Plectomycetes 171

contains two compatible nuclei, and functions as ascus mother cell. Out of the two nuclei of ascus mother cell one is male and
another is female (Miller, 1984).
According to some workers there is no crozier formation in the ascogenous hypha, and its terminal binucleate cell starts to func-
tion as an ascus mother cell. The two nuclei in the ascus mother cell fuse and form a diploid zygotic nucleus (Fig.15.4 F).
Diploid nucleus of the ascus divides by meiosis into four haploid nuclei, each of which divides mitotically, giving a total of
eight haploid nuclei (Fig.15.4 G). Each nucleus gets surrounded by cytoplasm and develops into an ascospore.
At the time when asci and ascospores are developing from the ascogenous hyphae, many sterile hyphae grow up from the base
of the archicarp. These sterile hyphae enclose the developing asci completely from all sides, and form a pseudoparenchymatous
sheath or peridium. This peridium is a wall of multicellular sterile hyphae. The entire structure (asci, ascogenous hyphae and pe-
ridium) appears like a hollow ball of nearly a pin-head size, and represents the fruiting body or ascocarp (Fig. 15.4 H). Because
of its closed nature it is of cleistothecium type.
The mature cleistothecia (Fig.15.4 H, J) are often yellow and globose, and contain a wall of multicellular, closely-packed sterile
hyphae. Many globose or pear-shaped asci are present inside. Each ascus contains eight ascospores (Fig. 15.4 J). Soon after the as-
cospore formation, the wall of the asci breaks down, and the ascospores are released into the cleistothecium. The cleistothecium
decays and the ascospores are released.
Each ascospore is pully-wheel shaped (Fig. 15.4 K), and attains a diameter of approximately 5 mm. The spore
wall is differentiated into an outer thick and sculpturous epispore and inner thin endospore. Each ascospore germinates into a new
mycelium (Fig. 15.4 A).

15.7 PENICILLIUM

15.7.1 Systematic Position (According to Ainsworth, 1973)

Division – Eumycota
Subdivision – Ascomycotina
Class – Plectomycetes
Order – Eurotiales
Family – Eurotiaceae
Genus – Penicillium

15.7.2 Nomenclatural Problem

Kirk et al. (2001) mentioned that Penicillium is “anamorphic Eupenicillium – Talaromyces” represented by 223 wide-
spread species. Eupenicillium is represented by 43 widespread species of Anamorph Penicillium belonging to family
Trichocomaceae, while 24 species of Talaromyces also belong to Anamorph Penicillium of the same family.
Like Aspergillus, Penicillium is also a form-genus based on conidial morphology. Because of the absence of perfect states
in a majority of the species, Penicillium also shows the same taxonomic and nomenclatural problems as shown by Aspergillus.
Its perfect state has been assigned many different generic names, such as Talaromyces and Eupenicillium.

15.7.3 Occurrence
Commonly called ‘green moulds’ or ‘blue moulds’, Penicillium species occur frequently on Citrus and other fruits, cheese and many
other similar foodstuffs. Like those of Aspergillus, its conidia are also present almost everywhere in the air, and therefore
Penicillium species are also common biological contaminants of the laboratory cultures. Contamination of the bacterial culture
of Staphylococcus by Penicillium notatum in the laboratory of Alexander Fleming in 1928 (see Raper, 1978) actually became the
basis of the discovery of the antibiotic, penicillin, the first ‘Wonder-drug’ of the world.
172 Fungi and Allied Microbes

Besides Citrus fruits, Penicillia occur commonly on apple, leather and other fabrics, and on all kinds of decaying materials and
spoiling foods. Some species cause skin and other diseases of man and animals.

15.7.4 Economic Importance

1. Penicillium notatum (Fleming, 1944) and P. chrysogenum (Raper, 1952, 1978) are used for the production of penicillin
throughout the world.
2. The antibiotic griseofulvin, used in skin and nail infection, is prepared from P. griseofulvum.
3. Cheese is fermented by P. camemberti and P. roqueforti.
4. Decaying and rotting of Citrus fruits is done by P. digitatum and P. italicum, whereas apples are decayed by P. ex-
pansum.
5. Some species destroy the leather and fabrics, whereas others are associated with some human and animal diseases.

15.7.5 Laboratory Culture

If a moist Citrus fruit or cheese piece is covered with a hollow large cup and placed in damp warm place, bluish-green patches of
Penicillium appear within a few days.

