Woese et al. (1990): 3 domains (Bacteria, Archaea, Eukarya).
Phylogenetic Tree of Life
Woese's Three Domains: Bacteria, Archaea, and Eukaryote, based on rRNA sequence data.
Number of Fungal Species
Approximately 100,000 fungal species have been described, with about 1,700 new species identified each year.
Hawksworth's Estimate (1991, 2001):
Ratio of fungi to plant species in the British Isles is 6:1 (12,000 fungi vs. 2,000 flowering plants).
Estimates based on this ratio suggest 1.5 million species of fungi worldwide.
Only a small fraction of fungal species has been described, and at the current rate, it would take over 800 years to describe all extant species.
More recent estimates, based on sequencing, suggest the number could be more than 5 million.
Definition of a Fungus
Fungus (singular): A eukaryotic, heterotrophic organism, devoid of chlorophyll, with chitin in cell walls, that obtains its nutrients by absorption, usually reproduces by spores, and usually exhibits a pronounced polarization of growing cells.
Fungi (plural).
Characteristics of Fungi
Eukaryotic: Possessing cells with a membrane-bound nucleus.
Prokaryotic: Lacking a membrane-bound nucleus.
Most fungi are comprised of hyphae that elongate by tip growth.
Hyphae collectively form the mycelium.
Hyphal Growth
Hyphal growth in Neurospora crassa is studied.
Fairy Rings
Fairy rings are naturally occurring rings or arcs of mushrooms.
Ringworm
Ringworm is a fungal infection of the skin.
Size of Mycelium
Mycelia can grow to be very large.
The largest living thing is a fungus in Oregon, covering 2,200 acres (3.5 square miles), estimated to be 2,400 years old and 605 tons in mass (Armillaria ostoyae).
Heterotrophic Nutrition
All fungi are heterotrophs with absorptive nutrition (saprotrophs).
They lack chlorophyll and are not autotrophic.
Cell Wall Structure
Unique cell wall structure containing chitin, a structural polymer of glucose.
Plants have cellulose, and animals lack cell walls.
Cell Compartmentalization
Fungi are multi-cellular organisms with a unique system of cell compartmentalization.
Septum Formation
Septum formation in Neurospora crassa is being studied.
Fungal hyphae of many species contain septa, which facilitate cell compartmentalization.
This compartmentalization distinguishes fungi from other multi-cellular organisms.
Fungal Cells
Exceptions: Yeasts are single-celled organisms, and many spores are single-celled.
Spores
Spores are reproductive propagules, usually microscopic.
Types of Spores
Sexual Spores (products of meiosis):
Ascospores
Basidiospores
Zygospores
Zoospores (meiotic)
Asexual Spores (products of mitosis):
Zoospores
Sporangiospores
Conidia
Learning Objectives
Discuss the kingdoms/domains of life.
Define 'Fungus'.
Understand the estimation of the number of fungal species in the world.
Gain familiarity with introductory mycology terminology.
Name the largest/oldest organism in the world.
Explain what is meant by a cell compartment.
Fungal Classification
How fungi are named and classified.
Examples: Chytridiomycota and Blastocladiomycota.
Examples of Fungi
Examples include Wood Ear, Turkey Tail, Orange Peel Fungus, Eyelash Fungus, Dog Vomit Fungus, Russula emetica, Stemonitis, P. coronata, Tilletia foetida, and Black spot of elm (Stegonophora ulmea).
Species Concepts
Morphological Species Concept:
Organisms that look the same are considered the same species.
Most fungal species are defined this way.
Biological Species Concept:
Interbreeding or potentially interbreeding natural populations constitute the same species.
Mating in the lab is often used to define species membership.
Phylogenetic Species Concept:
Individuals of the same species share a genealogical relationship inferred by phylogenetic analysis.
Assumes monophyly.
The most appropriate species concept depends on the taxa under investigation.
Fungal Phylogeny
Major groups: Chytridiomycota, Zygomycota, Basidiomycota, Ascomycota.
Fungal Evolution
Evolutionary adaptation of fungi over time.
Chytridiomycota
Approximately 1000 described species.
"True Fungi" with flagellate spores (Zoospores) in sporangia.
Do not make hyphae.
Thallus is coenocytic (septa are rare or absent).
Cell walls contain chitin and cellulose.
