Textbook Physiology and Genetics Selected Basic and Applied Aspects 2Nd Edition Timm Anke Ebook All Chapter PDF

Physiology and Genetics Selected
Basic and Applied Aspects 2nd Edition

Timm Anke
Visit to download the full and correct content document:
https://textbookfull.com/product/physiology-and-genetics-selected-basic-and-applied-
aspects-2nd-edition-timm-anke/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...
Systemic Lupus Erythematosus: Basic, Applied and

Clinical Aspects 2nd Edition George Tsokos (Editor)
https://textbookfull.com/product/systemic-lupus-erythematosus-
basic-applied-and-clinical-aspects-2nd-edition-george-tsokos-
editor/
Neotropical Social Wasps: Basic and applied aspects

Fabio Prezoto
https://textbookfull.com/product/neotropical-social-wasps-basic-
and-applied-aspects-fabio-prezoto/
Basic Physiology for Anaesthetists 2nd Edition David

Chambers
https://textbookfull.com/product/basic-physiology-for-
anaesthetists-2nd-edition-david-chambers/
Advancing Frontiers in Mycology amp Mycotechnology

Basic and Applied Aspects of Fungi Tulasi Satyanarayana
https://textbookfull.com/product/advancing-frontiers-in-mycology-
amp-mycotechnology-basic-and-applied-aspects-of-fungi-tulasi-
satyanarayana/
Examination Questions and Answers in Basic Anatomy and
Physiology 2nd Edition Martin Caon
https://textbookfull.com/product/examination-questions-and-
answers-in-basic-anatomy-and-physiology-2nd-edition-martin-caon/
Basic and Applied Bone Biology 2nd Edition David B.

Burr
https://textbookfull.com/product/basic-and-applied-bone-
biology-2nd-edition-david-b-burr/
Basic and Applied Phytoplankton Biology Perumal

Santhanam
https://textbookfull.com/product/basic-and-applied-phytoplankton-
biology-perumal-santhanam/
Marathon Running Physiology Psychology Nutrition and

Training Aspects 1st Edition Christoph Zinner
https://textbookfull.com/product/marathon-running-physiology-
psychology-nutrition-and-training-aspects-1st-edition-christoph-
zinner/
The Heparins: Basic and Clinical Aspects 1st Edition

David Green
https://textbookfull.com/product/the-heparins-basic-and-clinical-
aspects-1st-edition-david-green/
Edited by
Karl Esser
THE MYCOTA
A Comprehensive Treatise on Fungi
as Experimental Systems for Basic and Applied Research
Physiology and Genetics

Selected Basic and Applied Aspects
XV
Second Edition
Timm Anke
Anja Schüffler
Volume Editors
The Mycota
Edited by
K. Esser
The Mycota
I Growth, Differentiation and Sexuality

1st edition ed. by J.G.H. Wessels and F. Meinhardt
2nd edition ed. by U. Kües and R. Fischer
3rd edition ed. by J. Wendland
II Genetics and Biotechnology
1st and 2nd edition ed. by U. Kück
III Biochemistry and Molecular Biology
1st and 2nd edition ed. by R. Brambl and G. Marzluf
3rd edition ed. by D. Hoffmeister
IV Environmental and Microbial Relationships
1st edition ed. by D. Wicklow and B. Söderström
2nd edition ed. by C.P. Kubicek and I.S. Druzhinina
3rd edition ed. by I.S. Druzhinina and C.P. Kubicek
V Plant Relationships
1st edition ed. by G. Carroll and P. Tudzynski
2nd edition ed. by H.B. Deising
VI Human and Animal Relationships
1st edition ed. by D.H. Howard and J.D. Miller
2nd edition ed. by A.A. Brakhage and P.F. Zipfel
VII Systematics and Evolution
1st edition ed. by D.J. McLaughlin, E.G. McLaughlin, and P.A. Lemke{
2nd edition ed. by D.J. McLaughlin and J.W. Spatafora
VIII Biology of the Fungal Cell
1st and 2nd edition ed. by R.J. Howard and N.A.R. Gow
IX Fungal Associations
1st and 2nd edition ed. by B. Hock
X Industrial Applications
1st edition ed. by H.D. Osiewacz
2nd edition ed. by M. Hofrichter
XI Agricultural Applications
1st and 2nd edition ed. by F. Kempken
XII Human Fungal Pathogens
1st edition ed. by J.E. Domer and G.S. Kobayashi
2nd edition ed. by O. Kurzai
XIII Fungal Genomics
1st edition ed. by A.J.P. Brown
2nd edition ed. by M. Nowrousian
XIV Evolution of Fungi and Fungal-Like Organisms
Ed. by S. Pöggeler and J. Wöstemeyer
XV Physiology and Genetics: Selected Basic and Applied Aspects
1st edition ed. by T. Anke and D. Weber
2nd edition ed. by T. Anke and A. Schüffler
The Mycota
A Comprehensive Treatise on Fungi as
Experimental Systems for Basic and Applied
Research
Edited by K. Esser
Physiology and Genetics

XV Selected Basic and Applied
Aspects
2nd Edition
Volume Editors:
Timm Anke and Anja Schüffler
Series Editor
Professor Dr. Dr. h.c. mult. Karl Esser
Allgemeine Botanik
Ruhr-Universität
44780 Bochum, Germany
Tel.: +49 (234)32-22211
Fax.: +49 (234)32-14211
e-mail: Karl.Esser@rub.de
Volume Editors
Professor Dr. Timm Anke

Institut für Biotechnologie und Wirkstoff-Forschung gGmbH (IBWF)
Erwin-Schrödinger-Str. 56
67663 Kaiserslautern, Germany
e-mail: anke@rhrk.uni-kl.de
Dr. Anja Schüffler

Institut für Biotechnologie und Wirkstoff-Forschung gGmbH (IBWF)
Erwin-Schrödinger-Str. 56
67663 Kaiserslautern, Germany
Tel.: +49 (631)31672-19
Fax: +49 (631)31672-15
e-mail: schueffler@ibwf.de
ISBN 978-3-319-71739-5 ISBN 978-3-319-71740-1 (eBook)

https://doi.org/10.1007/978-3-319-71740-1
Library of Congress Control Number: 2017964284

