Champignon du mois – Juin 2021

Juin 2021


Penicillium brevicompactum

REGNE : Fungi
PHYLUM : Dikarya, Ascomycota, Pezizomycotina
CLASSE : Eurotiomycetes
ORDRE : Eurotiales
FAMILLE : Aspergillaceae

Penicillium brevicompactum, qui appartient au sous-genre Penicillium, à la section Brevicompacta et à la série Brevicompacta (comprenant également les espèces P. bialowiezense, P. fennelliae, P. kongii et P. neocrassum) [1], est un champignon filamenteux environnemental non pathogène dont l’intérêt est essentiellement d’ordre industriel. C’est en effet à partir de ce champignon que fut découvert l’un des immunosuppresseurs les plus largement utilisés aujourd’hui dans les transplantations d’organes solides et de cellules souches hématopoïétiques, l’acide mycophénolique.

 

Cette moisissure saprophyte peut être isolée de substrats très divers, puisqu’on la retrouve dans le sol où elle se développe à partir des débris végétaux en décomposition, mais aussi dans les céréales et particulièrement le maïs, les aliments, les textiles ou les peintures. Il s’agit d’une moisissure cosmopolite, présente dans les régions tempérées ou subtropicales, comme dans les régions polaires [2]. Penicillium brevicompactum est en effet une espèce psychrophile, halotolérante et xérophile ; l’optimum thermique de croissance de P. brevicompactum est de 23°C, mais la croissance est possible pour des températures comprises entre -2°C et 30°C (2) ; de plus, cette moisissure est capable de croître dans des milieux de pH acide (jusqu’à pH 5) ou présentant une faible teneur en eau (0,78 aw) [3].

 

Penicillium brevicompactum fait partie des Penicillium à croissance lente. Macroscopiquement, les colonies formées sont planes et peu extensives, mesurant après 7 jours d’incubation, 20 à 30 mm de diamètre sur milieu CYA (Czapek Yeast Agar) et 12 à 22 mm de diamètre sur milieu MEA (Malt Extract Agar). A maturité, les colonies sont de couleur gris-vert ou jaune-vert au recto (Figure 1A), avec un verso crème à brun (Figure 1B).

Microscopiquement, les spores, toujours unicellulaires et globuleuses à sub-globuleuses, sont produites par des cellules spécialisées, les phialides, rassemblées au sommet de filaments aériens, les conidiophores. Ces derniers sont fins, cloisonnés et non dilatés à leur extrémité, ce qui les différencient des conidiophores des Aspergillus. Penicillium brevicompactum est un Penicillium triverticillé : les phialides sont en effet portées par des métules, elles-mêmes insérées au sommet du conidiophore par l’intermédiaire de branches. Les pénicilles sont très larges, avec des métules larges, renflées à leur sommet et densément groupées (Figure 2). Les phialides sont au nombre de 6 à 10 par métule en moyenne ; elles sont ampulliformes et mesurent de 6-10 µm de long sur 2,5 à 3 µm de large. Les conidies, de petite taille (2,6 à 3,2 µm de diamètre), sont produites par bourgeonnement au sommet des phialides ; elles sont hyalines à vertes à maturité, avec une paroi lisse ou très finement rugueuse [2].

Figure 1 : Aspect des colonies de Penicillium brevicompactum sur gélose de Sabouraud après 8 jours d’incubation à 20-25°C. A : recto ; B : verso.
Figure 2 : Morphologie microscopique de Penicillium brevicompactum (objectif x 40). Du filament végétatif, part un conidiophore fin et cloisonné portant à son sommet un pénicille triverticillé (avec deux branches, portant chacune plusieurs métules, elles-mêmes portant plusieurs phialides).

