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72 Penicillium Link: Fries
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Penicillium is a very large and ubiquitous genus which currently contains 354 accepted species (Visagie et al.,
2014). Many species are common contaminants on various substrates and are known as potential mycotoxin
producers. Correct identification is therefore important when studying possible Penicillium contamination of
food. Human pathogenic species are rare, however opportunistic infections leading to mycotic keratitis,
otomycosis and endocarditis (following insertion of valve prosthesis) have been reported (Lyratzopoulos
et al., 2002). Note: Species within the subgenus Biverticillium, including Penicillium marneffei have been
transferred to the genus Talaromyces; colonies in this genus often have bright colours (Samson et al., 2011b).
RG-1 organisms
Morphological genus description: Colonies are usually fast growing, in shades of green, sometimes white,
mostly consisting of a dense felt of conidiophores (Fig. 72.1). Microscopically, chains of single-celled conidia
are produced in basipetal succession from a specialised conidiogenous cell called a phialide. The term basocatenate is often used to describe such chains of conidia where the youngest conidium is at the basal or
proximal end of the chain. In Penicillium, phialides may be produced singly in groups or from branched
metulae, giving a brush-like appearance (a penicillus). The penicillus may contain both branches and metulae
(penultimate branches which bear a whorl of phialides). All cells between the metulae and the stipes of the
conidiophores are referred to as branches. The branching pattern may be either simple (non-branched or
monoverticillate), one-stage branched (biverticillate-symmetrical), two-stage branched (biverticillateasymmetrical) or three- to more-stage branched (Fig. 72.2). Conidiophores are hyaline, smooth or roughwalled. Phialides are usually flask-shaped, consisting of a cylindrical basal part and a distinct neck, or
lanceolate (more or less with a narrow basal part tapering to a somewhat pointed apex). Conidia are in long
dry chains, divergent or in columns, are globose, ellipsoidal, cylindrical or fusiform, hyaline or greenish,
smooth or rough-walled (Fig. 72.3). Sclerotia are produced by some species.
Fig. 72.1. Penicillium species culture.
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis) 196
Phialides
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Metulae
Branches
Fig. 72.2. Types of conidiophore branching in Penicillium (a) monoverticillate; (b) biverticillate; (c) terverticillate; (d)
quaterverticillate (Visagie et al., 2014).
Molecular identification: ITS and/or β-tubulin loci are recommended for identification of Penicillium species
(Visagie et al., 2014; Yilmaz et al., 2014). ITS sequencing alone is often insufficient to resolve species, and
morphological characters may be required for confirmation.
Key features: Flask-shaped phialides arranged in groups from branched metulae forming a penicillus.
Antifungal susceptibility: (Table 72.1).
Table 72.1. Penicillium spp. (Australian national data); MIC μg/mL.
Antifungal No. <0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 70 4 6 11 14 21 9 5
ISAV 17 1 4 6 2 4
VORI 68 2 6 12 10 12 5 21
POSA 67 1 6 16 13 20 6 1 4
ITRA 70 1 3 13 12 24 11 2 4
Descriptions of Medical Fungi 197
(a) (b)
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20 ˜m
Fig. 72.3. (a) Simple conidiophore of P. chermesinum showing long chains conidia and (b) conidiophores of P.
aurantiogriseum showing two-stage branching.
20 ˜m
References: Raper and Thom (1949); Pitt (1979); McGinnis (1980); Onions et al. (1981); Ramirez (1982);
Samson et al. (1995, 2011b); Domsch et al. (2007); de Hoog et al. (2015).
DOI: 10.1079/9781800622340.0072
Descriptions of Medical Fungi 198
73 Phaeoacremonium parasiticum
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(Ajello et al.) W. Gams et al.
Synonymy: Phialophora parasiticum Ajello, Georg & Wang.
The genus Phaeoacremonium initially accommodated isolates with features similar to those seen in both
Acremonium and Phialophora. It differs from the former by having pigmented hyphae and conidiophores
and from the latter by having indistinct collarettes and warty conidiogenous cells (Revankar and Sutton,
2010).