# Springer International Publishing AG 2009, 2018
This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the
material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation,
broadcasting, reproduction on microfilms or in any other physical way, and transmission or information
storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology
now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, express or implied, with respect to the material contained herein or for any errors
or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims
in published maps and institutional affiliations.
Printed on acid-free paper
This Springer imprint is published by Springer Nature

The registered company is Springer International Publishing AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
Karl Esser
(born 1924) is retired Professor of General Botany
and Director of the Botanical Garden at the Ruhr-
Universität Bochum (Germany). His scientific work
focused on basic research in classical and molecular
genetics in relation to practical application. His stud-
ies were carried out mostly on fungi. Together with
his collaborators he was the first to detect plasmids in
higher fungi. This has led to the integration of fungal
genetics in biotechnology. His scientific work was
distinguished by many national and international
honors, especially three honorary doctoral degrees.
Timm Anke
(born 1944) studied biochemistry at the University of
Tuebingen where he got his PhD with a dissertation on
the biosynthesis of fungal siderophores. In 1973, he
joined the group of Fritz Lipmann at the Rockefeller
University in New York City where he was investigat-
ing the biosynthesis of valinomycin, a streptomycete
ionophore. After his return to Tuebingen in 1975, he
started to build up a group searching for new antibio-
tics from basidiomycetes within the framework of
Hans Zaehners Collaborative Research Center (SFB
76) focusing on the chemistry and biology of micro-
organisms. In 1981, he became full professor of bio-
technology at the University of Kaiserslautern from
which he retired in 2010. In addition, he headed the
Institute of Biotechnology and Drug Research IBWF e.
V. in Kaiserslautern from 1998 to 2010. One of his
outstanding achievements in the field of antibiotic
research is the discovery of the strobilurins, a major
class of agricultural fungicides for which he was
awarded the Karl Heinz Beckurts Prize in 1996.
Anja Schüffler
(born 1981) studied biology at the University of Kai-
serslautern. Her PhD focused on the characterization
of fungal natural products with antimicrobial activity.
In 2010, she received a DFG fellowship and joined
Prof. Fenical’s group at the Scripps Institute for
Oceanography in San Diego, where she studied anti-
bacterial natural products from streptomycetes. In
2011, she started a research fellowship at the Institute
of Biotechnology and Drug Research (IBWF, Kaiser-
slautern). Her scientific work focuses on bioactive
natural products and assay development, the biosyn-
thesis of secondary metabolites as well as the isolation
and taxonomy of fungi of diverse habitats.
Series Preface
Mycology, the study of fungi, originated as a sub discipline of botany and was a
descriptive discipline, largely neglected as an experimental science until the early
years of this century. A seminal paper by Blakeslee in 1904 provided evidence for
self-incompatibility, termed “heterothallism,” and stimulated interest in studies
related to the control of sexual reproduction in fungi by mating-type
specificities. Soon to follow was the demonstration that sexually reproducing
fungi exhibit Mendelian inheritance and that it was possible to conduct formal
genetic analysis with fungi. The names Burgeff, Kniep and Lindegren are all
associated with this early period of fungal genetics research.
These studies and the discovery of penicillin by Fleming, who shared a Nobel
Prize in 1945, provided further impetus for experimental research with fungi.
Thus, began a period of interest in mutation induction and analysis of mutants
for biochemical traits. Such fundamental research, conducted largely with Neu-
rospora crassa, led to the one gene:one enzyme hypothesis and to a second Nobel
Prize for fungal research awarded to Beadle and Tatum in 1958. Fundamental
research in biochemical genetics was extended to other fungi, especially to
Saccharomyces cerevisiae, and by the mid-1960s fungal systems were much
favored for studies in eukaryotic molecular biology and were soon able to
compete with bacterial systems in the molecular arena.
The experimental achievements in research on the genetics and molecular
biology of fungi have benefited more generally studies in the related fields of
fungal biochemistry, plant pathology, medical mycology, and systematics. Today,
there is much interest in the genetic manipulation of fungi for applied research.
This current interest in biotechnical genetics has been augmented by the develop-
ment of DNA-mediated transformation systems in fungi and by an understanding
of gene expression and regulation at the molecular level. Applied research initia-
tives involving fungi extend broadly to areas of interest not only to industry but to
agricultural and environmental sciences as well.
It is this burgeoning interest in fungi as experimental systems for applied as
well as basic research that has prompted publication of this series of books under
the title The Mycota. This title knowingly relegates fungi into a separate realm,
distinct from that of either plants, animals, or protozoa. For consistency through-
out this series of volumes, the names adopted for major groups of fungi (repre-
sentative genera in parentheses) are as follows:
Pseudomycota
Division: Oomycota (Achlya, Phytophthora, Pythium)
Division: Hyphochytriomycota
viii Series Preface
Eumycota
Division: Chytridiomycota (Allomyces)
Division: Zygomycota (Mucor, Phycomyces, Blakeslea)
Division: Dikaryomycota
Subdivision: Ascomycotina
Class: Saccharomycetes (Saccharomyces, Schizosaccharomyces)
Class: Ascomycetes (Neurospora, Podospora, Aspergillus)
Subdivision: Basidiomycotina
Class: Heterobasidiomycetes (Ustilago, Tremella)
Class: Homobasidiomycetes (Schizophyllum, Coprinus)
We have made the decision to exclude from The Mycota the slime molds which,
although they have traditional and strong ties to mycology, truly represent
nonfungal forms insofar as they ingest nutrients by phagocytosis, lack a cell
wall during the assimilative phase, and clearly show affinities with certain proto-
zoan taxa.
The series throughout will address three basic questions: what are the fungi,
what do they do, and what is their relevance to human affairs? Such a focused and
comprehensive treatment of the fungi is long overdue in the opinion of the
editors.
A volume devoted to systematics would ordinarily have been the first to
appear in this series. However, the scope of such a volume, coupled with the
need to give serious and sustained consideration to any reclassification of major
fungal groups, has delayed early publication. We wish, however, to provide a
preamble on the nature of fungi, to acquaint readers who are unfamiliar with
fungi with certain characteristics that are representative of these organisms and
which make them attractive subjects for experimentation.
The fungi represent a heterogeneous assemblage of eukaryotic microorgan-
isms. Fungal metabolism is characteristically heterotrophic or assimilative for
organic carbon and some nonelemental source of nitrogen. Fungal cells charac-
teristically imbibe or absorb, rather than ingest, nutrients and they have rigid cell
walls. The vast majority of fungi are haploid organisms reproducing either
sexually or asexually through spores. The spore forms and details on their
method of production have been used to delineate most fungal taxa. Although
there is a multitude of spore forms, fungal spores are basically only of two types:
(1) asexual spores are formed following mitosis (mitospores) and culminate
vegetative growth, and (2) sexual spores are formed following meiosis (meios-
pores) and are borne in or upon specialized generative structures, the latter
frequently clustered in a fruit body. The vegetative forms of fungi are either
unicellular, yeasts are an example, or hyphal; the latter may be branched to
form an extensive mycelium.
Regardless of these details, it is the accessibility of spores, especially the direct
recovery of meiospores coupled with extended vegetative haploidy, that have
made fungi especially attractive as objects for experimental research.
The ability of fungi, especially the saprobic fungi, to absorb and grow on rather
simple and defined substrates and to convert these substances, not only into
essential metabolites but into important secondary metabolites, is also noteworthy.
The metabolic capacities of fungi have attracted much interest in natural products
Series Preface ix
chemistry and in the production of antibiotics and other bioactive compounds.