Penicillium brevicompactum produit différents métabolites, notamment de la brévianamide A, la brévicompanine C, les cyclodepsipeptides pétrosifungines A et B, de l’asperphénamate, et de la botryodiploidine, mais cette moisissure est surtout connue pour la production d’acide mycophénolique (MPA). La synthèse de ce dérivé de l’acide orsellinique (Figure 3) fait intervenir plusieurs enzymes, codées par différents gènes organisés en cluster, appelés mpa (pour mycophenolic acid). Ce cluster dont la composition a été défini par Regueira et al. [4] est centré sur le gène mpaC codant une polycétide synthase de type I partiellement réductrice, et comprend 6 autres gènes [4,5] :

  • mpaA codant une prényltransférase,
  • mpaB codant une protéine de fonction inconnue,
  • mpaDE codant une protéine de fusion naturelle comportant un domaine cytochrome P450 et un domaine hydrolase,
  • mpaF codant une protéine conférant au champignon la résistance au MPA,
  • mpaG codant une O-méthyltransférase,
  • mpaH codant une enzyme de clivage oxydatif.
Figure 3 : Synthèse de l’acide mycophénolique chez Penicillium brevicompactum, d’après Regueira et al. (2011) et Zhang et al. (2019).. Les gènes impliqués dans la synthèse de ce métabolite (indiqués en rouge, alors que les protéines correspondantes sont indiquées en bleu) sont organisés en cluster. PKS : polycétide synthétase ; SAT : domaine starter ; KS : domaine kétosynthase ; AT : Domaine acyltransférase ; DH : Domaine déshydratase ; ACP : Domaine acyl carrier protein ; cMT : Domaine carboxy-méthyltransférase ; TE : Domaine thioestérase. DHMB : acide 3,5-dihydroxy-7-(hydroxyméthyl)-6-méthylbenzoique ; DHMP : 3,5-dihydroxy-6-méthylphthalide ; FDHMP : 4-farnésyl-3,5-dihydroxy-6-méthylphthalide

L’acide mycophénolique est un inhibiteur spécifique et non compétitif de l’inosine 5’ monophosphate déshydrogénase, et bloque ainsi la synthèse de novo des bases puriques. D’abord décrit comme antibiotique (actif vis-à-vis de Staphylococcus aureus et Bacillus anthracis), le MPA présente également des propriétés antivirales et antifongiques [6,7], mais ce métabolite secondaire est surtout connu pour ses propriétés immunosuppressives [8]. L’acide mycophénolique présente un effet cytostatique plus marqué sur les lymphocytes B et T que sur les autres types cellulaires, la prolifération des lymphocytes étant essentiellement dépendante de la synthèse de novo pour les bases puriques. La molécule est maintenant largement utilisée sous forme de mycophénolate sodique ou d’une prodrogue, le mycophénolate mofétil (ester 2-morpholinoéthylique du MPA) en prévention du rejet de greffe.

Pour plus d’informations, consulter :

  1. Jens C. Frisvad JC, Samson RA. Polyphasic taxonomy of Penicillium subgenus Penicillium. A guide to identification of food and air-borne terverticillate Penicillia and their mycotoxins. Stud Mycol. 2004;49:1–174.
  2. Botton B, Larpent JP. Moisissures utiles et nuisibles: importance industrielle. 2ème édition revue et complétée. Collection Biotechnologies. Masson, Paris. 1990. 512 pp.
  3. Pitt JI. Penicillium and related genera. In: de Blackburn C (Ed.), Food Spoilage Microorganisms. Elsevier. 2006;437 –50.
  4. Regueira TB, Kildegaard KR, Hansen BG, Mortensen UH, Hertweck C, Nielsen J. Molecular basis for mycophenolic acid biosynthesis in Penicillium brevicompactum. Appl Environ Microbiol. 2011;77(9):3035–43.
  5. Zhang W, Du L, Qu Z, Zhang X, Li F, Li Z, et al. Compartmentalized biosynthesis of mycophenolic acid. Proc Natl Acad Sci U S A. 2019;116:13305–10.
  6. Abraham EP. The effect of mycophenolic acid on the growth of Staphylococcus aureus in heart broth. Biochem J. 1945;39(5):398–408.
  7. Noto T, Sawada M, Ando K, Koyama K. Some biological properties of mycophenolic acid. J Antibiot. 1969;22:165–9.
  8. Bentley R. Mycophenolic acid: A one hundred year odyssey from antibiotic to immunosuppressant. Chem Rev. 2000;100:3801–26.

Claire Hoffmann1, Solène Le Gal2, Gilles Nevez2 et Jean-Philippe Bouchara1
(Groupe d’Etude des Interactions Hôte-Pathogène (EA 3142), Université d’Angers – Université de Bretagne Occidentale, CHU d’Angers1, CHU de Brest2)

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