Introduction to the Fungi

Characteristics of "fungi" in the broad sense

1. Achlorophyllous: Fungi cannot make their own food like plants. They are
heterotrophs and depend upon other organism for their carbon source.
Heterotrophs can further be divided into the following categories:
– Parasites: Organisms that derives their nutrition from the protoplasm
of another organism (=host).
– Saprobes: Organisms that obtains their carbon source (=food) from
the by-products of organisms or dead organisms. However, if the
opportunity arises, some saprobes may become parasitic. Such
organisms are said to be facultative parasites.
– Symbiosis: In the strict sense, this term refers to the habitual "living
together" of different species. As such, there are a number of different
categories of relationships that may fit under this term. However, we
will define it in its most common usage: "The intimate association of
two dissimilar organisms in a mutually beneficial relationship, e.g.
lichens and mycorrhizae." This type of symbiosis is specifically referred
to as a mutualistic symbiosis.
2. Eukaryotic: Fungi have membrane bound organelles, i.e. nucleus,
mitochondrion, E.R., etc. Once upon a time filamentous bacteria
called Actinomycetes were classified with fungi, but this is no longer
the case.
3. The body or assimilative part of the fungus (=thallus) usually takes
the following forms:
– Yeast: Unicellular fungi that reproduce, asexually, by budding or
fission (terms to be defined later).
– Mycelium: The collective, filamentous strands that make up the fungal
thallus. Strands of mycelium is referred to as hyphae (sing.=hypha).
Mycelium may be of two types:
• Septate: Mycelium that is divided into discreet cells by cell walls that are laid
down at regular intervals along the length of the mycelium. These cell walls
are called septa (sing.= septum).
• Coenocytic: Mycelium that is not divided up by septa and forms a continuous
tubular network. Septa, however, are present occasionally, especially where
reproductive structures occur and where the cell wall of the mycelium has
been compromised.
– Some species may have thalli that are mycelium and yeast. Such fungi
are said to be dimorphic (=two forms).
4. The assimilative stage of the fungal body, i.e. mycelium or yeast, has a
cell wall. In the strict sense organisms classified as fungi have cell walls
composed primarily of chitin. However, we will be also be covering
"fungi" that do not have chitin in their cell walls.
Fungi have a common nutritional mode: Absorption: The transport of
food from their substrate into their cell walls. The following events occur
in this mode of nutrition:
If the available food that the fungus is using is soluble, i.e. a simple organic
compound, such as simple sugars and amino acids, the mycelium or yeast
cells can transport the food directly through their cell wall.
If the available food is insoluble, i.e. a large, complex, organic compound,
such as lignin, cellulose and pectin, then production the food must first be
digested. Digestion is carried out by the production of various enzymes that
are substrate specific and will break down insoluble food material to soluble
compounds that can be transported through the cell wall. Although this
appears to be very different from the way in which we (animals) digest food,
it differs only in the sequence of events that takes place. Where we ingest
food and then digest it, fungi first digest their food before ingestion
5. Either sexual or asexual reproduction or both may occur by
spores.
• Spores and/or gametes can be motile or not.
• However, in the strict sense as fungi are currently defined,
only those organisms that produce nonmotile spores and
gametes are classified as fungi.
• Nevertheless, we will be going over organisms that have
motile spores, called zoospores, and motile gametes.
• In summary then, the organisms that we call fungi
represent a heterogenous group, i.e., they are
polyphyletic, that are not closely related as you will soon
see.
Ainsworth and Bisby (1971) Bessey (1950) Alexopoulos (1962)
Kingdom Fungi Kingdom Plantae
Division Mycota
Myxomycota Myxomycotina
 Acrasiomycetes Mycetozoa  Myxomycetes
 Hydromyxomycetes Eumycotina
 Myxomycetes  Plasmodiophormycetes
 Plasmodiophoromycetes
Eumycota
Mastigiomycotina Phycomyceteae  Chytridiomycetes
 Chytridiomycetes  Hyphochytridiomycetes
 Hyphochytridiomycetes  Oomycetes
 Oomycetes
Zygomycotina
 Zygomycetes • Zygomycetes
 Trichomycetes • Trichomycetes
Carpomycetae
Ascomycotina Ascomyceteae Ascomycetes
 Hemiascomycetes  Hemiascomycetidae
 Plectomycetes Pyrenomycetes  Euascomycetidae
 Pyrenomycetes o Plectomycetes
 Discomycetes o Pyrenomycetes
 Laboulbenomycetes o Discomycetes
 Loculoascomycetes o Laboulbeniomycetes
 Loculoascomycetidae
Basidiomycotina Class: Basidiomyceteae Basidiomycetes
• Teliomycetes Teliosporeae Heterobasidiomycetidae
• Hymenomycetes Heterobasidiae Homobasidiomycetidae
o Phragmobasidiomycetidae Hymenomyceteae Gasteromycetes
o Holobaasidiomycetidae Gasteromyceteae
• Gastromycetes
Deuteromycotina The Imperfect Fungi Deteromycetes
 Blastomycetes  Moniliales
 Hyphomycetes  Sphaeropsidales
 Coelomycetes  Melanconiales
 Basidiomycota