Fungi, especially yeasts, are important in fermentation processes. Other fungi are
important in the production of enzymes, citric acid, and other organic compounds
as well as in the fermentation of foods.
Fungi have invaded every conceivable ecological niche. Saprobic forms
abound, especially in the decay of organic debris. Pathogenic forms exist with
both plant and animal hosts. Fungi even grow on other fungi. They are found in
aquatic as well as soil environments, and their spores may pollute the air. Some
are edible; others are poisonous. Many are variously associated with plants as
copartners in the formation of lichens and mycorrhizae, as symbiotic endophytes
or as overt pathogens. Association with animal systems varies; examples include
the predaceous fungi that trap nematodes, the microfungi that grow in the
anaerobic environment of the rumen, the many insect associated fungi, and the
medically important pathogens afflicting humans. Yes, fungi are ubiquitous and
important. There are many fungi, conservative estimates are in the order of
100,000 species, and there are many ways to study them, from descriptive
accounts of organisms found in nature to laboratory experimentation at the
cellular and molecular level. All such studies expand our knowledge of fungi
and of fungal processes and improve our ability to utilize and to control fungi for
the benefit of humankind.
We have invited leading research specialists in the field of mycology to
contribute to this series. We are especially indebted and grateful for the initiative
and leadership shown by the Volume Editors in selecting topics and assembling
the experts. We have all been a bit ambitious in producing these volumes on a
timely basis and therein lies the possibility of mistakes and oversights in this first
edition. We encourage the readership to draw our attention to any error, omis-
sion, or inconsistency in this series in order that improvements can be made in
any subsequent edition.
Finally, we wish to acknowledge the willingness of Springer-Verlag to host
this project, which is envisioned to require more than 5 years of effort and the
publication of at least nine volumes.
Bochum, Germany KARL ESSER

Auburn, AL, USA PAUL A. LEMKE
April 1994 Series Editors
Volume Preface
More than 120,000 different fungal species have been described, and it is esti-
mated that there exist more than 1.5 106 species. Fungi have adopted many
different ways of living in very diverse habitats as saprophytes, pathogens,
symbionts or endophytes. Fungi and their products are used for the fermentation
and processing of food and feeds, for biological control and for the production of
vitamins and amino acids. Some of their secondary metabolites are used in
medicine, e.g. as antibiotics, immunosuppressants, cholesterol-lowering drugs
or agrochemical fungicides. Recently, progress in the field of mycology has been
substantial due to new methodological approaches and technologies, many of
them DNA-based, strongly adding to the motivation to compile a new volume of
Mycota XV Physiology and Genetics: Selected Basic and Applied Aspects.
Chapter “Fruiting Body Development in Ascomycetes” provides a general
overview about the morphology and development of fruiting bodies in ascomy-
cetes with discussion of regulatory networks such as signal transduction path-
ways, protein degradation mechanisms as well as transcriptional regulators and
chromatin modifiers. Chapter “Fungal Inteins: Distribution, Evolution and Appli-
cations” summarizes the current knowledge of inteins, their occurrence, evolu-
tion and application. Inteins are internal protein sequences which are excised
from a precursor protein (extein) whose N- and C-termini are subsequently
ligated to yield two stable proteins, the mature protein and the intein. In the
interaction of yeasts, killer toxins play an important role. Their structures, modes
of action and resistance as well as possible applications are discussed in chapter
“Yeast Killer Toxins: Fundamentals and Applications”. Chapter “The Fungal
MCC/Eisosome Complex: An Unfolding Story” deals with the fungal MCC/eiso-
some complex which plays an important role in plasma membrane organization
and diverse plasma membrane-associated processes. In chapter “The Genus
Periglandula and Its Symbiotum with Morning Glory Plants (Convolvulaceae)”,
the enigma of why ergot alkaloids are equally present in fungi (Clavicipitaceae)
and higher plants (Convolvulaceae) is addressed and solved. Chapter “Volatiles
in Communication of Agaricomycetes” presents a comprehensive survey on
communication activities of Agaricomycetes on all organismal scales and com-
munity levels in which signalling by fungal volatile organic compounds (VOCs) is
recognized. The substantial progress in elucidating the lifestyle, metabolism and
genetics of endophytic fungi is addressed in the chapter “Endophytic Fungi,
Occurrence, and Metabolites”. Basidiomycetes are a rich source of unique sec-
ondary metabolites in most cases not found in other fungi. The chapter “Second-
ary Metabolites of Basidiomycetes” offers a survey of new compounds isolated
since 2008. Chapter “Identification of Fungicide Targets in Pathogenic Fungi”
xii Volume Preface
covers the current molecular targets of antifungal compounds and discusses