Oil Cytoplasm
15.7.6 Somatic Structures globules Septum
Cell wall Nucleus
pore
The mycelium is well-branched, and consists of many septate
hyphae, which are slender, branched, tubular and generally hya-
line (Fig. 15.5 A). The septa have central pore, which allows the A
Conidia Conidia
cytoplasm and nuclei to flow from cell to cell. Some of the hy-
phae enter into the substratum to absorb the food material, whereas
others remain on surface to produce conidiophores and conidia.
Each cell is usually uninucleate, but may also be bi- or multi-
Phialide Phialide
nucleate. The cytoplasm encloses mitochondria, ribosomes,
endoplasmic reticulum and other usual inclusions of a fungal Metula
cell. Reserve food is present in the form of oil globules. Conidio- Ramus
phore

15.7.7 Vegetative Reproduction

B
It takes place by fragmentation. The vegetative mycelium breaks Conidia
up into two or more fragments, each of which develops into
a new mycelium.
Conidiophore

15.7.8 Asexual Reproduction C

Conidia D
Conidia developing on conidiophores are the means of asexual
Phialide
reproduction. Electron microscopic studies of Penicillium conid- G
Conidio-
ia have been made by Kolesnik (1981, Photoplate 3 A-J) and phore
many others. F H
E
A conidiophore is an erect, tubular hyphal outgrowth which
may develop from any cell of the mycelium, but not from a spe-
cialized foot cell as in Aspergillus. The conidiophores of only Fig. 15.5 Penicillium. A, A portion of hypha; B, Un-
some species are unbranched as in P. spinulosum (Fig. 15.5 B) branched conidiophore of P. spinulosum;
and P.thomii. But in a majority of species the conidiophores C, Branched conidiophore of P expansum;
D-E, Conidiophore and conidia of P. digi-
are branched as in P. expansion (Fig. 15.5 C) and P. digi-
tatum; F-H, Conidia and their germination.
 Plectomycetes 173

tatum. Successive whorls of conidiophore branches terminate into clusters of phialides or sterigmata. If the conidiophore
is unbranched, the phialides are borne directly on its tip (Fig. 15.5 B). But in species with branched conidiophores, the ultimate
branches, which contain the tufts of phialides, are called metulae (sing. metula). Usually the conidiophores show more branch-
ings, and these lower branches that support the metulae are called rami (sing. ramus) as in P. expansun (Fig. l5.5 C) and
P. digitatum (Fig. l5.5 D). A conidiophore with phialides and terminal chains of conidia appears like that of an artist’s
brush, and hence is technically called penicillus(L. penicillum, small brush). Only because of its brush-like appearance the
generic name Penicillium is given. From the tip of bottle-shaped phialides develop conidia.
The conidia (Plate 3 A-J) are globose to ovoid in shape with smooth or rough wall. Usually, they are uninucleate (Fig. 15.5 F)
but in some species they are multinucleate. They are green, blue, or yellow-coloured structures, and are produced in enor-
mous quantities to provide the characteristic colour to the colony. Martinez et al. (1982) studied the morphology of the
conidia of 211 species or Penicillium, and classified them into six major groups: smooth-walled (7% of the species), deli-
cately roughened (13%), warty (28%), echinate (10%), striate with low irregular ridges (36%) and striate with scarce high
ridges and bars (6%).
According to Alexopoulos and Mims (1979) the conidia in Penicillium develop exactly in the same manner as in Asper-
gillus. At the time of the formation of conidia (Fig. 15.5 E), the nucleus of a phialide divides, and one daughter nucleus
migrates into the narrow portion at the apex. The apex is then cut off by a cross-wall, and this separated uninucleate cell
develops into a conidium. The conidium later on secretes its own spore wall.
Electron microscopic studies of Fletcher (1971) in P. clavigerum (Fig. 15.6) suggest that at the time of the conidium-
formalion an apical plug of material (Fig. 15.6 A) lines the neck of the phialide. This apical plug remains separate from
the phialide wall. The primary wall of the conidium is formed by this apical plug. A centripetal ingrowth of the wall
material (Fig. 15.6 B) develops into a septum. Thus septum separates the protoplast of the conidium from that of the phi-
alide (Fig. 15.6 C). The septal pore is closed and the conidial protoplast is separated completely. Inside the primary wall
of the conidium develops a new wall layer (Fig. 15.6 D). The wall that separates two conidia functions as a connective
(Fig. 15.6 D). The primary wall of each conidium becomes folded, and because of this the conidium appears rough.

Apical plug
Connective

Phialide wall Conidium

Surface
layer
Septal pore

Nucleus Septum Septum

New wall
A B C D

Fig. 15.6 A–D, Penicillium clavigerum, showing development of conidia as viewed under electron microscope (after
Fletcher, 1971)

The conidia are dispersed by wind. On getting the suitable conditions of moisture and temperature, each conidium
swells, and germinates by producing a germ tube. The latter becomes septate and develops into the mycelium.
174 Fungi and Allied Microbes

A different terminology has been used to describe the Penicillium

different conidiogenous structures in Penicillium by Sam-
son and Pitt (1985) and Kirk et al. (2001), as is clear from
Fig. 15.7.
Conidiogenous cell
Cell supporting
15.7.9 Sexual Reproduction conidiogenous cell

