Lichens

Lichens are an intimate symbiosis, in which two species live together as a type of composite organism. Lichens are an obligate mutualism between a fungus mycobiont and an alga or blue-green bacterium phycobiont.

Each lichen mutualism is highly distinctive, and can be identified on the basis of its size, shape, color, and biochemistry. Even though lichens are not true "species" in the conventional meaning of the word, lichenologists have developed systematic and taxonomic treatments of these mutualisms.

The fungal partner in the lichen mutualism gains important benefits through access to photosynthetic products of the alga or blue-green bacterium. The phycobiont profits from the availability of a relatively moist and protected habitat, and greater access to inorganic nutrients.

The most common fungi in lichens are usually species of Ascomycetes, or a few Basidiomycetes. The usual algal partners are either species of green algae Chlorophyta or blue-green bacteria of the family Cyanophyceae. In general, the fungal partner cannot live without its phycobiont, but the algae is often capable of living freely in moist soil or water. The largest lichens can form a thallus up to 3 ft (1 m) long, although most lichens are smaller than a few inches or centimeters in length. Lichens can be very colorful, ranging from bright reds and oranges, to yellows and greens, and white, gray, and black hues.

Most lichens grow very slowly. Lichens in which the phycobiont is a blue-green bacterium have the ability to fix nitrogen gas into ammonia. Some lichens can commonly reach ages of many centuries, especially species living in highly stressful environments, such as alpine or arctic tundra.

Lichens can grow on diverse types of substrates. Some species grow directly on rocks, some on bare soil, and others on the bark of tree trunks and branches. Lichens often grow under exposed conditions that are frequently subjected to periods of drought, and sometimes to extremes of hot and cold. Lichen species vary greatly in their tolerance of severe environmental conditions. Lichens generally respond to environmental extremes by becoming dormant, and then quickly becoming metabolically active again when they experience more benign conditions.

Lichens are customarily divided into three growth forms, although this taxonomy is one of convenience, and is not ultimately founded on systematic relationships. Crustose lichens form a thallus that is closely appressed to the surface upon which they are growing. Foliose lichens are only joined to their substrate by a portion of their thallus, and they are somewhat leaf-like in appearance. Fruticose lichens rise above their substrate, and are much branched and bushy in appearance.

Most lichens regenerate asexually as lichen symbioses, and not by separate reproduction of their mycobiont and phycobiont. Reproduction is most commonly accomplished by small, specialized fragments of thallus known as soredia, consisting of fungal tissue enclosing a small number of algal cells. The soredia generally originate within the parent thallus, then grow out through the surface of the thallus, and detach as small bits of tissue that are dispersed by the wind or rain. If the dispersing soredium is fortunate enough to lodge in a favorable microenvironment, it develops into a new thallus, genetically identical to the parent.

Because they are capable of colonizing bare rocks and other mineral substrates, lichens are important in soil formation during some ecological successions. For example, lichens are among the first organisms to colonize sites as they are released from glacial ice. In such situations, lichens can be important in the initial stages of nitrogen accumulation and soil development during post-glacial primary succession.

Lichens are important forage for some species of animals. The best known example of this relationship involves the northern species of deer known as caribou or reindeer (Rangifer tarandus) and the so-called reindeer lichens (Cladina spp.) that are one of their most important foods, especially during winter.

Some species of lichens are very sensitive to air pollutants. Consequently, urban environments are often highly impoverished in lichen species. Some ecologists have developed schemes by which the intensity of air pollution can be reliably assayed or monitored using the biological responses of lichens in their communities. Monitoring of air quality using lichens can be based on the health and productivity of these organisms in places variously stressed by toxic pollution. Alternatively, the chemical composition of lichens may be assayed, because their tissues can effectively take up and retain sulfur and metals from the atmosphere.

Some lichens are useful as a source of natural dyes. Pigments of some of the more colorful lichens, especially the orange, red, and brown ones, can be extracted by boiling and used to dye wool and other fibers. Other chemicals extracted from lichens include litmus, which was a commonly used acid-base indicator prior to the invention of the pH meter.

Some of the reindeer lichens, especially Cladina alpestris, are shaped like miniature shrubs and trees. Consequently, these plants are sometimes collected, dried, and dyed, and are used in "landscaping" the layouts for miniature railroads and architectural models.

In addition, lichens add significantly to the aesthetics of the ecosystems in which they occur. The lovely orange and yellow colors of Caloplaca and Xanthoria lichens add much to the ambience of rocky seashores and tundras. The intricate webs of filamentous Usnea lichens hanging in profusion from tree branches give a mysterious aspect to humid forests. These and other, less charismatic lichens are integral components of their natural ecosystems. These lichens are intrinsically important for this reason, as well as for the relatively minor benefits that they provide to humans.