Studies in Mycology 74 (March 2013) Development of Aspergillus niger Jan Dijksterhuis and Han Wösten, editors CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands An institute of the Royal Netherlands Academy of Arts and Sciences Development of Aspergillus niger STUDIES IN MYCOLOGY 74, 2013 Studies in Mycology The Studies in Mycology is an international journal which publishes systematic monographs of filamentous fungi and yeasts, and in rare occasions the proceedings of special meetings related to all fields of mycology, biotechnology, ecology, molecular biology, pathology and systematics. For instructions for authors see www.cbs.knaw.nl. EXECUTIVE EDITOR Prof. dr dr hc Robert A. Samson, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] LAYOUT EDITOR Manon van den Hoeven-Verweij, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] SCIENTIFIC EDITORS Prof. dr Dominik Begerow, Lehrstuhl für Evolution und Biodiversität der Pflanzen, Ruhr-Universität Bochum, Universitätsstr. 150, Gebäude ND 44780, Bochum, Germany. E-mail: [email protected] Prof. dr Uwe Braun, Martin-Luther-Universität, Institut für Biologie, Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, D-06099 Halle, Germany. E-mail: [email protected] Dr Paul Cannon, CABI and Royal Botanic Gardens, Kew, Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K. E-mail: [email protected] Prof. dr Lori Carris, Associate Professor, Department of Plant Pathology, Washington State University, Pullman, WA 99164-6340, U.S.A. E-mail: [email protected] Prof. dr Pedro W. Crous, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Prof. dr David M. Geiser, Department of Plant Pathology, 121 Buckhout Laboratory, Pennsylvania State University, University Park, PA, U.S.A. 16802. E-mail: [email protected] Dr Johannes Z. Groenewald, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Prof. dr David S. Hibbett, Department of Biology, Clark University, 950 Main Street, Worcester, Massachusetts, 01610-1477, U.S.A. E-mail: [email protected] Dr Lorelei L. Norvell, Pacific Northwest Mycology Service, 6720 NW Skyline Blvd, Portland, OR, U.S.A. 97229-1309. E-mail: [email protected] Prof. dr Alan J.L. Phillips, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, Quinta de Torre, 2829-516 Caparica, Portugal. E-mail: [email protected] Dr Amy Y. Rossman, Rm 304, Bldg 011A, Systematic Botany & Mycology Laboratory, Beltsville, MD, U.S.A. 20705. E-mail: [email protected] Dr Keith A. Seifert, Research Scientist / Biodiversity (Mycology and Botany), Agriculture & Agri-Food Canada, KW Neatby Bldg, 960 Carling Avenue, Ottawa, ON, Canada K1A OC6. E-mail: [email protected] Prof. dr Hyeon-Dong Shin, Division of Environmental Science & Ecological Engineering, Korea University, Seoul 136-701, Korea. E-mail: [email protected] Dr Roger Shivas, Manager, Plant Biosecurity Science, Biosecurity Queensland, Department of Employment, Economic Development and Innovation, DEEDI, GPO Box 267, Brisbane, Qld 4001, Dutton Park 4102, Queensland, Australia. E-mail: [email protected] Dr Marc Stadler, InterMed Discovery GmbH, Otto-Hahn-Straße 15, D-44227 Dortmund, Germany. E-mail: [email protected] Prof. dr Jeffrey K. Stone, Department of Botany & Plant Pathology, Cordley 2082, Oregon State University, Corvallis, OR, U.S.A. 97331-2902. E-mail: [email protected] Dr Richard C. Summerbell, 27 Hillcrest Park, Toronto, Ont. M4X 1E8, Canada. E-mail: [email protected] Prof. dr Brett Summerell, Royal Botanic Gardens and Domain Trust, Mrs. Macquaries Road, Sydney, NSW 2000, Australia. E-mail: [email protected] Prof. dr Ulf Thrane, Department of Systems Biology, Center for Microbial Biotechnology, Technical University of Denmark, Søltofts Plads 221, DK-2800 Kgs. Lyngby, Denmark. E-mail: [email protected] Copyright 2013 CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. You are free to share — to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author"s moral rights. Publication date:15 March 2013 Published and distributed by CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. Internet: www.cbs.knaw.nl. E-mail: [email protected]. ISBN/EAN : 978-90-70351-93-9 Online ISSN : 1872-9797 Print ISSN : 0166-0616 Cover: Top from left to right: Very early conidiophore and