By the end of this reading you should be able to:

• Explain the roles fungi can play in ecosystems
• Describe mutualistic relationships of fungi with plant roots and photosynthetic organisms
• Describe the beneficial relationship between some fungi and insects

Fungi play a crucial role in the balance of ecosystems. They colonize most habitats on Earth, preferring dark, moist conditions. They can thrive in seemingly hostile environments, such as the tundra, thanks to a most successful symbiosis with photosynthetic organisms like algae to produce lichens. Fungi are not obvious in the way large animals or tall trees appear. Yet, like bacteria, they are the major decomposers of nature. With their versatile metabolism, fungi break down organic matter, which would not otherwise be recycled.

Although fungi are primarily associated with humid and cool environments that provide a supply of organic matter, they colonize a surprising diversity of habitats, from seawater to human skin and mucous membranes. Chytrids are found primarily in aquatic environments. Other fungi, such as Coccidioides immitis, which causes pneumonia when its spores are inhaled, thrive in the dry and sandy soil of the southwestern United States. Fungi that parasitize coral reefs live in the ocean. However, most members of the Kingdom Fungi grow on the forest floor, where the dark and damp environment is rich in decaying debris from plants and animals. In these environments, fungi play a major role as decomposers and recyclers, supplying other groups of organisms with some of the nutrients they need to live.

The food web would be incomplete without organisms that decompose organic matter. Some elements—such as nitrogen and phosphorus—are required in large quantities by biological systems, and yet are not abundant in the environment. The action of fungi releases these elements from decaying matter, making them available to other living organisms (Fig. 1). Trace elements present in low amounts in many habitats are essential for growth and would remain tied up in rotting organic matter if fungi and bacteria did not return them to the environment via their metabolic activity.

Symbiosis is the ecological interaction between two organisms that live together. The definition does not describe the quality of the interaction. When both members of the association benefit, the symbiotic relationship is called a mutualism. Fungi form mutualistic associations with many types of organisms, including cyanobacteria, algae, plants, and animals.

One of the most remarkable associations between fungi and plants is the establishment of mycorrhizae. Mycorrhiza, which comes from the Greek words myco meaning fungus and rhizo meaning root, refers to the association between vascular plant roots and their symbiotic fungi. Somewhere between 80 and 90 percent of all plant species have mycorrhizal partners. In a mycorrhizal association, the fungal mycelia use their extensive network of hyphae and large surface area in contact with the soil to channel water and minerals from the soil into the plant. In exchange, the plant supplies the products of photosynthesis to fuel the metabolism of the fungus.

There are a number of types of mycorrhizal. Ectomycorrhizae (“outside” mycorrhiza) depend on fungi enveloping the roots in a sheath (called a mantle) and a Hartig net of hyphae that extends into the roots between cells (Fig. 3a).  Arbuscular mycorrhiza sometimes called endomycorrhizae (Fig. 3b),  form arbuscules that penetrate root cells and are the site of the metabolic exchanges between the fungus and the host plant (Fig 3 & 4). Orchids rely on a third type of mycorrhiza. Orchids are epiphytes that form small seeds without much storage to sustain germination and growth. Their seeds will not germinate without a mycorrhizal partner. After nutrients in the seed are depleted, fungal symbionts support the growth of the orchid by providing necessary carbohydrates and minerals. Some orchids continue to be mycorrhizal throughout their lifecycle.

Other examples of fungus–plant mutualism include the endophytes: fungi that live inside tissue without damaging the host plant. Endophytes release toxins that repel herbivores or confer resistance to environmental stress factors, such as infection by microorganisms, drought, or heavy metals in soil.

A well-accepted theory proposes that fungi were instrumental in the evolution of the root system in plants and contributed to the success of flowering plants. Fossil records indicate that fungi preceded plants on dry land. The first association between fungi and photosynthetic organisms on land involved moss-like plants and endophytes. These early associations developed before roots appeared in plants. Slowly, the benefits of the endophyte and rhizoid interactions for both partners led to present-day mycorrhizae; up to about 90 percent of today’s vascular plants have associations with fungi in their rhizosphere. The fungi involved in mycorrhizae display many characteristics of primitive fungi; they produce simple spores, show little diversification, do not have a sexual reproductive cycle, and cannot live outside of a mycorrhizal association. The plants benefited from the association because mycorrhizae allowed them to move into new habitats because of increased uptake of nutrients, and this gave them a selective advantage over plants that did not establish symbiotic relationships.