future directions of fungicide research. Helminths can pose serious problems to
animal and human health. It is therefore quite remarkable that fungi produce low
molecular weight compounds specifically interfering with reactions not present in
mammalian hosts, thus paving the way for selective nontoxic medications or
agrochemicals. For the discovery of avermectin, successfully used in the therapy
of roundworm parasites, Prof. Satoshi Omura, author of chapter “Helminth
Electron Transport Inhibitors Produced by Fungi”, was awarded the Nobel
Prize in Physiology or Medicine in 2015. Chapter “Cyclic Peptides and Depsipep-
tides from Fungi” describes the occurrence, structures and biological activities of
peptides and depsipeptides produced by fungi and discusses the importance of
these compounds as lead compounds for agricultural and pharmaceutical appli-
cations.
In chapter “Polyketide Synthase–Nonribosomal Peptide Synthetase Hybrid
Enzymes of Fungi”, recent efforts in engineering-selected fungal species to make
them amenable to efficient genetic modifications for identifying genes responsi-
ble for the biosynthesis of secondary metabolites are addressed. This review also
discusses how the engineered fungi are used in deciphering the mechanism of
natural product biosynthesis, primarily through heterologous reconstitution of
biosynthetic pathways of interest. Fungal polyketides are among the prominent
fungal metabolites. As addressed in chapter “Biosynthesis of Fungal Polyketides”,
their biosynthesis is increasingly well understood at chemical, biochemical and
genetic levels, thus offering a chance to obtain sufficient quantities of complex but
potentially valuable therapeutics. The mycotoxins ochratoxin, citrinin and patu-
lin are often found as contaminants of foods. New results concerning the regula-
tion and the simultaneous occurrence of ochratoxin, citrinin and patulin
producing penicillia in certain habitats are presented in chapter “Aspects of the
Occurrence, Genetics and Regulation of Biosynthesis of the Three Food Relevant
Penicillium Mycotoxins: Ochratoxin A, Citrinin and Patulin”.
We do hope that readers enjoy reading this volume of The Mycota. We are
very grateful to the contributing authors, whose expertise and efforts have made
this project possible. We thank Dr. Andrea Schlitzberger of Springer Verlag for
her support and engagement during the preparation of this volume.
Kaiserslautern and Mannheim, Germany Timm Anke

October 2017 Anja Schüffler
Contents
Fruiting-Body Development in Ascomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

S. PÖGGELER, M. NOWROUSIAN, I. TEICHERT, A. BEIER, U. KÜCK
Fungal Inteins: Distribution, Evolution, and Applications . . . . . . . . . . . . . 57

SKANDER ELLEUCHE, STEFANIE PÖGGELER
Yeast Killer Toxins: Fundamentals and Applications . . . . . . . . . . . . . . . . . . . 87

RAFFAEL SCHAFFRATH, FRIEDHELM MEINHARDT, ROLAND KLASSEN
The Fungal MCC/Eisosome Complex: An Unfolding Story . . . . . . . . . . . 119

KRISZTINA KOLLÁTH-LEIß, FRANK KEMPKEN
The Genus Periglandula and Its Symbiotum with Morning Glory Plants
(Convolvulaceae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
E. LEISTNER, U. STEINER
Volatiles in Communication of Agaricomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . 149

URSULA KÜES, WEERADEJ KHONSUNTIA, SHANTA SUBBA, BASTIAN DÖRNTE
Endophytic Fungi, Occurrence, and Metabolites . . . . . . . . . . . . . . . . . . . . . . . . 213

MYSORE V. TEJESVI, ANNA MARIA PIRTTILÄ
Secondary Metabolites of Basidiomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231

ANJA SCHÜFFLER
Identification of Fungicide Targets in Pathogenic Fungi . . . . . . . . . . . . . . . 277

ANDREW J. FOSTER
Helminth Electron Transport Inhibitors Produced by Fungi . . . . . . . . . . . 297

MIHOKO MORI, KENICHI NONAKA, ROKURO MASUMA, SATOSHI ŌMURA,
KAZURO SHIOMI
Cyclic Peptides and Depsipeptides from Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . 331

HEIDRUN ANKE, HARTMUT LAATSCH
Polyketide Synthase–Nonribosomal Peptide Synthetase Hybrid Enzymes

of Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
SHINJI KISHIMOTO, YUICHIRO HIRAYAMA, KENJI WATANABE
xiv Contents
Biosynthesis of Fungal Polyketides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385

RUSSELL J. COX, ELIZABETH SKELLAM, KATHERINE WILLIAMS
Aspects of the Occurrence, Genetics, and Regulation of Biosynthesis of the

Three Food Relevant Penicillium Mycotoxins: Ochratoxin A, Citrinin,
and Patulin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 413
ROLF GEISEN, MARKUS SCHMIDT-HEYDT, DOMINIC STOLL, NAJIM TOUHAMI
Biosystematic Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435
Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447

