Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
What are Fungi?
Biology of Fungi
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life Living organisms on
earth first arose about 3.5 billion years ago Prokaryotic Anaerobic
Oldest fossils of fungi are about 460 million years old Fossilized perithecium of what is believed to be a
fungus of the genus Savoryella. Note the ascospores (arrow) within the ascocarp. Source: www.ucmp.berkeley.edu/fungi/fungifr.html
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.)
Coincides with the rapid expansion of multi-cellular organisms
Major multicellular eukaryotes are divided into Kingdoms Animals Plants Fungi
Artistic vision of mushrooms among plants. Source: interactive.usc.edu/members/jchen/
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.)
Each of these three kingdoms differ in their basic cellular structure and mode of nutrition (defined by Whittaker, 1969) Plants - photosynthetic, cellulosic cell walls Animals - digestive systems, wall-less cells Fungi - absorptive nutrition, chitinous walls
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.)
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.) The estimates for the expansion of
multicellular organisms are based upon phylogenetic analyses of Carl Woese Examined ribosomal RNA (rRNA)
Present in prokaryotes and eukaryotes Relatively stable, but changes occur over time;
thereby acting as a chronometer Distinguished three separate groups
(Domains) of living organisms
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.) Domains - rRNA sequence differences
correlate with differences in cellular structure and physiology Bacteria - “true bacteria” Archaea - “ancient prokaryotes” Eucarya - eukaryotes
Taxonomic grouping of “Kingdom” lies beneath that of “Domain”
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Figure 1.1, Deacon (2006) Fungal Biology, 4th ed.
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.) Though the fossil evidence suggests
fungi were present on earth about 450 million years ago, aquatic fungi (Phylum Chytridiomycota) most likely were present about a million years before this time
About 354 - 417 million years ago, fungi evolved with primitive land plants
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in the Tree of Life (cont.) These plant-associated fungi probably
helped their photosynthetic partners gather nutrients from the harsh soils of the time
These fungi were the early ancestors of the present day phylum Glomeromycota
Despite plant-fungus co-evolution, fungi are more closely related to animals
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom
Mycology is the study of fungi Myco- = fungi -ology = the study of
Mycology originally arose as a branch of botany because fungi were once believed to be “achlorophyllic” plants
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Why Study the Fungi? There are over 100,000 species of
known fungi and probably 15 times that many that have yet to be discovered
Fungi are an extremely important part of the ecosystem Recycling of minerals and carbon Cause plant and animal diseases Source of food, medicines, and chemicals Important models in scientific research
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi
Greeks (about 300 B.C.) believed truffles were produced by thunder
Oldest illustration of fungi was found among the ruins of Pompeii (A.D. 79) that depicted edible mushrooms
A truffle. Source: www.truffle-and-truffe.com
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi Euripides (A.D. 450-456) recorded the
first mushroom poisoning deaths
Euripides. Source: uk.wikipedia.org Amanita muscaria, one of the most
poisonous mushrooms in the world. Source: www.myco-vaud.ch
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) During a period of the Middle Ages
(A.D. 1470 to 1670), books on ‘herbals’ were published that included illustrations and descriptions of fungi Remarkably similar to those of Greeks Used same methods to divide mushrooms
and truffles into poisonous and non-poisonous varieties
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) The German herbalist Jerome
Bock wrote in A.D. 1552: “Fungi and truffles are neither herbs, nor roots, nor flowers, nor seeds, but merely the superfluous moisture of earth, of trees, or rotten wood, and of other rotting things. This is plain from the fact that all fungi and truffles, especially those that are used for eating, grow commonly in thundery and wet weather.” [From Ainsworth (1976) Introduction to the History of Mycology, as cited in
Moore-Landecker (1996) Fundamentals of the Fungi, 4th ed.]
Jerome Bock.