About sexual reproduction in most of the species of Penicil- Branch cell

Conidiophore Branch
lium, nothing is known. Only a few species are known to
reproduce sexually. The sexual or perfect state of Penicillium
has been assigned to different genera, such as Talaromyces,
Eupenicillium and Carpenteles (Webster, 1980). Alexopoulus
and Mims (1979) mentioned that Carpenteles and Eupeni-
cillium are the two names of the same genus, and now the
valid name is Eupenicillium.
Fig. 15.7 Diagram to describe the different conidiog-
A few species that show sexual reproduction are Penicil- enous structures in Penicillium (after Samson
lium vermiculatum (= Talaromyces vermiculatus) and P. stip- & Pitt, 1985).
itatum (= T. stipitatus). Cleistothecial type of fruiting bodies
are formed. According to Miller (1984) the details of formation of cleistothecia, ascus formation and release of ascospores
are very similar to that of Aspergillus.
Dangeard (1907) studied the sexual reproduction in Penicillium vermiculatum (= Talaromyces vermiculatus), and
noted the following details:
The mycelium contains uninucleate cells (Fig.15.8 A), from which develops a uninucleate ascogonium. The ascogo-
nium elongates, and its nucleus divides several times to produce as many as 64 nuclei (Fig. 15.8 B). Simultaneously, a
uninucleate antheridium also develops and coils spirally around the multinucleate ascogonium (Fig. 15.8 B). The antheridial

Mycelium
Ascogonium
G

Antheridium
er
m
in
at

A
io

Ascospores
n

Peridium Sterile hyphae

B
Ascospores
Ascogoniu
Ascus with
binucleate
cells
am ?
?
og is
y
ry ios

E
Ka Me

Fig. 15.8 A–F, Sexual reproduction in Penicillium vermiculatum (= Talaromyces vermiculatus) (after Dangeard, 1907).
 Plectomycetes 175

tip touches the ascogonium at one point, and the walls of the two, at the point of contact, dissolve. Side by side the ascogonium
gets segmented in the form of many binucleate cells (Fig. 15.8 C). According to Dangeard the antheridial nucleus does
not migrate into the ascogonium. It remains very much there in the antheridium, even after the development of ascogenous
hyphae. This indicates that antheridia are formed but they remain non-functional. From the binucleate cells of the septate
ascogonium develop the ascogenous hyphae. The nuclei in the ascogenous hyphae, therefore, are derived from the original
nucleus of the ascogonium. Many sterile hyphae develop and enclose the young asci from all sides. The fruiting body, therefore,
is of cleistothecium type. Many asci, each with eight ascospores, are seen in the cleistothecium (Fig. 15.8 E). Dangeard (1907)
did not observe the actual processes of karyogamy and meiosis. But two nuclei of each ascogenous hypha (Fig. 15.8 D) might
have fused to form a diploid nucleus in the young ascus mother cell, and this zygotic nucleus might have passed through meiosis
to form eight ascospores in each ascus (Fig. 15.8 E). The asci are globose or pear-shaped bodies. Their walls soon dissolve, and
the ascospores are released in the cleistothecium. As in Aspergillus, the ascospores are pully-wheel shaped and uninucleate
bodies. Each ascospore germinates into a fresh mycelium.

15.8 DIFFERENCE BETWEEN TALAROMYCES AND EUPENICILLIUM

Table 15.2 Difference between Talaromyces and Eupenicillium

No. Talaromyces Eupenicillium

1. Fruiting body grows indefinitely and its size keeps on Fruiting body atttains a definite size.
increasing even after the maturation of ascospores.
2. Cleistothecial wall is made up of loosely Cleistothecial wall is hard and made up of thick-walled
arranged interwoven hyphae. pseudoparenchyma.
3. Asci are produced from croziers in most of the species. The asci generally develop on short branches of the
ascogenous hyphae.

15.9 DIFFERENCE BETWEEN ASPERGILLUS AND PENICILLIUM

Table 15.3 Difference between Aspergillus and Penicillium.

No. Aspergillus Penicillium

1. The conidiophores are unseptate and unbranched. The conidiophores are septate and branched.
2. A conidiophore develops from a specialized,thick-walled cell Conidiophore develops from any vegetative cell of the
called foot cell. mycelium; foot cells are absent.
3. Each conidiophore enlarges into a swollen vesicle at its tip. Vesicle is not formed at the tip of conidiophore,
and the entire structure appears like that of an artist’s brush.
4. Metulae are not present below the phialides. Metulae are present.
5. Mature conidia are yellow, green, brown or black in colour. Usually the mature conidia are green in colour.
6. Wall of the cleistothecium is loose. Wall of the cleistothecium is comparatively thick.

15.10 ROLE OF ASPERGILLUS AND PENICILLIUM IN BIOTECHNOLOGY

After Saccharomyces cerevisiae, two most important organisms used in biotechnology are Aspergillus and Penicillium.
Brief consideration of some of the examples the role of these two organisms in biotechnology is undermentioned.