Division: Basidiomycota
Class: Basidiomycetes
Order: Agaricales (Mushrooms)
Order: Lycoperdales, Phallales and
Nidulariales (Puffballs)
Order: Aphyllophorales (Polypores)
Order: Tremellales, Dacrymycetales and
Auriculariales (Jelly Fungi)
Class: Teliomycetes (Rust)
Class: Ustomycetes (Smuts)
 Basidiomycota
This division has many features in common with the
Ascomycota:
mycelia with chitinous cell walls that are regularly septate,
presence of an extended dikaryon stage, yeast stage and
presence of macroscopic fruiting bodies, in some taxa,
and conidia are produced if an asexual stage is present.
As in the Ascomycota and Zygomycota, the characteristic
that defines this subdivision is the sexual spore stage.
The sexual spores produced are basidiospores that are
typically borne, exogenously, on horn-like sterigmata
(sing.=sterigma) of basidia (sing.= basidium) (Fig. 1).
Figure 1: Unicellular
basidium, with four
sterigmata and
basidiospores. Basidium
illustrated to the left is
commonly used as
representative of the
typical basidium.
The morphology of the basidium, however, is variable
and a few of the variations are shown in the
micrographs below (Figs. 2-4), and the variations
observed below were once thought to be of
considerable significance in the phylogeny of the
Basidiomycota.
Figure 2: Cruciate-septate
basidium. This basidium is
divided into four chambers.
The basidium is named for
the "cross" that can be seen
when viewed,from above,
through the microscope.
 Figure 3: Transversely
septate basidium. This
basidium resembles
hyphal cells, with
sterigmata . Because of
its lack of
differentiation, this was
once considered to be a
primitive basidium.
 Figure 4: Tunning
fork basidium. The
basidium is named
for its obvioius
resemblance to a
tunning fork. This
basidium
produces only two
basidiospores.
Figure 5: Transversely
septate basidiospores
germinating from a
rust teliospore.
Basidiocarp is absent.
 SEM micrograph of
the distal end of a
basidium with its four
basidiospores, each
attached at the end of
a sterigma.
LIFE CYCLE OF BASIDIOMYCOTA
 Clamp Connection Formation
It is believed that the formation of clamp connections, where they occur in the
dikaryons of some species, function in ensuring that each cell is binucleate. In
describing their formation, below (Fig. 1), note that this is the case.

Figure 1. Formation of Clamp Connections: a.

Terminal cell of hypha. Growth only takes place at
hyphal tips; b. Hyphal tip elongating. c.
Synchronous division of nuclei and the beginning of
hyphal branch that will become the clamp
connection. One nucleus (b) migrates into the new
clamp. d. Septum forms at base of the clamp
trapping nucleus b. Nuclei a' and b' migrate to the
hyphal tip, while nucleus a migrates away from the
tip. e. Septum forms below clamp forming new cell
at hyphal tip. Fusion of the clamp to the adjacent
cell releases nucleus b to the adjacent cell. Now
both the terminal and subterminal are binucleate,
each with a compatible pair of nuclei.
 Basidiomycetes

