Growth and Reproduction

The vegetative body of a fungus is called a thallus and can be unicellular or multicellular. Some fungi are dimorphic because they can go from being unicellular to multicellular depending on environmental conditions. Unicellular fungi are generally referred to as yeasts.Saccharomyces cerevisiae (baker’s yeast) and Candida species (the agents of thrush, a common fungal infection) are examples of unicellular fungi.

Most fungi are multicellular organisms. They display two distinct morphological stages: vegetative and reproductive. The vegetative stage is characterized by a tangle of slender thread-like structures called hyphae (singular, hypha), whereas the reproductive stage can be more conspicuous. A mass of hyphae is called a mycelium (Figure 13.22). It can grow on a surface, in soil or decaying material, in a liquid, or even in or on living tissue. Although individual hypha must be observed under a microscope, the mycelium of a fungus can be very large with some species truly being “the fungus humongous.” The giant Armillaria ostoyae (honey mushroom) is considered the largest organism on Earth, spreading across over 2,000 acres of underground soil in eastern Oregon; it is estimated to be at least 2,400 years old.

Most fungal hyphae are divided into separate cells by end walls called septa (singular, septum). In most divisions (like plants, fungal phyla are called divisions by tradition) of fungi, tiny holes in the septa allow for the rapid flow of nutrients and small molecules from cell to cell along the hyphae. They are described as perforated septa. The hyphae in bread molds (which belong to the division Zygomycota) are not separated by septa. They are formed of large cells containing many nuclei, an arrangement described as coenocytic hyphae.

Fungi thrive in environments that are moist and slightly acidic, and can grow with or without light. They vary in their oxygen requirements. Most fungi are obligate aerobes, requiring oxygen to survive. Other species, such as the Chytridiomycota that reside in the rumen of cattle, are obligate anaerobes, meaning that they cannot grow and reproduce in an environment with oxygen. Yeasts are intermediate: They grow best in the presence of oxygen but can use fermentation in the absence of oxygen. The alcohol produced from yeast fermentation is used in wine and beer production, and the carbon dioxide they produce carbonates beer and sparkling wine, and makes bread rise.

Fungi can reproduce sexually or asexually. In both sexual and asexual reproduction, fungi produce spores that disperse from the parent organism by either floating in the wind or hitching a ride on an animal. Fungal spores are smaller and lighter than plant seeds, but they are not usually released as high in the air. The giant puffball mushroom bursts open and releases trillions of spores: The huge number of spores released increases the likelihood of spores landing in an environment that will support growth (Figure 13.23).

 

How Fungi Obtain Nutrition

Like animals, fungi are heterotrophs: They use complex organic compounds as a source of carbon rather than fixing carbon dioxide from the atmosphere, as some bacteria and most plants do. In addition, fungi do not fix nitrogen from the atmosphere. Like animals, they must obtain it from their diet. However, unlike most animals that ingest food and then digest it internally in specialized organs, fungi perform these steps in the reverse order. Digestion precedes ingestion. First, exoenzymes, enzymes that catalyze reactions on compounds outside of the cell, are transported out of the hyphae where they break down nutrients in the environment. Then, the smaller molecules produced by the external digestion are absorbed through the large surface areas of the mycelium. As with animal cells, the fungal storage polysaccharide is glycogen rather than starch, as found in plants.

Fungi are mostly saprobes, organisms that derive nutrients from decaying organic matter. They obtain their nutrients from dead or decomposing organic matter, mainly plant material. Fungal exoenzymes are able to break down insoluble polysaccharides, such as the cellulose and lignin of dead wood, into readily absorbable glucose molecules. Decomposers are important components of ecosystems, because they return nutrients locked in dead bodies to a form that is usable for other organisms. This role is discussed in more detail later. Because of their varied metabolic pathways, fungi fulfill an important ecological role and are being investigated as potential tools in bioremediation. For example, some species of fungi can be used to break down diesel oil and polycyclic aromatic hydrocarbons. Other species take up heavy metals such as cadmium and lead.

Importance to Ecosystems

Food webs would be incomplete without organisms that decompose organic matter and fungi are key participants in this process. Decomposition allows for cycling of nutrients such as carbon, nitrogen, and phosphorus back into the environment so they are available to living things, rather than being trapped in dead organisms. Fungi are particularly important because they have evolved enzymes to break down cellulose and lignin, components of plant cell walls that few other organisms are able to digest, releasing their carbon content.