List of Contributors
HEIDRUN ANKE
Institute for Biotechnology and Drug Research, IBWF gGmbH, Kaiserslautern,
Germany
ANNA BEIER
Lehrstuhl für Allgemeine und Molekulare Botanik, Ruhr-Universität, Bochum,
Bochum, Germany
RUSSELL J. COX
Institute for Organic Chemistry and Biomolekulares Wirkstoffzentrum
(BMWZ), Leibniz Universität Hannover, Hannover, Germany
BASTIAN DÖRNTE
Molecular Wood Biotechnology and Technical Mycology, University of
Goettingen, Göttingen, Germany
SKANDER ELLEUCHE
Miltenyi Biotec GmbH, Bergisch Gladbach, Germany
ANDREW J. FOSTER
University of Exeter, Biosciences, Exeter, UK
ROLF GEISEN
Max Rubner-Institut, Karlsruhe, Germany
YUICHIRO HIRAYAMA
Department of Pharmaceutical Sciences, University of Shizuoka, Shizuoka, Japan
FRANK KEMPKEN
Botanisches Institut, Christian-Albrechts-Universität zu Kiel, Kiel, Germany
WEERADEJ KHONSUNTIA
SHINJI KISHIMOTO
xvi List of Contributors
ROLAND KLASSEN
Fachgebiet Mikrobiologie, Institut für Biologie, Universität Kassel, Kassel,
Germany
KRISZTINA KOLLÁTH-LEIß
Botanisches Institut, Christian-Albrechts-Universität zu Kiel, Kiel, Germany
ULRICH KÜCK
Bochum, Germany
URSULA KÜES
HARTMUT LAATSCH
University of Goettingen, Institute of Organic and Biomolecular Chemistry,
Goettingen, Germany
ECKHARD LEISTNER
Institut für Pharmazeutische Biologie, Rheinische Friedrich Wilhelm-Universität
Bonn, Bonn, Germany
ROKURO MASUMA
The Kitasato Institute for Life Sciences and Graduate School of Infection Control
Sciences, Kitasato University, Tokyo, Japan
FRIEDHELM MEINHARDT
Institut für Molekulare Mikrobiologie und Biotechnologie, Westfälische
Wilhelms-Universität Münster, Münster, Germany
MIHOKO MORI
KENICHI NONAKA
MINOU NOWROUSIAN
Bochum, Germany
SATOSHI ŌMURA
List of Contributors xvii
STEFANIE PÖGGELER
Abteilung Genetik eukaryotischer Mikroorganismen, Institut für Mikrobiologie
und Genetik, Georg-August-Universität Göttingen, Göttingen, Germany
Göttingen Center for Molecular Biosciences (GZMB), Georg-August-University
Göttingen, Göttingen, Germany
ANNA MARIA PIRTTILÄ

Genetics and Physiology, University of Oulu, Oulu, Finland
ANJA SCHÜFFLER
Institut für Biotechnologie und Wirkstoff-Forschung gGmbH, Kaiserslautern,
Germany
RAFFAEL SCHAFFRATH
Fachgebiet Mikrobiologie, Institut für Biologie, Universität Kassel, Kassel,
Germany
MARKUS SCHMIDT-HEYDT
KAZURO SHIOMI
ELIZABETH SKELLAM
ULRIKE STEINER
Institut für Nutzpflanzenwissenschaften und Ressourcenschutz
(INRES), Rheinische Friedrich Wilhelm-Universität Bonn, Bonn, Germany
DOMINIC STOLL
SHANTA SUBBA
INES TEICHERT
Bochum, Germany
MYSORE V. TEJESVI
Genetics and Physiology, University of Oulu, Oulu, Finland
NAJIM TOUHAMI
xviii List of Contributors
KENJI WATANABE
KATHERINE WILLIAMS
Fruiting-Body Development in Ascomycetes
S. PÖGGELER1, M. NOWROUSIAN2, I. TEICHERT2, A. BEIER2, U. KÜCK2
CONTENTS 2. Transcriptional Changes During

Development . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
I. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
3. Chromatin Modifiers . . . . . . . . . . . . . . . . . . . 39
A. Fungal Sexual Development . . . . . . . . . . . . . . . 2
V. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
B. Fruiting-Body Morphology . . . . . . . . . . . . . . . . 5
II. Systems to Study Fruiting-Body References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
A. Neurospora crassa . . . . . . . . . . . . . . . . . . . . . . . . . 5
B. Podospora anserina . . . . . . . . . . . . . . . . . . . . . . . 8
C. Sordaria macrospora . . . . . . . . . . . . . . . . . . . . . . 8 I. Introduction
D. Aspergillus nidulans . . . . . . . . . . . . . . . . . . . . . . 8
E. Emerging Model Systems . . . . . . . . . . . . . . . . . . 9 Fruiting bodies are multicellular structures,
1. Aspergillus fumigatus, A. flavus, and
A. parasiticus . . . . . . . . . . . . . . . . . . . . . . . . . . . 9 which protect the products of meiosis, the sex-
2. Botrytis cinerea . . . . . . . . . . . . . . . . . . . . . . . . . 9 ual spores. They occur during the sexual life
3. Trichoderma reesei . . . . . . . . . . . . . . . . . . . . . . 9 cycle of the Dikarya, a group that encompasses
4. Penicillium species . . . . . . . . . . . . . . . . . . . . . . 9 the ascomycetes and basidiomycetes (Hibbett
5. Fusarium graminearum and et al. 2007; Peraza-Reyes and Malagnac 2016)
F. verticillioides . . . . . . . . . . . . . . . . . . . . . . . . . 9
6. Pyronema confluens . . . . . . . . . . . . . . . . . . . . . 10
(Fig. 1). However, only filamentous species
III. Factors Influencing Fruiting-Body show the development of fruiting bodies,
Development . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 while yeasts never exhibit comparable struc-
A. Environmental Factors . . . . . . . . . . . . . . . . . . . . 10 tures.
1. Light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 In this chapter, which is an extension and
2. Nutrients . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
B. Endogenous Factors . . . . . . . . . . . . . . . . . . . . . . . 14
update of a previous review in this series (Pög-
1. Primary and Secondary Metabolites . . . . 15 geler et al. 2006b), we will give an overview of
2. Pheromones . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 the development of fruiting bodies in ascomy-
IV. Regulatory Networks . . . . . . . . . . . . . . . . . . . . . . . . 21 cetes, including an outline of some model
A. Signal Transduction Pathways . . . . . . . . . . . . 21 ascomycetes, which have been used to study
1. MAP Kinase Signaling . . . . . . . . . . . . . . . . . . 21
2. STRIPAK . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
fruiting-body development at the molecular
3. NOX Complexes . . . . . . . . . . . . . . . . . . . . . . . . 25 level. Further, we will summarize factors that
B. Protein Degradation . . . . . . . . . . . . . . . . . . . . . . 25 can either be environmental or endogenous,
1. Autophagy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 which control this process. Finally, regulatory
2. Proteasomal Degradation . . . . . . . . . . . . . . . 29 networks will be mentioned that govern
C. Transcriptional Regulation . . . . . . . . . . . . . . . . 32
fruiting-body development. This includes sig-
1. Transcription Factors . . . . . . . . . . . . . . . . . . . 32
nal transduction pathways, protein degradation
mechanisms, and transcriptional regulatory
1
Institut für Mikrobiologie und Genetik, Genetik eukaryo- networks. Ultimately, we observe that novel
tischer Mikroorganismen, Universität Göttingen, Göttingen,
experimental approaches such as quantitative
Germany; e-mail: spoegge@gwdg.de
2
Lehrstuhl für Allgemeine und Molekulare Botanik, Ruhr-
mass spectrometry, functional genomics, or
Universität, Bochum, Bochum, Germany; e-mail: ulrich. super resolution microscopy have begun to
kueck@rub.de improve our knowledge about the mechanistic
Physiology and Genetics, 2nd Edition