Source: www.nndb.com
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) Other cultures also believed the fungi
originated from thunder and lightning Hindu god Soma was a child of
thunderstorms who offered hallucinogenic fluids from Amanita muscaria, one of the world’s most poisonous mushrooms
Similar legends existed in Guatemala and Mexico
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.)
Modern times in the study of fungi began with the invention of the microscope (about 1590-1600) by Hans and Zacharias Janssen of Holland
Zacharias Janssen. Source: www.astrophotoclub.com/history/telescope.htm
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) Robert Hooke used
the microscope to make the first drawings of a microscopic fungus (Mucor or Rhizopus) in 1665 and published them in his book Micrographia
Hooke’s drawing in Micrographia. Source: www.unb.br/ib/cel/microbiologia
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) In 1699, fungi are found
to be a component of lichens
Anton van Leeuwenhoek observes yeasts using a microscope (1673)
British Soldier Lichen. Source: www.buenavistatownship.org
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
A Brief History of Fungi (cont.) Hereafter, the study of fungi
exploded with many different contributions over the next 300 years, including the one gene-one enzyme hypothesis (Beadle and Tatum, 1941) and the 2001 Nobel Prize for cell division studies in yeast
Source: www.laskerfoundation.org
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom
Fungi are simple, eukaryotic microbes Many are microscopic Studies typically
employ standard microbiological techniques
Macroscopic (above; from Kendrick) and microscopic (below;
from Cooper) fungi
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Mycologists (fungal biologists) have traditionally studied not only the true fungi (e.g., mildew), but also fungus-like organisms (e.g., slime molds) Physarum polycephalum, a slime mold, growing
out of Petri dishes (upper image) and a closer view of the plasmodium phase (lower image) Source: waynesword.palomar.edu/slime1.htm
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
The kingdom Mycota is comprised of the true fungi
True fungi have the following features: Eukaryotic Transmission electron micrograph of a fungal cell
showing typical eukaryotic structures. Source: www.stchas.edu/faculty/zfitzgerald/fungi.jpg
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Fungal features (cont.): Typically grow as
filaments, termed hyphae (sing., hypha) via apical growth [the latter differs from the growth of other filamentous organisms]
Scanning electron micrograph of a fungal hyphae growing on the surface of a leaf. Source: www.abdn.ac.uk/ims/h-em/images/sem4/pages/fungal-hyphae-on-leaf.html
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Apical growth of a fungal hypha of Sclerotinia sclerotiorum. Source: Fungal Cell Biology Group (www.fungalcell.org)
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Hyphal strand of a Streptomyces species. Note the newly dividing cell (arrow). Source: zoology.okstate.edu/zoo_lrc/biol1114/sample_tests/preview_material/exam1/s03/preview-exam1_s03.htm
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Fungal features (cont.): Fungal hyphae
repeatedly branch to form a network of filaments termed a mycelium (sing., mycelia)
Drawing of a mycelium. Source: www8.nos.noaa.gov/coris_glossary/index.aspx?letter=m
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Fungal features (cont.): Some fungi grow as a single-celled entity,
termed a yeast, that grows either by a budding process or via binary fission
Budding yeast (left) and fission yeast (left). Sources: www.biochem.wisc.edu/yeastclub and www.steve.gb.com/science/model_organisms.html
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.) Fungal features (cont.):
Some fungi can switch growth forms between a hyphal phase and a yeast phase, a property known as dimorphism Typically induced by environmental
conditions A number of such fungi are disease-
causing agents of humans and animals
Dimorphism of Candida albicans. Source: www.explorepub.com/articles/darkfield_charts/fungus9.html
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Fungal features (cont.): Heterotrophic (chemo-organotrophs) -
require preformed organic compounds Absorb nutrients after degradation by
exogenously released enzymes
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.) Fungal features (cont.):
Unique cell wall components Chitin Glucans Rare instances of cellulose, but definitely fungal
cell walls are not as rich in this polymer as are plants
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.) Fungal features (cont.):
Typically have haploid nuclei Hyphae often have a number of haploid nuclei
present in each cell Some yeasts have a single diploid nucleus
Reproduce both sexually and asexually, typically through the production of spores
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.)