Fungi are also involved in ecologically important coevolved symbioses, both mutually beneficial and pathogenic with organisms from the other kingdoms. Mycorrhiza, a term combining the Greek roots myco meaning fungus and rhizo meaning root, refers to the association between vascular plant roots and their symbiotic fungi. Somewhere between 80–90 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. Ectomycorrhizae (“outside” mycorrhiza) depend on fungi enveloping the roots in a sheath (called a mantle) and a net of hyphae that extends into the roots between cells. In a second type, the Glomeromycota fungi form arbuscular mycorrhiza. In these mycorrhiza, the fungi form arbuscles, a specialized highly branched hypha, which penetrate root cells and are the sites of the metabolic exchanges between the fungus and the host plant. Orchids rely on a third type of mycorrhiza. Orchids form small seeds without much storage to sustain germination and growth. Their seeds will not germinate without a mycorrhizal partner (usually Basidiomycota). 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.

Lichens blanket many rocks and tree bark, displaying a range of colors and textures. Lichens are important pioneer organisms that colonize rock surfaces in otherwise lifeless environments such as are created by glacial recession. The lichen is able to leach nutrients from the rocks and break them down in the first step to creating soil. Lichens are also present in mature habitats on rock surfaces or the trunks of trees. They are an important food source for caribou. Lichens are not a single organism, but rather a fungus (usually an Ascomycota or Basidiomycota species) living in close contact with a photosynthetic organism (an alga or cyanobacterium). The body of a lichen, referred to as a thallus, is formed of hyphae wrapped around the green partner. The photosynthetic organism provides carbon and energy in the form of carbohydrates and receives protection from the elements by the thallus of the fungal partner. 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.

Fungi have evolved mutualistic associations with numerous arthropods. 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. Fungi are cultivated in these gardens, digesting the cellulose that the ants cannot break down. Once smaller sugar molecules are produced and consumed by the fungi, they 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.

Importance to Humans

Although we often think of fungi as organisms that cause diseases and rot food, fungi are important to human life on many levels. As we have seen, they influence the well-being of human populations on a large scale because they help nutrients cycle in ecosystems. They have other ecosystem roles as well. For example, as animal pathogens, fungi help to control the population of damaging pests. These fungi are very specific to the insects they attack and do not infect other animals or plants. The potential to use fungi as microbial insecticides is being investigated, with several species already on the market. For example, the fungus Beauveria bassiana is a pesticide that is currently being tested as a possible biological control for the recent spread of emerald ash borer. It has been released in Michigan, Illinois, Indiana, Ohio, West Virginia, and Maryland.

The mycorrhizal relationship between fungi and plant roots is essential for the productivity of farmland. Without the fungal partner in the root systems, 80–90% of trees and grasses would not survive. Mycorrhizal fungal inoculants are available as soil amendments from gardening supply stores and promoted by supporters of organic agriculture.

We also eat some types of fungi. Mushrooms figure prominently in the human diet. Morels, shiitake mushrooms, chanterelles, and truffles are considered delicacies (Figure 13.27). The humble meadow mushroom, Agaricus campestris, appears in many dishes. Molds of the genus Penicillium ripen many cheeses. They originate in the natural environment such as the caves of Roquefort, France, where wheels of sheep milk cheese are stacked to capture the molds responsible for the blue veins and pungent taste of the cheese.

 

Fermentation—of grains to produce beer, and of fruits to produce wine—is an ancient art that humans in most cultures have practiced for millennia. Wild yeasts are acquired from the environment and used to ferment sugars into CO₂ and ethyl alcohol under anaerobic conditions. It is now possible to purchase isolated strains of wild yeasts from different wine-making regions. Pasteur was instrumental in developing a reliable strain of brewer’s yeast, Saccharomyces cerevisiae, for the French brewing industry in the late 1850s. It was one of the first examples of biotechnology patenting. Yeast is also used to make breads that rise. The carbon dioxide they produce is responsible for the bubbles produced in the dough that become the air pockets of the baked bread.

Many secondary metabolites of fungi are of great commercial importance. Antibiotics are naturally produced by fungi to kill or inhibit the growth of bacteria, and limit competition in the natural environment. Valuable drugs isolated from fungi include the immunosuppressant drug cyclosporine (which reduces the risk of rejection after organ transplant), the precursors of steroid hormones, and ergot alkaloids used to stop bleeding. In addition, as easily cultured eukaryotic organisms, some fungi are important model research organisms including the red bread mold Neurospora crassa and the yeast, S. cerevisiae.