The Mycota XV
T. Anke, A. Schüffler (Eds.)
© Springer International Publishing AG 2018
2 S. Pöggeler et al.
Fig. 1 Phylogenetic tree of Ascomycota. Characteristic sphaerella graminicola, Neosartorya fischeri, Neuros-
species are given as examples. Branch lengths are pro- pora crassa, Phaeosphaeria nodorum, Pyronema
portional to genetic distances [adapted from Traeger confluens, Saccharomyces cerevisiae, Schizosaccharo-
et al. (2013)]. Species used to construct the phyloge- myces pombe, Sclerotinia sclerotiorum, Sordaria macro-
netic tree: Agaricus bisporus, Arthrobotrys oligospora, spora, Tuber melanosporum, Yarrowia lipolytica. S.
Blumeria graminis, Coccidioides immitis, Aspergillus pombe, L. bicolor, and A. bisporus served as outgroups
nidulans, Gibberella zeae, Laccaria bicolor, Myco-
processes that lead to the formation of multi- netic mating partners (Peraza-Reyes and
cellular structures. Malagnac 2016). As a result of meiotic divi-
sions, fungi produce four haploid spores,
which may be doubled or multiplied by one or
A. Fungal Sexual Development several postmeiotic mitoses.
The ascus is the meiosporangium of the
Fungi propagate either asexually or sexually. Ascomycota. These sac-like sporangia carry
Asexual propagation is characterized by mitotic the ascospores, the products of meiosis. In
divisions, and as a result, endospores within mycelial ascomycetes, asci are usually formed
sporangia or exospores like conidia are gener- inside developmentally complex fruiting bodies
ated. In contrast, sexual propagation is charac- that are called the ascomata or ascocarps. In
terized by karyogamy and meiotic divisions, contrast to filamentous ascomycetes, ascos-
and fungi share this feature with most other pores of unicellular ascomycetes (yeasts) are
eukaryotes. Generally, sexual reproduction is never found in fruiting bodies. The develop-
thought to be the source of genetic diversity. ment of fruiting bodies is a rather complex
During meiotic divisions, recombination cellular process that requires special environ-
occurs between chromosomes of two heteroge- mental and genetic conditions, which control
Fruiting-Body Development in Ascomycetes 3
the expression of developmentally regulated DOPA (L-3,4-dihydroxyphenylalanine) path-

