Classifications of Fungi
Overview
By the end of this section, you will be able to:
- Classify fungi into the five major phyla
- Describe each phylum in terms of major representative species and patterns of reproduction
The kingdom Fungi contains five major phyla that were established according to their mode of sexual reproduction or using molecular data. Polyphyletic, unrelated fungi that reproduce without a sexual cycle, are placed for convenience in a sixth group called a “form phylum”. Not all mycologists agree with this scheme. Rapid advances in molecular biology and the sequencing of 18S rRNA (a part of RNA) continue to show new and different relationships between the various categories of fungi.
The five true phyla of fungi are the Chytridiomycota (Chytrids), the Zygomycota (conjugated fungi), the Ascomycota (sac fungi), the Basidiomycota (club fungi) and the recently described Phylum Glomeromycota. An older classification scheme grouped fungi that strictly use asexual reproduction into Deuteromycota, a group that is no longer in use.
Note: “-mycota” is used to designate a phylum while “-mycetes” formally denotes a class or is used informally to refer to all members of the phylum.
Chytridiomycota: The Chytrids
The only class in the Phylum Chytridiomycota is the Chytridiomycetes. The chytrids are the simplest and most primitive Eumycota, or true fungi. The evolutionary record shows that the first recognizable chytrids appeared during the late pre-Cambrian period, more than 500 million years ago. Like all fungi, chytrids have chitin in their cell walls, but one group of chytrids has both cellulose and chitin in the cell wall. Most chytrids are unicellular; a few form multicellular organisms and hyphae, which have no septa between cells (coenocytic). They produce gametes and diploid zoospores that swim with the help of a single flagellum.
The ecological habitat and cell structure of chytrids have much in common with protists. Chytrids usually live in aquatic environments, although some species live on land. Some species thrive as parasites on plants, insects, or amphibians (Figure), while others are saprobes. The chytrid species Allomyces is well characterized as an experimental organism. Its reproductive cycle includes both asexual and sexual phases. Allomyces produces diploid or haploid flagellated zoospores in a sporangium.
Zygomycota: The Conjugated Fungi
The zygomycetes are a relatively small group of fungi belonging to the Phylum Zygomycota. They include the familiar bread mold, Rhizopus stolonifer, which rapidly propagates on the surfaces of breads, fruits, and vegetables. Most species are saprobes, living off decaying organic material; a few are parasites, particularly of insects. Zygomycetes play a considerable commercial role. The metabolic products of other species of Rhizopus are intermediates in the synthesis of semi-synthetic steroid hormones.
Zygomycetes have a thallus of coenocytic hyphae in which the nuclei are haploid when the organism is in the vegetative stage. The fungi usually reproduce asexually by producing sporangiospores (Figure). The black tips of bread mold are the swollen sporangia packed with black spores (Figure). When spores land on a suitable substrate, they germinate and produce a new mycelium. Sexual reproduction starts when conditions become unfavorable. Two opposing mating strains (type + and type –) must be in close proximity for gametangia from the hyphae to be produced and fuse, leading to karyogamy. The developing diploid zygospores have thick coats that protect them from desiccation and other hazards. They may remain dormant until environmental conditions are favorable. When the zygospore germinates, it undergoes meiosis and produces haploid spores, which will, in turn, grow into a new organism. This form of sexual reproduction in fungi is called conjugation (although it differs markedly from conjugation in bacteria and protists), giving rise to the name “conjugated fungi”.
Ascomycota: The Sac Fungi
The majority of known fungi belong to the Phylum Ascomycota, which is characterized by the formation of an ascus (plural, asci), a sac-like structure that contains haploid ascospores. Many ascomycetes are of commercial importance. Some play a beneficial role, such as the yeasts used in baking, brewing, and wine fermentation, plus truffles and morels, which are held as gourmet delicacies. Aspergillus oryzae is used in the fermentation of rice to produce sake. Other ascomycetes parasitize plants and animals, including humans. For example, fungal pneumonia poses a significant threat to AIDS patients who have a compromised immune system. Ascomycetes not only infest and destroy crops directly; they also produce poisonous secondary metabolites that make crops unfit for consumption. Filamentous ascomycetes produce hyphae divided by perforated septa, allowing streaming of cytoplasm from one cell to the other. Conidia and asci, which are used respectively for asexual and sexual reproductions, are usually separated from the vegetative hyphae by blocked (non-perforated) septa.