Phaeoacremonium currently consists of 46 species with P. parasiticum and P. krajdenii recognised as the
predominant species associated with human infections (Mostert et al., 2005). Other species have also been
isolated from clinical cases, i.e. P. alvesii, P. amstelodamense, P. griseorubrum, P. minimum, P. rubrigenum,
P. tardicrescens, and P. venezuelense. Infections caused by P. parasiticum include subcutaneous abscesses,
thorn-induced arthritis, and disseminated infection (Revankar and Sutton, 2010; Gramaje et al., 2015).
RG-2 organism
Morphological description: Cultures usually slow growing, suede-like with radial furrows, initially whitish-grey
becoming olivaceous-grey with age (Fig. 73.1). Hyphae hyaline, later becoming brown and some becoming
rough-walled. Phialides are brown, thick-walled, slender, acular to cylindrical slightly tapering towards the
tip, 15-50 μm long, often proliferating, with small, funnel-shaped collarettes (Fig. 73.2). Conidia, often in
balls, are hyaline, thin-walled, cylindrical to sausage-shaped, 3-6 x 1-2 μm, later inflating. Maximum growth
temperature 40
o
C.
Molecular identification: ITS and/or β-tubulin sequencing (Mostert et al., 2006; Gramaje et al., 2015).
10 ˜m
Fig. 73.1. Phaeoacremonium parasiticum culture and microscopy of a single phialide.
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
199
Key features: The identification of the different Phaeoacremonium species can be performed by combining
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cultural, morphological and ITS, β-tubulin sequence data (Mostert et al., 2006; Gramaje et al., 2015). Relying
on morphological characteristics alone can be difficult for species identification.
20 ˜m
Fig. 73.2. Phaeoacremonium parasiticum phialides with small, funnel-shaped collarettes (arrow).
Antifungal susceptibility: (Table 73.1).
Table 73.1. Phaeoacremonium parasiticum (Badali et al., 2015; and Australian national data); MIC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 32 ≥64
AmB 24 1 3 9 5 6
ISAV 19 1 3 5 6 4
VORI 24 3 1 6 13 1
POSA 19 5 8 5 1
ITRA 24 1 1 3 19
References: Mostert et al. (2005, 2006); Revankar and Sutton (2010); Badali et al. (2015); de Hoog et al.
(2015); Gramaje et al. (2015).
DOI: 10.1079/9781800622340.0073
Descriptions of Medical Fungi 200
The genus Phialophora contains more than 40 species, most are saprophytes commonly found in soil or on
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decaying wood. Some human pathogens with phialidic conidiogenesis previously assigned to Phialophora
have been moved to other genera, namely, Phaeoacremonium and Pleurostoma. Phialophora verrucosa,
P. americana, P. bubakii, P. europaea and P. reptans remain of medical interest (Revankar and Sutton, 2010).
Both P. verrucosa and P. americana produce their conidia from phialides with conspicuous darkened collarettes,
however sequencing has demonstrated a close relatedness, suggesting that these species may be synonymous
(de Hoog et al., 1999). P. verrucosa is primarily an agent of chromoblastomycosis although other reported
infections include endocarditis, keratitis, and osteomyelitis (Li et al., 2017).
RG-2 organism
Morphological description: Colonies are slow growing, initially dome-shaped, later becoming flat, suede-like
and olivaceous to black in colour. Phialides are flask-shaped or elliptical with distinctive funnel-shaped,
darkly pigmented collarettes (Fig. 74.1). Conidia are ellipsoidal, smooth-walled, hyaline, mostly 3.0-5.0 x
1.5-3.0 μm, and aggregate in slimy heads at the apices of the phialide.
Key features: Characteristic flask-shaped phialides with distinctive funnel-shaped, darkly pigmented
collarettes.
Molecular identification: ITS sequencing recommended (de Hoog et al., 1999).
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
201
20 ˜m
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20 ˜m
Fig. 74.1. Phialophora verrucosa phialides with distinctive funnel-shaped collarettes (arrow) and conidia.
Antifungal susceptibility: (Table 74.1).
Table 74.1. Phialophora verrucosa limited data (Australian national data); MIC μg/mL.