genes. Fruiting bodies are highly complex ways. Some can also be generated by the
structures, which contain several different tis- L-tyrosine degradation pathway (Langfelder
sues protecting the asci. For example, 15 differ- et al. 2003). One of the best-characterized mel-
ent cell types were recognized in fruiting bodies anin biosynthetic pathways is the DHN melanin
of the Sordariomycete Neurospora crassa (Bistis pathway, which has been verified for many
et al. 2003). For a coordinated fruiting-body members of the Pezizomycotina. Melanins sta-
development, enzymes involved in cell wall bio- bilize the cell wall and provide protection
genesis and metabolism are required, as well as against UV light-induced DNA damage. An
genes responsible for the cytoskeleton structure investigation with S. macrospora showed that
and organization. Here we will mention some expression of melanin biosynthesis genes is
representative examples, and a more detailed correlated with fruiting-body development.
description on this subject can be found in our For example, melanin gene expression is highly
previous review (Pöggeler et al. 2006a). The repressed in submerged cultures, where no sex-
ami1 gene from Podospora anserina, for exam- ual development occurs. Similarly, sterile
ple, is necessary for nuclear positioning, most mutants of S. macrospora showed only reduced
likely by regulating components of the dynein transcript levels of melanin biosynthesis genes
pathway. This gene was shown to be responsi- (Engh et al. 2007). Finally, mutants with a defect
ble for male fertility, and deletion results in a in melanin biosynthesis from Ophiostoma pili-
delayed formation of fruiting bodies in the ferum and Podospora anserina showed defects
corresponding mutants (Bouhouche et al. in the formation of fruiting bodies. These
2004). The outer shell of the fruiting body, the observations are consistent with early reports
peridium, is an essential structure to protect the for N. crassa, P. anserina, and Tuber species,
meiosporangia with the ascospores. The perid- where correlation between melanin biosynthe-
ium consists of bundles of filamentous cells, sis and the reproductive cycle was suggested
and their cell walls have three main constitu- (Hirsch 1954; Esser 1966; Prade et al. 1984;
ents, namely, chitin, mannan, and b-glucan. Ragnelli et al. 1992; Teichert and Nowrousian
Though the related biosynthetic pathways 2011).
have intensively been investigated, it has not Very important proteins of the cell wall are
been demonstrated with certainty that the the hydrophobins and lectins. Although they
corresponding genes are preferentially have mainly been characterized in higher basi-
expressed in fruiting-body tissues. For Sordaria diomycetes, where they are implied in mush-
macrospora functional analysis of the class VII room formation, they have also a function in
(division III) chitin synthase gene (chs7) has the Pezizomycotina. Cryparin, a class II hydro-
shown that it is dispensable for fruiting-body phobin, was found mainly in the cell walls
formation, but the corresponding mutant dis- of fruiting bodies from the chestnut blight
played sensitivity toward cell wall stress (Trae- pathogen, Cryphonectria parasitica. Deletion
ger and Nowrousian 2015). Another result mutants lacking the cryparin gene were unable
comes from Tuber borchii, where three genes to generate wild-type-like fruiting bodies. Thus,
for chitin synthesis were investigated. Albeit this pathogen needs hydrophobins for its fit-
they are constitutively expressed in vegetative ness under natural conditions (Kazmierczak
mycelium, they show a differential expression et al. 2005).
in sporogenic or vegetative tissue of the fruiting On the genetic level, there are many genes
bodies (Balestrini et al. 2000). In contrast, sev- regulating the sexual cycle of ascomycetes.
eral chs mutants from N. crassa and Aspergillus Important master genes involved in the general
nidulans show severe defects in perithecial control of sexual development are part of the
development (Fajardo-Somera et al. 2015). mating-type loci. They have been found so far in
Important pigments of the cell walls are all ascomycetes, irrespective of whether they
melanins. They are synthesized either through produce fruiting bodies or not. Their regulatory
the DHN (1,8-dihydroxynaphthalene) or the role during the sexual cycle has been thoroughly
Fig. 2 Examples of mating-type loci of heterothallic conserved genes, such as sla2 (cytoskeleton assembly
members of the Dothideomycetes (Cochliobolus hetero- control factor) and apn2 (DNA lyase). An exception is
strophus) (Wirsel et al. 1998), Eurotiomycetes (Penicil- the MAT locus from C. heterostrophus with the following
lium chrysogenum) (Böhm et al. 2013, 2015), and flanking genes: GAP, GTPase-activating protein; ß-Glc,
Sordariomycetes (N. crassa). For comparison the ß-glucosidase. Abbreviations: “a” and “HMG” indicate
mating-type locus encoding four open reading frames genes encoding transcription factors with conserved
from the homothallic fungus S. macrospora (Sordario- DNA-binding domains
mycetes) is shown. The flanking regions often carry
studied in the yeasts Saccharomyces cerevisiae tem is found, e.g., in P. anserina or Neurospora
and Schizosaccharomyces pombe, which pro- tetrasperma.
duce asci, but no fruiting bodies. The role of Usually the mating-type loci of heterothallic
mating-type genes during fruiting-body devel- species contain dissimilar sequences, albeit they
opment is by far less well understood, although are located at identical chromosomal positions.
some studies have shown that mating-type Thus, mating-type loci do not represent alleles
genes are directly involved in fruiting-body of a given gene but rather dissimilar DNA
development (Nelson and Metzenberg 1992; sequences which are called idiomorphs. MAT
Pöggeler et al. 1997) (see also Sect. IV.C.1). In loci from the Pezizomycotina carry one or
general, two types of fungal breeding systems more open reading frames of which at least
are distinguished. Heterothallism involves two one codes for a mating-type transcription factor
individuals with opposing mating types, while (TF). In general, the MAT1-1 locus of hetero-
homothallism refers to sexual reproduction by thallic species contains one to three open
selfing. In the latter case, individual strains do reading frames, while only a single gene is
not need a mating partner to propagate sexually. found in MAT1-2 loci. In contrast to baker’s
Pseudohomothallism finally can be considered yeast, species of the Pezizomycotina carry no
to be an exceptional type of heterothallism. The silent mating-type loci. Thus, mating-type
term was used for species that contain asci with switching as observed in yeast does usually not
four ascospores, each carrying two nuclei with occur in heterothallic filamentous ascomycetes.
opposite mating-type genes. Thus, after germi- Mating-type loci encode TFs that are
nation, these resulting heterokaryotic mycelia directly involved in the sexual life cycle. Figure 2
can undergo selfing. This type of breeding sys- displays the general structure of mating-type
loci from members of the Eurotiomycetes, the 4. Apothecia are open to cup-shaped fruiting
Dothideomycetes, and the Sordariomycetes. The bodies that have a hymenium layer on their
MAT1-1-1 gene encodes a TF that is character- surface carrying the asci. The spores are
ized by an a DNA-binding domain, while the actively discharged, and examples of spe-
MAT1-2-1 gene codes for TFs with a high- cies that have apothecia are Ascobolus
mobility group (HMG) DNA-binding domain. immersus, Pyronema confluens, and Morch-
A detailed description of mating-type locus- ella sp. within the Pezizomycetes and
encoded TFs is given in Sect. IV.C.1. Botrytis cinerea within the Leotiomycetes.
B. Fruiting-Body Morphology
II. Systems to Study Fruiting-Body
During their sexual life cycle, filamentous fungi
of subdivision Pezizomycotina generate fruit- Development
ing bodies that were historically used for their
taxonomic classification. Current classification Fruiting-body development has been studied in
systems that rely on molecular data show that a wide range of different ascomycetous species.
these conventional classifications contain non- Here we describe four model systems, which
monophyletic groups (Schoch et al. 2009; were used intensively for investigations on
Ebersberger et al. 2012). However, different fruiting-body development. Further, we will
fruiting-body morphologies are important mention some emerging model ascomycetes
traits in fungal ecology, and the foremost com- that were used recently for studying specific
mon types of fruiting bodies (Esser 1982) are aspects of the sexual life cycle, including
described below (Figs. 3 and 4). fruiting-body formation.
1. Cleistothecia are closed, spherical fruiting
bodies that distribute the ascospores after
A. Neurospora crassa
disintegration of the peridium of the fruit-
ing bodies. Typically, members of the The model fungus Neurospora crassa is a het-
Eurotiomycetes such as Aspergillus fumiga- erothallic species of the Sordariaceae and has a
tus, Aspergillus nidulans, and Penicillium rather complex sexual life cycle. In general, two
chrysogenum generate cleistothecia. mating types can be distinguished, which are
2. Pseudothecia are spherical fruiting bodies called “A” (MAT1-1) and “a” (MAT1-2). Both
that contain cavities (loculi) that contain strains generate macro- and microconidia,
the gametangia. Spores are actively dis- which can be considered as male gametangia.
charged through openings which arise In addition, both strains form female gametan-
from local lysis of the peridium. Pseudothe- gia that are called ascogonia. Female gametan-
cia are, for example, found in the Dothideo- gia are surrounded by supporting hyphae,
mycetes, e.g., Venturia inaequalis. which after 2–3 days generate a protoperithe-
3. Perithecia are closed flask-like fruiting cium (young fruiting body). During fertiliza-
bodies that look similar to the pseudothe- tion, the female gametangium generates a
cia. Within perithecia, sterile hyphae are distinct uptake hypha called trichogyne. This
found that enclose the generative tissue trichogyne will fuse with male gametes, which
(hymenium). The hymenium generates can be macro- or microconidia as described
asci with usually eight ascospores, which above. Alternatively, a fusion with vegetative
are actively discharged from the perithe- hyphae has also been described. However,
cium through a preformed opening, the self-fertilization of each strain is prevented by
ostiole. Perithecia are typical fruiting bod- an incompatibility mechanism. Thus, tricho-
ies of members of the Sordariomycetes, gynes can only fuse with male gametes from
such as N. crassa, P. anserina, and S. macro- an opposite mating-type partner. In summary,
spora. the protoperithecium with a trichogyne from
Fig. 3 Typical fruiting bodies of ascomycetes. (a) Peri- spora; the arrow points to the ostiolum that is used to
thecia of the homothallic Sordariomycete S. macro- discharge the eight-spored asci. (b) Cleistothecia from
Fig. 4 Asci from different fungi of the Pezizomycotina. one shows ascospores with several nuclei, which appear
(a) Ascus from the Dothideomycete Keissleriella quad- after several mitotic divisions. (d) Asci from the fungus
riseptata [from Tanaka et al. (2015)]. (b) Asci from E. crustaceum. (e) Ascospores from E. crustaceum. (f)
Sordaria macrospora, obtained from a cross between a Asci from Podospora anserina contain four spores or
wild type (black spores) and a spore color mutant (lu five spores. Usually, each ascospore carries two nuclei;
with yellow spores). (c) Fluorescence microscopy of S. however, in rare cases, asci contain smaller spores with
macrospora asci and ascospores. YFP-tagged histones only a single nucleus. Arrows indicate small spores with
label nuclei. In the upper ascus, clearly one to two only a single nucleus compared to the regular ascos-
nuclei are visible in each ascospore, while the lower pores with two nuclei
an “A” strain can only be fertilized by a nucleus within one cell. After several conjugated divi-
of an “a” strain and vice versa. The fusion of the sions, ascus development will start with the
male gamete with the trichogyne will lead to the formation of ascogenous hyphae, which gener-
induction of the dikaryotic phase. During this ate the so-called crozier cell, which undergoes
phase, two genetically different nuclei exist conjugated divisions resulting in three cells,
⁄