Asexual reproduction is frequent and involves the production of conidiophores that release haploid conidiospores (Figure). Sexual reproduction starts with the development of special hyphae from either one of two types of mating strains (Figure). The “male” strain produces an antheridium and the “female” strain develops an ascogonium. At fertilization, the antheridium and the ascogonium combine in plasmogamy without nuclear fusion. Special ascogenous hyphae arise, in which pairs of nuclei migrate: one from the “male” strain and one from the “female” strain. In each ascus, two or more haploid ascospores fuse their nuclei in karyogamy. During sexual reproduction, thousands of asci fill a fruiting body called the ascocarp. The diploid nucleus gives rise to haploid nuclei by meiosis. The ascospores are then released, germinate, and form hyphae that are disseminated in the environment and start new mycelia (Figure).
Art Connection
Which of the following statements is true?
- A dikaryotic ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A diploid ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A haploid zygote that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A dikaryotic ascus that forms in the ascocarp undergoes plasmogamy, meiosis, and mitosis to form eight ascospores.
Basidiomycota: The Club Fungi
The fungi in the Phylum Basidiomycota are easily recognizable under a light microscope by their club-shaped fruiting bodies called basidia (singular, basidium), which are the swollen terminal cell of a hypha. The basidia, which are the reproductive organs of these fungi, are often contained within the familiar mushroom, commonly seen in fields after rain, on the supermarket shelves, and growing on your lawn (Figure). These mushroom-producing basidiomyces are sometimes referred to as “gill fungi” because of the presence of gill-like structures on the underside of the cap. The “gills” are actually compacted hyphae on which the basidia are borne. This group also includes shelf fungus, which cling to the bark of trees like small shelves. In addition, the basidiomycota includes smuts and rusts, which are important plant pathogens; toadstools, and shelf fungi stacked on tree trunks. Most edible fungi belong to the Phylum Basidiomycota; however, some basidiomycetes produce deadly toxins. For example, Cryptococcus neoformans causes severe respiratory illness.
The lifecycle of basidiomycetes includes alternation of generations (Figure). Spores are generally produced through sexual reproduction, rather than asexual reproduction. The club-shaped basidium carries spores called basidiospores. In the basidium, nuclei of two different mating strains fuse (karyogamy), giving rise to a diploid zygote that then undergoes meiosis. The haploid nuclei migrate into basidiospores, which germinate and generate monokaryotic hyphae. The mycelium that results is called a primary mycelium. Mycelia of different mating strains can combine and produce a secondary mycelium that contains haploid nuclei of two different mating strains. This is the dikaryotic stage of the basidiomyces lifecyle and and it is the dominant stage. Eventually, the secondary mycelium generates a basidiocarp, which is a fruiting body that protrudes from the ground—this is what we think of as a mushroom. The basidiocarp bears the developing basidia on the gills under its cap.
Art Connection
Which of the following statements is true?
- A basidium is the fruiting body of a mushroom-producing fungus, and it forms four basidiocarps.
- The result of the plasmogamy step is four basidiospores.
- Karyogamy results directly in the formation of mycelia.
- A basidiocarp is the fruiting body of a mushroom-producing fungus.
Asexual Ascomycota and Basidiomycota
Imperfect fungi—those that do not display a sexual phase—use to be classified in the form phylum Deuteromycota, , a classification group no longer used in the present, ever-developing classification of organisms. While Deuteromycota use to be a classification group, recent moleclular analysis has shown that the members classified in this group belong to the Ascomycota or the Basidiomycota classifications. Since they do not possess the sexual structures that are used to classify other fungi, they are less well described in comparison to other members. Most members live on land, with a few aquatic exceptions. They form visible mycelia with a fuzzy appearance and are commonly known as mold.
Reproduction of the fungi in this group is strictly asexual and occurs mostly by production of asexual conidiospores (Figure). Some hyphae may recombine and form heterokaryotic hyphae. Genetic recombination is known to take place between the different nuclei.
The fungi in this group have a large impact on everyday human life. The food industry relies on them for ripening some cheeses. The blue veins in Roquefort cheese and the white crust on Camembert are the result of fungal growth. The antibiotic penicillin was originally discovered on an overgrown Petri plate, on which a colony of Penicillium fungi killed the bacterial growth surrounding it. Other fungi in this group cause serious diseases, either directly as parasites (which infect both plants and humans), or as producers of potent toxic compounds, as seen in the aflatoxins released by fungi of the genus Aspergillus.