Mostly aquatic or soil-inhabiting saprobes, some are parasitic.
Chytridiomycota - Notable Species
Batrachochytrium dendrobatidis: pathogen of frogs and toads, responsible for amphibian decline worldwide.
Chytridiales
Olpidium brassicae: resting sporangia and zoospoorangia in rhizodermal cells.
Blastocladiales
Blastocladiella emersonii: Differentiation of sporangium depending on CO2 levels; complex life cycle.
Chytridiomycota vs. Blastocladiomycota
Chytridiomycota: no hyphae, zoospores.
Blastocladiomycota: with hyphae (no septa), zoospores.
Blastocladiomycota
Often referred to as 'Chytrids'.
"True Fungi" with flagellate spores (Zoospores) in sporangia.
They do make hyphae, septa are rare or absent.
Complex life cycles.
Mostly aquatic or soil-inhabiting saprobes.
Blastocladiomycota - Notable Species
Allmoyces macrogynous: A zoosporic aquatic fungus that makes hyphae that lack septa, exhibits complex life cycle with alteration of generations.
Blastocladiales Life Cycle
Alomyces macrogynus: Includes settlement of zygote, zygote germination, karyogamy, copulation, meiosis, and alternation of generations.
Learning Objectives Recap
Know why Latin is used to name and describe species.
Know the six major groups (phyla) of fungi (Chytridiomycota and Blastocladiomycota).
Discuss similarities and differences between the six major groups.
Know the name of the most common spores produced by each major group.
Describe each of the three species concepts described in class and when to use each.
Know the names of notable species.
Zygomycota and Zoopagomycota
Examples: Strawberry-Rhizopus time lapse - Rhizopus stolonifer eating your bagel.
Zygomycota
Unifying character: The Zygospore
Asexual spores are sporangiospores made in a sporangium.
With true hyphae that are mainly coenocytic, no septa in most newly developed hyphae.
Terrestrial
Cell walls have: Chitin, chitosan (non-acecylated chitan) and polyglucuronic acid
Note: some authors call this group Mucoromycota
Typical Zygomycota
Reproduces quickly
Makes many offspring
Utilizes simple sugars
Important Zygomycota
Rhizopus stolanifer (Black bread mold; model system for mating)
Phycomyces blakesleanus (model system for mating)
Pilobolus spp. (fascinating spore discharge)
Countless members of the Glomerales (90% of land plants form arbuscular mycorhizae with these fungi)
Rhizopus
Asexual Sporagia
Sexual Zygospores
Mating Type
Discuss Mating Type. Here + and –
Not the same as male and female, since + mating type can make male or female structures and – can make male or female structures.
These + or – is determined by a single locus, with a different idiomorphic gene present at the locus for each.
Heterothallic : requires two compatible thalli to produce sexual spores
Homothallic : capable of sexual reproduction from a single thallus
Rhizopus Life Cycle
Asexual and Sexual Reproduction
Rhizopus stolonifer
Asexual development: growth of aerial hyphae, sporangium differentiation, disease symptoms
Rhizopus sexualis
Sexual development: zygophore, progametangium, gametangium and zygospore
Acceleration of a top fuel drag racer = 15.8 m/s^2
Acceleration of bullet form .357 magnum handgun = 3.2 \times 10^5 m/s^2
Acceleration of Pilobolus sporangium = 2.1 \times 10^5 m/s^2
Zygomycota vs Zoopagomycota
Zygomycota: hyphae, no septa, zygospores, sporangiospores
Zoopagomycota: hyphae, no septa, zygospores, sporangiospores (~conidia), pathogens of animals
Zoopagomycota
All make a ZYGOSPORE as the sexual spore
Asexual spores are sporangiospores made in a sporangium…sporangia often germinate directly.
With true hyphae that are mainly coenocytic, no septa in most newly developed hyphae.
terrestrial
Most are typically insect or other animal pathogens.
