Fungi

The fungi an often overlooked group of unicellular and multicellular organisms, have a profound
influence on ecology and human health. Along with bacteria, they are important decomposers
and disease-causing organisms. Fungi are found everywhere—from the tropics to the tundra and
in both terrestrial and aquatic environments. Fungi made it possible for plants to colonize land by
associating with rootless stems and aiding in the uptake of nutrients and water. Mushrooms and
toadstools grow rapidly under proper conditions and produce large numbers of fungal spores. A
single Armillaria fungus can cover 15 hectares underground and weigh 100 tons, making it the
largest organism in the world based on area. Some puffball fungi are almost a meter in diameter
and may contain 7 trillion spores—enough to circle the Earth’s equator! Yeasts are fungi that are
used to make bread and beer and some fungi are prized as food, but other fungi cause disease in
plants and animals. These eukaryotic microbes are particularly problematic because fungi are
animals’ closest relatives. Drugs that can kill fungi often have toxic effects on animals, including
humans.

Fungal Mycelium-This mycelium, composed of hyphae, is growing through leaves on the

forest floor
Mycologists, scientists who study fungi and fungus like protists, believe there may be as many as
1.5 million fungal species. Fungi exist either as single-celled yeasts or in multicellular form.
Their
reproduction may be either sexual or asexual, and they exhibit an unusual form of mitosis. They
are specialized to extract and absorb nutrients from their surroundings by secreting enzymes.
Other major features of fungi include a cell wall composed of chitin and reproduction by spores.
Recent phylogenetic analysis of DNA sequences and protein sequences indicates that fungi are
more closely related to animals than to plants. Fossils and molecular data indicate that animals
and fungi last shared a common ancestor.
The phylogenetic relationships among fungi have been the cause of much debate. Traditionally,
four fungal phyla were recognized, based primarily on characteristics of the cells undergoing
meiosis: Chytridiomycota (“chytrids”), Zygomycota (“zygomycetes”), Ascomycota
(“ascomycetes”), and Basidiomycota (“basidiomycetes”). The chytrids and zygomycetes are not
monophyletic. The understanding of fungal phylogeny is going through rapid changes, aided by
increasing molecular sequence data (table). In 2007, mycologists agreed on seven monophyletic
phyla: Microsporidia, Blastocladiomycota, Neocallimastigomycota, Chytridiomycota,
Glomeromycota, Basidiomycota, and Ascomycota. Blastomycetes and neocallimastigomycetes
were formerly grouped with the chytrids. Confounding our understanding of the complex
relationships among fungi is a growing body of evidence that horizontal gene transfer has
occurred in many fungal species.
Group Typical Key Characteristics Approximate
Examples Number
of Living Species
Chytridiomycota Allomyces Aquatic, flagellated fungi that 750
produce haploid gametes by sexual
reproduction or diploid zoospores by
asexual reproduction.
Zygomycota Rhizopus, Multinucleate hyphae lack septa, 1050
Pilobolus except for reproductive structures;
fusion of hyphae leads directly to
formation of a zygote in
zygosporangium, in which meiosis
occurs just before it germinates;
asexual reproduction is most
common.
Glomeromycota Glomus Form arbuscular mycorrhizae.
Multinucleate hyphae lack septa. 230
Reproduce asexually.

Fungi may be unicellular or filamentous

The Chytridomycota and Microsporidia are ancient fungi and, as such, are unicellular organisms.
During the evolution of the higher fungi, filamentous growth forms developed. The filaments are
strings of cells called hyphae. Some hyphae are continuous or branching tubes filled with
cytoplasm and multiple nuclei. The hyphae of the more advanced fungal groups (Ascomycota
and Basidiomycota) are typically made up of long chains of cells joined end-to-end and divided
by cross-walls called septa (singular, septum). The septa rarely form a complete barrier, except
when they separate the reproductive cells. Even fungi with septa can be considered one long cell.
Cytoplasm characteristically flows or streams freely throughout the hyphae, passing through
major pores in the septa. Because of this streaming, nutrients synthesized or obtained by any part
of the hyphae may be carried to the actively growing tips. As a result, fungal hyphae may grow
very rapidly when food and water are abundant and the temperature is optimum. For example,
you may have seen mushrooms suddenly appear in a lawn overnight after a rain in summer.

