are the highly distinctive plant structures formed by some herbivorous insects as their own microhabitats. They are plant tissue which is controlled by the insect. Galls act as both the habitat and food source for the maker of the gall. The interior of a gall can contain edible nutritious starch and other tissues. Some galls act as "physiologic sinks", concentrating resources in the gall from the surrounding plant parts.[3] Galls may also provide the insect with physical protection from predators.[4][5]

Insect galls are usually induced by chemicals injected by the larvae or the adults of the insects into the plants, and possibly mechanical damage. After the galls are formed, the larvae develop inside until fully grown, when they leave. In order to form galls, the insects must seize the time when plant cell division occurs quickly: the growing season, usually spring in temperate climates, but which is extended in the tropics.

The meristems, where plant cell division occurs, are the usual sites of galls, though insect galls can be found on other parts of the plant, such as the leaves, stalks, branches, buds, roots, and even flowers and fruits. Gall-inducing insects are usually species-specific and sometimes tissue-specific on the plants they gall.

Inductor insects

Gall-inducing insects include gall wasps, gall midges, gall flies, Agromyzidae aphids (such as Melaphis chinensis, Pemphigus spyrothecae, and Pemphigus betae), scale insects, and psyllids. Various galls produced by insects and mites are listed below:

Fungi

Many rust fungi induce gall formation, including Western gall rust, which infects a variety of pine trees and cedar-apple rust. Galls are often seen in Millettia pinnata leaves and fruits. Leaf galls appear like tiny clubs; however, flower galls are globose. Exobasidium often induces spectacular galls on its hosts.

It is worth noting that the fungus Ustilago esculenta associated with Zizania latifolia, a wild rice, produces an edible gall highly valued as a food source in the Zhejiang and Jiangsu provinces of China.[6]

Bacteria and viruses

Agrobacterium tumefaciens and Pseudomonas savastanoi are examples of gall-causing bacteria. Gall forming Virus was found on rice plants in central Thailand in 1979 and named rice gall dwarf. Symptoms consisted of gall formation along leaf blades and sheaths, dark green discoloration, twisted leaf tips and reduced numbers of tillers. Some plants died in the glasshouse in later stages of infection. The causal agent was transmitted by Nephotettix nigropictus after an incubation of 2 weeks. Polyhedral particles of 65 nm diameter in the cytoplasm of phloem cells were always associated with the disease. No serologic relationship was found between this virus and that of rice dwarf.

Nematodes

Nematodes are microscopic worms that live in the soil. Some nematodes (Meloidogyne species or root-knot nematodes) cause galls on the roots of susceptible plants. The galls are small, individual and beadlike in some hosts. In other plant species galls may be massive accumulations of fleshy tissue more than an inch in diameter. Some ectoparasitic nematodes (nematodes that live outside the plant in the soil), such as sting and stubby-root nematodes, may cause root tips to swell.Nitrogen-fixing bacteria (Rhizobium species) cause swellings on the roots of most legumes (such as clover, peas and beans). These swellings, called nodules, are easily distinguished from root-knot galls by differences in how they are attached to the root and their contents. Nodules are loosely attached to the root, while root-knot galls originate from infection at the center of the root, so they are an integral part of the root. In addition, fresh Rhizobium nodules have a milky pink-to-brown liquid inside them, while root-knot galls have firmer tissues and contain female root-knot nematodes (creamy white beads less than 1/32 inch in diameter) inside the gall tissues.

Other plants

Mistletoe can form galls on its hosts.

Uses

Galls are rich in resins and tannic acid and have been used in the manufacture of permanent inks (such as iron gall ink) and astringent ointments, in dyeing, and in tanning. A high-quality ink has long been made from the Aleppo gall, found on oaks in the Middle East. The Talmud, ([7]) records using gallnuts as part of the tanning process as well as a dye-base for ink.

The larvae in galls are useful for a survival food and fishing bait; see the Indigenous Australian foods Bush coconut and Mulga apple. Nutgalls also produce purpurogallin.

The gall of Rhus chinensis, Galla chinensi, has long been considered to possess many medicinal properties.[8]

See also

Footnotes

  1. Merriam-Webster Online Dictionary, accessed Nov. 16, 2007 ("an abnormal outgrowth of plant tissue usually due to insect or mite parasites or fungi and sometimes forming an important source of tannin")
  2. medical-dictionary.thefreedictionary.com
  3. Larson, K. C., and T. G. Whitham. 1991. Manipulation of food resources by a gall-forming aphid: the physiology of sink-source interactions. Oecologia 88, P.15 – 21.
  4. Weis, A. E., and A. Kapelinski. 1994. Variable selection on Eurosta’s gall size. II. A path analysis of the ecological factors behind selection. Evolution 48, P.734 – 745.
  5. Graham N. Stone and Karsten Schonrogge (2003) The adaptive significance of insect gall morphology. TRENDS in Ecology and Evolution 18(10):512-522
  6. Terrell, E.E. and L.R. Batra. Zizania latifolia and Ustilago esculenta, a grass-fungus association. Economic Botany 36(3):274-285.
  7. Bavli, tractate Gittin:19a
  8. Zhang J, Li L, Kim SH, Hagerman AE, Lü J. 2009. "Anti-cancer, anti-diabetic and other pharmacologic and biological activities of penta-galloyl-glucose." Pharm Res 26: 2066–2080.

Further reading

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