Glomeromycota
The Glomeromycota is a newly established phylum which comprises about 230 species that all live in close association with the roots of trees. Fossil records indicate that trees and their root symbionts share a long evolutionary history. It appears that all members of this family form arbuscular mycorrhizae: the hyphae interact with the root cells forming a mutually beneficial association where the plants supply the carbon source and energy in the form of carbohydrates to the fungus, and the fungus supplies essential minerals from the soil to the plant.
The glomeromycetes do not reproduce sexually and do not survive without the presence of plant roots. Although they have coenocytic hyphae like the zygomycetes, they do not form zygospores. DNA analysis shows that all glomeromycetes probably descended from a common ancestor, making them a monophyletic lineage.
Section Summary
Chytridiomycota (chytrids) are considered the most primitive group of fungi. They are mostly aquatic, and their gametes are the only fungal cells known to have flagella. They reproduce both sexually and asexually; the asexual spores are called zoospores. Zygomycota (conjugated fungi) produce non-septated hyphae with many nuclei. Their hyphae fuse during sexual reproduction to produce a zygospore in a zygosporangium. Ascomycota (sac fungi) form spores in sacs called asci during sexual reproduction. Asexual reproduction is their most common form of reproduction. Basidiomycota (club fungi) produce showy fruiting bodies that contain basidia in the form of clubs. Spores are stored in the basidia. Most familiar mushrooms belong to this division. Fungi that have no known sexual cycle were classified in the form phylum Deuteromycota, which the present classification puts in the phyla Ascomycota and Basidiomycota. Glomeromycota form tight associations (called mycorrhizae) with the roots of plants.
Art Connections
Figure Which of the following statements is true?
- A dikaryotic ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A diploid ascus that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A haploid zygote that forms in the ascocarp undergoes karyogamy, meiosis, and mitosis to form eight ascospores.
- A dikaryotic ascus that forms in the ascocarp undergoes plasmogamy, meiosis, and mitosis to form eight ascospores.
Hint:
Figure A
Figure Which of the following statements is true?
- A basidium is the fruiting body of a mushroom-producing fungus, and it forms four basidiocarps.
- The result of the plasmogamy step is four basidiospores.
- Karyogamy results directly in the formation of mycelia.
- A basidiocarp is the fruiting body of a mushroom-producing fungus.
Hint:
Figure D
Review Questions
The most primitive phylum of fungi is the ________.
- Chytridiomycota
- Zygomycota
- Glomeromycota
- Ascomycota
Hint:
A
Members of which phylum produce a club-shaped structure that contains spores?
- Chytridiomycota
- Basidiomycota
- Glomeromycota
- Ascomycota
Hint:
B
Members of which phylum establish a successful symbiotic relationship with the roots of trees?
- Ascomycota
- Deuteromycota
- Basidiomycota
- Glomeromycota
Hint:
D
The fungi that do not reproduce sexually use to be classified as ________.
- Ascomycota
- Deuteromycota
- Basidiomycota
- Glomeromycota
Hint:
B
Free Response
What is the advantage for a basidiomycete to produce a showy and fleshy fruiting body?
Hint:
By ingesting spores and disseminating them in the environment as waste, animals act as agents of dispersal. The benefit to the fungus outweighs the cost of producing fleshy fruiting bodies.
For each of the four groups of perfect fungi (Chytridiomycota, Zygomycota, Ascomycota, and Basidiomycota), compare the body structure and features, and provide an example.
Hint:
Chytridiomycota (Chytrids) may have a unicellular or multicellular body structure; some are aquatic with motile spores with flagella; an example is the Allomyces. Zygomycota (conjugated fungi) have a multicellular body structure; features include zygospores and presence in soil; examples are bread and fruit molds. Ascomycota (sac fungi) may have unicellular or multicellular body structure; a feature is sexual spores in sacs (asci); examples include the yeasts used in bread, wine, and beer production. Basidiomycota (club fungi) have multicellular bodies; features includes sexual spores in the basidiocarp (mushroom) and that they are mostly decomposers; mushroom-producing fungi are an example.