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small droplet, termed the hylar droplet, is seen at
the base of the spore, then the spore is “popped” from
the sterigma into the space between the gills, where it
falls vertically, free from the fruitbody. After a period
of dormancy the haploid basidiospores germinate
to produce monokaryotic hyphae, which repeat the
whole cycle.
One of the most intriguing aspects of this life cycle
is the manner in which the dikaryon develops. This
is shown diagrammatically in Fig. 2.20. The dikaryotic
tip cell extends and synthesizes protoplasm until it
attains a critical cytoplasmic volume. Then a small
backwardly-projecting branch arises, and the two nuclei
divide synchronously – one of the nuclei dividing
along the axis of the hypha, and the other dividing so
that one of the daughter nuclei enters the branch. The
branch is then cut off by the development of a sep-
tum (cross-wall) and a septum also develops in the main
axis of the hypha. The effect of this is to create a tip
cell with two nuclei of different compatibility types, a
penultimate cell with a single nucleus, and a branch
with a single nucleus, isolated by septa. The branch then
fuses with the penultimate cell, by dissolution of the
wall, and the nucleus migrates into the penultimate cell,
restoring the dikaryotic condition. This process occurs
every time the nuclei divide. The resulting “bumps” on
the hyphae are easily seen by light microscopy, and are
termed clamp connections(Fig. 2.20). However, not
all members of the Basidiomycota produce clamp con-
nections, and yet they still grow as dikaryons, suggesting
that other regulatory mechanisms are involved.
If we refer back to the development of asci in Fig.
2.16, we see a remarkable similarity in the behavior

32 CHAPTER 2

Fig. 2.19Scanning electron micrograph of the basidia of
a mushroom, Coprinus cinereus. Note the inflated basidia,
each with four sterigmata, and the basidiospores, some
of which have already been released, or collapsed during
preparation of the specimen. (Courtesy of Dr Chris
Jeffree.)

Germination

Oidia

Homing

Plasmogamy
Dikaryotic
hyphae

Fruitbody
(Karyogamy)

Basidium

Basidiospores
Monokaryotic
hyphae

Fig. 2.18A generalized life cycle of a mush-
room; see text for details. (Courtesy of Maria
Chamberlain.)

basidiospores produce a dikaryotic mycelium that is
homothallic (self-fertile). However, the vast majority of
mushroom-producing fungi are heterothallic – about
90%. We will discuss the compatibility systems of
these fungi in Chapter 5.
The way in which the basidiospores are released
is not fully understood, but just before their release a

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