Fungal spores: Scanning electron micrograph of fungal spores from Aspergillus

The mycelium
A mass of connected hyphae is called a mycelium (plural, mycelia). be many meters long. The
mycelium grows into the soil, wood, or other material and digestion of the material begins
quickly. In two of the four major groups of fungi, reproductive structures formed of interwoven
hyphae, such as mushrooms, puffballs, and morels, are produced at certain stages of the life
cycle. These
structures expand rapidly because of rapid growth of the hyphae.
Cell walls with chitin
The cell walls of most fungi are formed of polysaccharides, including chitin. In contrast, cell
walls of plants and many protists contain cellulose, not chitin. Chitin is a modified cellulose
consisting
of linked glucose units to which nitrogen groups have been added; this polymer is then cross-
linked with proteins. Chitin is the same material that makes up the major portion of the hard
shells, or exoskeletons, of arthropods, a group of animals that includes insects and crustaceans.
Chitin is one of the shared traits that has led scientists to believe that fungi are more closely
related to animals than they are to plants.
Fungal cells may have more than one nucleus
Fungi are different from most animals and plants in that each cell (or hypha) can house one, two,
or more nuclei. A hypha that has only one nucleus is called monokaryotic; a cell with two
nuclei is dikaryotic. In a dikaryotic cell, the two haploid nuclei exist independently. Dikaryotic
hyphae have some genetic properties of diploids, because both genomes are transcribed.
Sometimes, many nuclei intermingle in the common cytoplasm of a fungal mycelium, which can
lack distinct cells. If a dikaryotic or multinucleate hypha has nuclei that are derived from two
genetically distinct individuals, the hypha is called heterokaryotic. Hyphae whose nuclei are
genetically similar to one another are called homokaryotic.
Mitosis is not followed by cell division
Mitosis in multicellular fungi differs from that in most other organisms. Because of the linked
nature of the cells, the cell itself is not (This word and the term mycology, the study of fungi, are
both derived from the Greek word mykes, meaning fungi.) The mycelium of a fungus constitutes
a system.
Fungi can reproduce both sexually and asexually
Many fungi are capable of producing both sexual and asexual spores. When a fungus reproduces
sexually, two haploid hyphae of compatible mating types may come together and fuse
Fungi are heterotrophs that absorb nutrients
All fungi obtain their food by secreting digestive enzymes into their surroundings and then
absorbing the organic molecules produced by this external digestion. The fungal body plan
reflects
this approach. Unicellular fungi have the greatest surface area-to-volume ratio of any fungus,
maximizing the surface area for absorption. Extensive networks of hyphae also provide an
enormous surface area for absorptive nutrition in a fungal mycelium. Many fungi are able to
break down the cellulose in wood, cleaving the linkages between glucose subunits and then
absorbing the glucose molecules as food. Most fungi also digest lignin, an insoluble organic
compound that strengthens plant cell walls. The specialized metabolic pathways of fungi allow
them to obtain nutrients from dead trees and from an extraordinary range of organic compounds,
including tiny roundworms called nematodes. The mycelium of the edible oyster mushroom
Pleurotus ostreatus excretes a substance that paralyzes nematodes that feed on the fungus. When
the worms become sluggish and inactive, the fungal hyphae envelop and penetrate their bodies.
Then the fungus secretes digestive juices and absorbs the nematode’s nutritious contents, just
like it would from a plant source. This fungus usually grows within living trees or on old stumps,
obtaining the bulk of its glucose through the enzymatic digestion of cellulose and lignin from
plant cell walls. The nematodes it consumes apparently serve mainly as a source of nitrogen—a
substance almost always in short supply in biological systems. Other fungi are even more active
predators than Pleurotus, snaring, trapping, or firing projectiles into nematodes, rotifers, and
other small animals on which they prey.