Lichens

Lichens are not a single organism, but rather an example of a mutualism, in which a fungus lives in close contact with a photosynthetic organism (a eukaryotic alga or a prokaryotic cyanobacterium). Generally, neither the fungus nor the photosynthetic organism can survive alone outside of the symbiotic relationship. The body of a lichen, referred to as a thallus, is formed from fungal hyphae wrapped around the photosynthetic partner (Fig. 6). The photosynthetic organism provides carbon and energy in the form of carbohydrates. Some cyanobacteria fix nitrogen from the atmosphere, contributing nitrogenous compounds to the association. In return, the fungus supplies minerals and protection from dryness and excessive light by encasing the algae in its mycelium. The fungus also attaches the symbiotic organism to the substrate.

The thallus of lichens grows very slowly, expanding its diameter a few millimeters per year. Both the fungus and the alga participate in the formation of dispersal units, soredia, for reproduction. These structures are clusters of algal cells surrounded by mycelia and are dispersed by wind and water to form new lichens.

Lichens are extremely sensitive to air pollution, especially to abnormal levels of nitrogen and sulfur. The U.S. Forest Service and National Park Service can monitor air quality by measuring the relative abundance and health of the lichen population in an area. Lichens fulfill many ecological roles. Caribou and reindeer eat lichens, and they provide cover for small invertebrates that hide in the mycelium. In the production of textiles, weavers used lichens to dye wool for many centuries until the advent of synthetic dyes.

Lichens are used to monitor the quality of air. Read more on this site (http://openstaxcollege.org/l/lichen_monitrng) from the United States Forest Service.

Fungi have evolved mutualisms with numerous insects in Phylum Arthropoda: jointed, legged invertebrates. Arthropods depend on the fungus for protection from predators and pathogens, while the fungus obtains nutrients and a way to disseminate spores into new environments. The association between species of Basidiomycota and scale insects is one example. The fungal mycelium covers and protects the insect colonies. The scale insects foster a flow of nutrients from the parasitized plant to the fungus. In a second example, leaf-cutting ants of Central and South America literally farm fungi. They cut disks of leaves from plants and pile them up in gardens (Fig. 7). Fungi are cultivated in these disk gardens, digesting the cellulose in the leaves that the ants cannot break down. Once smaller sugar molecules are produced and consumed by the fungi, the fungi, in turn, become a meal for the ants. The insects also patrol their garden, preying on competing fungi. Both ants and fungi benefit from the association. The fungus receives a steady supply of leaves and freedom from competition, while the ants feed on the fungi they cultivate.

Animal dispersal is important for some fungi because an animal may carry spores considerable distances from the source. Fungal spores are rarely completely degraded in the gastrointestinal tract of an animal, and many are able to germinate when they are passed in the feces. Some dung fungi actually require passage through the digestive system of herbivores to complete their lifecycle. The black truffle—a prized gourmet delicacy—is the fruiting body of an underground mushroom. Almost all truffles are ectomycorrhizal, and are usually found in close association with trees. Animals eat truffles and disperse the spores. In Italy and France, truffle hunters use female pigs to sniff out truffles. Female pigs are attracted to truffles because the fungus releases a volatile compound closely related to a pheromone produced by male pigs.

Fungi have colonized nearly all environments on Earth, but are frequently found in cool, dark, moist places with a supply of decaying material. Fungi are saprobes that decompose organic matter. Many successful mutualistic relationships involve a fungus and another organism. Many fungi establish complex mycorrhizal associations with the roots of plants. Some ants farm fungi as a supply of food. Lichens are a symbiotic relationship between a fungus and a photosynthetic organism, usually an alga or cyanobacterium. The photosynthetic organism provides energy derived from light and carbohydrates, while the fungus supplies minerals and protection. Some animals that consume fungi help disseminate spores over long distances.