The class Basidiomycetes includes those members that

produce their basidia and basidiospores on or in a
basidiocarp.
The morphology of the basidium is variable
However, rDNA sequencing analyst (Swann and Taylor,
1993), septal pore morphology and cell wall
biochemistry (McLaughlin et al, 1995) have determined
that far too much emphasis was placed on this
characteristic and all members of the Basidiomycota
that produce basidiocarps are now included in a single
class, the Basidiomycetes,
 Order: Agaricales
.This is the order that is commonly
referred to as mushrooms.
Basidiocarps of this order typically
are "fleshy" and have a stipe
(=stalk), pileus (=cap), and lamellae
(=gills) where the basidia and
basidiospores are borne (Fig. 1-4).
The Basidiospores in this order of
fungi are forcibly ejected from the
basidium, into the area between
the lamellar edges, which then
allows the spores to fall free from
the mushroom and be dispersed by
wind.
Figure 2: Low magnification of a cross
section through the lamella of a
mushroom
Figure 3: Higher magnification of section
through the lamella of mushroom.
Basidiospores now visible on the upper
and lower edge of lamella.
Figure 4: High magnification of basidia.
Center basidium shows two
basidiospores of four on sterigmata.
Figure 1: Various stage
of mushroom
development (right) and
mature, haploid
basidiospores on
basidia on the surface
of the lamella.
Figure 2: Mature
basidium with haploid
basidiospore that are
typically of different
mating strains
 Figure 3: Dikaryon
formation resulting
from fusion of a pair
of compatible
monokaryons. Clamp
connects may or
may not be present.
Figure 4: Formation
of basidiocarp.
Hymenial cells
(right), in lamella will
undergo karyogamy
and develop basidia
and basidiospores
Figure 5: The zygote
5a is the only diploid
stage. Four nucleate
stage after meiosis
5b. Formation of
basidiospores, but
nuclei have not
migrated into spores
5c. Nuclear
migration into
basidiospores 5d.
Figure 5: Amanita marmorata
ssp myrtacearum. Commonly
associated with Eucalyptus,
Melaleuca and Casuarina.
Figure 6: Cortinarius clelandii.
Commonly associated with
Eucalyptus, Melaleuca and
Casuarina.
Figure 7: Laccaria fraterna.
Commonly associated with
Eucalyptus, Melaleuca and
Casuarina.
Figure 8: Suillus salmonicolor.
Associated with species of Pinus.
Note that this species produces
a porous hymenium rather than
gills.
Figure 9: Amanita muscaria.
Associated with species of Pinus.
Order: Aphyllophorales
Species in this order are often coriaceous, leathery to
woody, but may also be fleshy. The basidiocarps and
hymenia are more variable than in the Agaricales.
Examples of members of this order can be observed
below (Figs. 10-14). As in the case of the Agaricales, the
basidiospores are forcibly ejected from the basidium and
are then dispersed by wind.
Figure 10: Ramaria fragilima, a
coral fungus. The basidia and
basidiospores are formed at the
tips of the basidiocarp.
Figure 11: Pycnoporous
sanguineus, a polypore. The
basidia and basidiospores are
formed in the pores of the
hymenium surface on the
basidiocarps (see Fig 12).

Figure 12: Porous hymenium from

a prepared slide.

Figure 13: Ganoderma

applanatum, another polypore.

Figure 14: Hydnellum peckii. The

basidia and basidiospores are
formed on the spines of the
hymenium surface of the
basidiocarp.
 Gasteromycetes (Puff balls)
The Gasteromycetes represent a number of orders that are not
closely related. The common name "puffballs", refers to the
basidiospores being "puffed" from the basidiocarp, in some
species. Unlike the Agaricales and Aphyllophorales, the puff
balls do not forcibly eject their basidiospores. This has led the
puffballs to evolve several interesting means of basidiospore
dispersal (Figs. 15-18).
The terminology having to do with basidiocarp structure also
differ. A hymenium is not formed in this group of fungi. Basidia
and basidiospores are formed throughout the fertile area of the
basidiocarp called the gleba. The part of the basidiocarp that
encloses the gleba is referred to as the peridium. These terms
will be referred to as we look at the examples below.
Figure 15. Lycoperdon perlatum is a species
belonging to the order Lycoperdales. The peridia
of the basidiocarps are the globose sacks that
you see on the left. They enclose the gleba,
which is a powdery mass of basidiospores at
maturity. Basidiospores are released through the
ostiole when raindrops or small mammals
impact the pliable peridium and causes the
basidiospores to "puff" out