Important Zoopagomycota
Basidiobolus ranarum (Pathogen of animals -humans)
Entomophthora muscae (Insect pathogen)
Learning Objectives Recap
Know the six major groups (phyla) of fungi (Zygomycota and Zoopagomycota today)
Be able to discuss similarities and difference between the six major groups
Know the name of the most common asexual and sexual spores produced by each major group
Be able to describe the spore discharge mechanism of Pilobolus
Be familiar with the life cycle of Rhizopus as a means of understanding asexual and sexual reproduction
Understand mating type with respect to the Zygomycota
Ascomycota and Conidiating Fungi
Discussing Ascomycota and deuteromycota (conidiating fungi)
Ascomycota
Production of an Ascus (Asci plural)
An Ascus is a sac-like structure filled with ascospores
Contains approximately 75% of the described species
The Most diverse and arguably the most important phylum
Ascomycota Characteristics
Have regularly spaced septa (usually with a central pore)
Cell walls contain chitin and glucans
Never produce motile spores
When produced, conidia (the asexual spores) are never produced by cytoplasmic cleavage in a sporangium
Ascomycota Phylogeny
Certainly a monophyletic group
Common ancestor circa 300-500 million years ago
Closely related to the Basidiomycota
Ascomycota Reproduction
Generally Ascomycetes are thought to reproduce both sexually and asexually
Some have only a sexual state known while others have only an asexual stage known.
Ascomycota characteristics
Thallus varies from single-celled to well-developed mycelium with regularly spaced septa. Most septa possess a single central pore.
Small spherical structures called Woronin bodies which are typically associated with each septum. Some yeasts have septa with micropores similar to a sieve
Woronin bodies
Spherical structures associated with the septum in ascomycetes
Textbook Ascus formation
Gametangial Contact/ Fusion
Ascus formation process
Diagrams illustrating the process of ascogonium and antheridium fusion leading to ascus formation
Crozier formation
asci initial
General life cycle
Diagram illustrating a generalized life cycle
Types of Ascocarps
Perithecium (perithecia pl.) a closed ascocarp with a pore (ostiole) at the top, and a wall of its own.
Ascocarp Variety
Apothecium (apothecia pl.) an open ascocarp
Ascocarp Variety
Cleistothecium (cleistothecia pl.) a completely closed ascocarp
Ascocarp Variety
Ascostroma (ascostromata pl.) a stromatic ascocarp bearing asci directly in locules within the stroma (also called pseudothecium)
Locule a cavity within a stroma
Stroma a somatic structure on which or in which fructifications are formed
Examples of Ascomycota
Chlorociboria sp.
Bulgaria inquinans
Examples of Pezizales
Cookeina sp.
Sarcoscypha coccinea
Urnula sp.
Aleuria aurantia, the orange peel fungus
Scutellinia scutellata, its orange apothecia rimmed with dark hairs
Ascobolus
The ascus and ascospore discharge
Truffles
Tuber texenses
Mitosporic Fungi
Deuteromycota (Ascomyctoa)
Ascomycetes Sexual/Asexual Stages
Sexual Stage= “perfect” stage or Meiotic stage produceing meiospores
Asexual Stage= “imperfect” stage or Mitotic stage produceing mitospores
Mitosporic Fungi
What is a conidium ? Conidia plural
Conidium a non-motile, asexual spore formed ‘one at a time’ of a conidiogenous cell.
Cell wall of conidium and the conidiogenous cell is continuous.
They are not formed by cleavage of cyotplasm as is the case for sporangiospores and zoospores we have already encountered in the class. Consequently they are not surrounded by an additional sporangial wall.
Holomorph Terms
Holomorph, Anamorph, Teleomorph concept:
Holomorph = All forms (morphs) of the fungus, either latent or expressed
Anamorph = Asexual (imperfect, mitotic) form(s)
Teleomorph = Sexual (perfect, meiotic) form
To properly classify a fungus, you must know its teleomorph. If no sexual stage is known, then the fungus must be given a temporary name based upon its asexual stage and placed in the Deuteromycota/ Mitosporic Fungi. If a sexual stage is discovered then the species must be renamed and placed in the appropriate phylum.
Conidia formation
Formation of conidia in Erysiphales; basipetal, blastic
Condiophores
Development in Apergillus nidulans
Conidiphore Development
Conidiogenesis is phialidic, blastic, basipetal
Learning Objectives Recap
Know the major groups of fungi (Ascomycota today)
Be able to discuss similarities and difference between the major groups
Know the name of the most common asexual and sexual spores produced by each major groups
Be able to explain the holomorph, teleomorph, anamorph concept
Know the general life cycle for the Ascomycota
Know the four types of ascocarp described today
Know the difference between conidia and sporangiospores