Antifungal No. ≤0.016 0.03 0.06 0.125 0.25 0.5 1 2 4 8 16 ≥32
AmB 7 1 4 2
VORI 7 1 3 3
POSA 5 2 4 1
ITRA 7 1 4 2
References: Ellis (1971); McGinnis (1978b, 1980); de Hoog et al. (1999, 2015); Domsch et al. (2007);
Revankar and Sutton (2010); Li et al. (2017).
DOI: 10.1079/9781800622340.0074
Descriptions of Medical Fungi 202
Members of the genus Phoma have a worldwide distribution and are ubiquitous in nature, with over 200
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species having been described from soil, as saprophytes on various plants, and as pathogens to plants and
humans.
RG-1 organism
Morphological description: Colonies are spreading, greyish-brown, powdery or suede-like and produce large,
globose, membranous to leathery, darkly pigmented, ostiolate pycnidia (Fig. 75.1). Conidia are produced in
abundance within the pycnidia on narrow thread-like phialides, which are hardly differentiated from the
inner pycnidial wall cells. Conidia are globose to cylindrical, one-celled, hyaline, and are usually extruded in
slimy masses from the apical ostiole.
Molecular identification: ITS, D1/D2, β-tubulin and 18S sequencing has been used to identify Phoma species
(de Gruyter et al., 2009; Aveskamp et al., 2010). Note: Public sequence databases, particularly GenBank,
contain many sequences from incorrectly identified species, making identifications of coelomycetous fungi
very difficult. Comparison to Type strain sequences, and use of confirmatory morphological studies will be
important.
Key features: Ostiolate pycnidia producing masses of slimy, hyaline, single-celled conidia.
References: Punithalingam (1979); McGinnis (1980); Sutton (1980); Rippon (1988); Montel et al. (1991);
Samson et al. (1995); de Hoog et al. (2015).
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
203
20 ˜m
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Fig. 75.1. Phoma spp. showing a pycnidium with an apical ostiole.
DOI: 10.1079/9781800622340.0075
Descriptions of Medical Fungi 204
Two species previously classified in the genus Candida have now been transferred to the genus Pichia. Pichia
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kudriavzevii (formerly Candida krusei) and Pichia norvegensis (formerly Candida norvegensis).
76.1. Pichia kudriavzevii Boidin et al.
Synonymy: Candida krusei (Castellani) Berkhout; Issatchenkia orientalis Kudryavtesev.
Pichia kudriavzevii is regularly associated with some forms of infant diarrhoea and occasionally with
systemic disease. It has also been reported to colonise the gastrointestinal, respiratory and urinary tracts of
patients with granulocytopenia. Environmental isolations have been made from beer, milk products, skin,
faeces of animals and birds.
RG-2 organism
Culture: Colonies (SDA) white to cream coloured, smooth, glabrous, yeast-like.
Microscopy: Predominantly small, elongated to ovoid blastoconidia, 2-5 x 4-5 μm.
India ink preparation: Negative - no capsules present.
Dalmau plate culture: Abundant long, wavy, branched pseudohyphae with elongated to ovoid blastoconidia,
budding off in verticillate branches.
Molecular identification: ITS sequencing recommended.
MALDI-ToF MS: Able to accurately identify this species.
Germ tube formation: Negative
Physiological tests: (+ Positive, – Negative, v Variable, w Weak, s Slow, n not done).
Fermentation:
Glucose + Sucrose – Lactose –
Galactose – Maltose – Trehalose –
Growth reactions:
Glucose + L-Sorbose – myo-Inositol –
Sucrose – L-Rhamnose – DL-Lactate +
Raffinose – D-Xylose – D-Gluconate –
Melibiose – L-Arabinose – 2-Keto-D-gluconate –
Galactose – D-Arabinose – D-Glucosamine +
Lactose – D-Ribose – N-Acetyl-D-glucosamine +
Trehalose – Glycerol + D-Glucuronate –
Maltose – Erythritol – Nitrate –
Melezitose – Ribitol – Urease –
Methyl-α-D-glucoside – Galactitol – 0.1% Cycloheximide v
Soluble starch – D-Mannitol – Growth at 37oC +
Cellobiose – D-Glucitol – Growth at 40oC +
© CAB International 2023. Descriptions of Medical Fungi (eds S. Kidd, C.Halliday and D. Ellis)
205
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