aerial hyphae, formation of conidia on phialides, fully developed and young conidiophore. Bottom from left to right: Growing apical hypha with elongated organelles, older hyphae with septum and rounded vacuoles, aerial hyphae and stipe. Development of Aspergillus niger edited by Jan Dijksterhuis CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands and Han Wösten Microbiology and Kluyver Centre for Genomics of Industrial Fermentations, Utrecht University, Padualaan 8, 3584 CH Utrecht, The Netherlands CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands An institute of the Royal Netherlands Academy of Arts and Sciences Preface PREFACE This Issue of Studies in Mycology deals with vegetative growth and development of Aspergillus inThis general issue of Studiesand A. in Mycologyniger in deals particular with vegetative. he first growth description and during of Aspergillusthe First World dates War. backIn the tooriginal 1729. paper Pier we find: “In Antoniodevelopment Micheli of Aspergillus (1679 -in1737) general described and A. niger Aspergillus in particular. as one1913 ofZahorski 1400 was novel granted genera a patent of plantin the sUnited in his States on a NovaThe first plantarum description ofAspergillus genera (Figdates back1). Atos 1729. a clerical Pier Antonio, Micheli method recognized for producing the citric similarity acid by betweenfermenting sugarthe solutions Micheli (1679–1737) described Aspergillus as one of 1400 novel with Sterigmatocystis nigra. This is one of the many names that sporulating structure of the fungus and the aspergillum, a liturgical device used in the Catholic genera of plants in his Nova plantarum genera (Fig. 1). As a clerical, has been used to designate fungi of the black Aspergillus group. ChurchMicheli recognised to sprinkle the similarity holy water between during the sporulating a service. structure Zahorski, however, states that Sterigmatocystis differs distinctly of the fungus and the aspergillum, a liturgical device used in the from Aspergillus”. It was the American company Chas. Pfizer & Co. Catholic Church to sprinkle holy water during a service. Inc that started large scale production of citric acid in 1923 using surface cultures of the fungus. This issue starts with a review on molecular mechanisms underlying differentiation processes in the vegetative mycelium and Figduring 1. Early asexual pictures and sexual of Aspergillus development conidiophores of aspergilli as(Krijgsheld et depictedal. 2013). by ThePier other Antonio articles Micheli in this in 1729.issue focus on germination of conidia (van Leeuwen et al. 2013a, b), formation of heterogeneous Aspergillusmicro-colonies niger (van wasVeluw described et al. 2013) in and 1867 differentiation in in a sporulatingmanuscript colonies entitled of A. niger (Bleichrodt“Physiologie et al. 2013).des Fig 1. Early pictures of Aspergillus conidiophores as depicted by Pier Antonio mucMichelié indin 1729.ées” by the French botanist Philippe Edouard TheLéon articles van of vanTieghem Leeuwen. et Heal. show isolated that the RNAthis composition fungus from molded galls with the main aim to study theof productiondormant conidia of gallicis highly acid different by a from process that of of germinating conidia (i.e. of conidia during isotropic and polarised growth). The fungal fermentation. Gallic acid was important for a variety of applications including ink Aspergillus niger was described in 1867 in a manuscript transcriptome of conidia changes most dramatically during the first productionentitled “Physiologie. From des the mucédinées” molded gallsby the heFrench isolated botanist two fungitwo hours namely of germination Penicillium enabling initiationglaucum of protein and synthesisa and AspergillusPhilippe Edouard speci Léones van “with Tieghem. ornamented He isolated this spores fungus, fromthat wasrespiration. similar Theto Aspergillusantifungal natamycin glaucus, does neither but that,affect differential bymolded its blackishgalls with thecolor main that aim wasto study cons theerved production on ofdifferent gallic media,expression by ofthe genes mu norsk ygermination odor it producedof A. niger conidia and during the someacid by aother process characteristics, of fungal