Figure 16. Geastrum tripex is also a member of

the Lycoperdales and disperses its basidiospore
by the same mechanism as the above example.
This species is commonly called an earth-star
because of the stellate dehiscence of the outer,
leathery peridium. The inner peridium, which
encloses the basidiospores, is soft and pliable.
 Jelly Fungi
"Jelly Fungi" because of the jelly-like consistency of the
basidiocarp. gelatinous.
This type of basidiocarp becomes shrunken and
shriveled, when dried, but with available moisture
revives to its former consistency
The life cycle of this order is the same as that in
mushrooms. There are several orders in this group of
fungi and they are delimited by the morphology of their
basidium. The basidia are typically septate, the
exception being the tunning fork basidium, which is
aseptate, but is deeply lobed and produce only two
basidiospores. Three orders will be described below:
The Tremellales, Auriculariales and Dacrymycetales.
 Order: Tremellales
This order produces cruciate septate basidia (Fig. 1).
This basidium type is so-called because when viewed,
from above, under a compound microscope, the
basidium can be seen to be divided, evenly, into four
chambers by septa that intersect at right angles. The
intersection of the septa forms a cross or crucifix. The
basidiocarps are too variable to describe. However, we
will look at a few species of Tremella that can be found
in Hawai‘i (Fig 2-3).
Figure 1: A cruciate septate basidium.
Unfortunately, this is not a top view of
the basidium. Thus, you will be unable
to see the cruciate septa. A single
septum can be seen, along the globose,
basal part of the basidium. The red
coloration of the basidium is from the
1% phloxine stain that has been used to
mount this section.
Figure 2: Tremella fuciformis has a white,
glistening, foliose, and sometimes
translucent basidiocarp, when fresh. This
is the same species that is cultivated in
Japan, People's Republic of China and
Taiwan.
Figure 3: Tremella boraborensis has a
black, cerebriform basidiocarp, with
hollow lobes.
 Order: Auriculariales
This order is characterized by having a
transversely septate basidium (Fig. 4). As is
the case of the Tremellales, many species
have a gelatinous basidiocarp, reminiscent of
the consistency of jelly. There is only one
common, obvious species in Hawai‘i,
Auricularia cornea (Fig. 5)
Figure 4: A transversely septate basidium of
the genus Helicogloea. This genus was used
rather than Auricularia because the basidia
of Auricularia are compactly held together
by a gelatinous matrix that will not come
apart, making it impossible to make observe
an individual basidium.

Figure 5: A cluster of Auricularia cornea

growing on a fallen log. This species is
commonly cultivated in Asia and the South
Pacific. Locally, it is called "pepeiao" or "ear".
Order: Dacrymycetales
This order is characterized by presence of a tunning ford
basidium (Fig. 6). The basidiocarps are typically some shade of
yellow-orange and is gelatinous, as in the other two orders. Only
a few common species occur in Hawai‘i (Fig. 7-9)

Figure 6: A tunning fork basidium. The name is based on its resemblance

to a tunning fork. It differs from other basidia in that only two
basidiospores are produced per basidium.

Figure 7: Dacryopinax spathularia. Although common, in Hawai‘i, its small

size has probably contributed to its anonymity.

Figure 8: Calocera cornea. This species is both small and uncommon in

Hawai‘i.

Figure 9: Dacrymyces palmatus. A common temperate species that does not

occur in Hawai‘i.
Figure 6: A tunning fork basidium. The
name is based on its resemblance to a
tunning fork. It differs from other basidia
in that only two basidiospores are
produced per basidium.

Figure 7: Dacryopinax spathularia. Although

common, in Hawai‘i, its small size has
probably contributed to its anonymity.

Figure 8: Calocera cornea. This species is

both small and uncommon in Hawai‘i.

Figure 9: Dacrymyces palmatus. A common

temperate species that does not occur in
Hawai‘i.
 Uredinomycetes
Order: Uredinales (Rusts)
This class is composed of a single order of obligate parasites that are
pathogenic to ferns, gymnosperms and flowering plants. Basidiocarps
are not produced in this order and the life cycle is quite different than
that of the Basidiomycetes. In Puccinia graminis (Wheat Rust), the
species that will be used as the representative for this order, there are
five spore stages that are produced and two hosts are required in the
completion ofthe life cyle. The five stages produced are:
Stage 0:Spermagonium
Stage I: Aecium
Stage II: Uredium
Stage III: Telium
Stage IV: Basidium
As a means of comparing the life cycle of Puccinia graminis with that of
the mushroom life cycle, a summary of the life cycle will begin with the
basidiospore.
• In common cladistic usage, a monophyletic group is
a taxon (group of organisms) which forms a clade,
meaning that it contains all the descendants of the
possibly hypothetical closest common ancestor of
the members of the group. The term is synonymous
with the uncommon term holophyly. Monophyletic
groups are typically characterized by shared derived
characteristics (synapomorphies).
• Klad atau klade adalah suatu kelompok taksonomi
yang memiliki satu leluhur bersama dan semua
keturunannya juga berasal dari moyang tersebut.
Kelompok semacam itu dikatakan sebagai kelompok
monofiletik, yang dapat digambarkan melalui
analisis filogenetika, seperti dalam diagram pohon
atau suatu kladogram.
A polyphyletic group is any group other than a
monophyletic group or a paraphyletic group, which
like a paraphyletic group contains only some of the
descendants of their closest common ancestor, but
unlike a paraphyletic group is not characterized by the
missing descendants forming one (or more)
monophyletic groups.