Asexual mitotically-derived spores (conidia; orange arrow in above figure) of the fungus Scedosporium apiospermum and the
meiotically-derived spores (ascospores within a specialized structure termed a cleistothecium; white arrow in figure to the
right) of the sexual form of the same organism given the designation Pseudallescheria boydii
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Defining the Fungal Kingdom (cont.) Fungal features (cont.):
Other differences [Deacon, Table 1.1] between fungi and animals and plants include: Histone types Sensitivity of microtubules to inhibitors Manner of lysine biosynthesis Membrane sterols Organellar structure/morphology
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi
Plant parasites Irish potato blight of
the 1840s Dutch elm disease Disappearance of
frogs in Costa Rica Phytophthora infestans growing into the leaf of a potato plant (above) and the
resulting rotting tuber from infection by this fungus (right). Sources: www.seedquest.com/News/releases/2005/may/12297.htm and
www.science.siu.edu/plant-biology/PLB117/JPEG%20files/potato.blight.jpg
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi
Plant symbionts Lichens (can also
form with cyanobacteria)
Mycorrhiza
Mycorhizzal fungus associated with roots of a pine. Source: www.virtualmuseum.ca/Exhibitions/Mushroom/English/Species/mycorrhizal.html
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi (cont.)
Human pathogens About 200 known species of fungi are
known to infect humans Diverse diseases including:
Dandruff “ring worm” Pneumocystis infection of HIV-infected persons Candidiasis
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi (cont.)
Biological control agents Mycoparasites (other fungi) Entomopathogens (insects) Nematophagous (nematodes)
Nematode trapping fungus. Source: Kendrick
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi (cont.) Decomposition
Cellulose (plant material) Rumen fungi in cows Dry rot Dry rot due to the fungus Serpula lacrymans.
Source: www.grzyby.pl/gatunki/Serpula_lacrymans.htm
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Major Activities of Fungi (cont.) Toxin production (mycotoxins)
Aflatoxins (peanuts and grains) Mushroom poisoning
Moldy corn due to the aflatoxin producer Aspergillus flavus. Source: www.ipm.iastate.edu/ipm/icm/2001/10-22-2001/earrot.html
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in Biotechnology
Foods and flavorings Edible mushrooms
5 million tons produced worth $14 billion (1994) Diverse types now widely available in
supermarkets Alcoholic beverages Breads, cheeses, soy sauce Quorn mycoprotein
Lecture: What are Fungi? BIOL 4848/6948 - Fall 2009
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BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in Biotechnology (cont.)
Fungal metabolites Two categories
Primary - intermediates or end products of common metabolic pathways essential for normal cellular function
Secondary - diverse range of compounds formed by specific pathways of a given organism and not essential for growth (but may provide some selection advantage)
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in Biotechnology (cont.)
Fungal metabolites (cont.) Examples of primary metabolites
Citric acid (estimated 200,000 tons produced in the year 2000) [soft drinks]
Gluconic acid (estimated annual production of 100,000 tons) [food additive]
Itaconic acid (estimated annual production of 80,000 tons) [paint and adhesive manufacture]
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in Biotechnology (cont.)
Fungal metabolites (cont.) Examples of secondary metabolites
β-lactam antibiotics, e.g., penicillins and cephalosporins
Non-β-lactam antibiotics, e.g., griseofulvin, gliotoxin, ciclosporins
Pullulan - film-wrap for food in Japan Chitosan - sewage clarification, plant defense
initiator
BIOL 4848/6948 (v. F09) Copyright © 2009 Chester R. Cooper, Jr.
Fungi in Biotechnology (cont.)
Enzymes and enzymic conversions Extracellular enzymes Commercially valuable roles
Food industry Bioconversions
Heterologous gene products - expression of foreign proteins by fungi having medical/industrial applications