Fig. 3 (continued) A. nidulans with small surrounding iella quadriseptata [from Tanaka et al. (2015)]. (a), (b),
Hülle cells. (c) Cleistothecia from the homothallic fun- and (c) are scanning electron micrographs and (d)–(f)
gus Eupenicillium crustaceum. (d) Perithecia from light microscopy; (b) courtesy of G. Braus (Göttingen,
Podospora anserina show typical hairs (arrow) at the Germany); (e) from Traeger et al. (2013); (f) copyright
neck of the perithecia. (e) Apothecia from P. confluens. from Elsevier Press
(f) Pseudothecium from the Dothideomycete Keissler-
two basal and one upper cell. The two nuclei in ate macro- or microconidia, and thus, only the
the upper dikaryotic cell undergo karyogamy sexual cycle contributes to the propagation of
followed by meiosis (Peraza-Reyes and Malag- this fungus. The sexual cycle can be completed
nac 2016). In N. crassa, a postmeiotic mitosis in the laboratory within 1 week, since ascos-
follows before spore formation starts. Thus, pores require no heat shock or resting period
each ascus contains eight linearly ordered for germination (Esser and Straub 1958). The
ascospores. After maturation, perithecia have sexual cycle starts with the formation of asco-
a size of about 300 mm, while ascospores have gonia. However, so far the molecular mechan-
a size between 15 and 30 mm. Importantly, isms leading to the formation of the dikaryotic
ascospore germination occurs only after a heat hyphae are not understood. After karyogamy
shock. Fruiting-body formation in N. crassa of two nuclei in the abovementioned crozier
was investigated in diverse genetic, biochemi- cells, meiosis will follow to generate the mei-
cal, and molecular studies (Davis 1995). otic products as a source for ascospore forma-
tion. Similar to ascus formation in N. crassa,
meiosis is followed by a postmeiotic mitosis.
B. Podospora anserina As a result, eight ascospores within a single
ascus are derived from a single dikaryotic
P. anserina is a coprophilic fungus with a pseu- mother cell.
dohomothallic mating system, which shows As mentioned above, sexual reproduction
similarities to the life cycle of N. crassa. The is a source of genetic diversity. Usually strains
mating-type strains are designated “+” (MAT1-2) of opposite mating types from heterothallic
and “” (MAT1-1). However, there are some species (e.g., N. crassa or P. anserina) are used
distinct differences compared to N. crassa. As for conventional genetic recombination stud-
male gametes, microconidia, but no macroco- ies. However, it has been shown for many spe-
nidia, are generated that germinate under species that recombination can also occur between
cific physiological conditions. Secondly, the two strains of a homothallic species. In these
asci usually contain only four spores, which cases, the strains are distinguished by at least a
are generated as a result of specific nuclear single mutation. Homothallic species such as S.
distribution mechanisms. After meiosis and macrospora and other Sordaria species are used
postmeiotic mitosis, spore-wall formation cov- for conventional genetic analysis (Teichert et al.
ers two genetically distinct nuclei. Usually one 2014a).
nucleus carries the “+” and the other the “”
mating type. With a frequency of about 3%,
five- or six-spored asci are generated. They D. Aspergillus nidulans
carry either two or four smaller spores that
carry only a single nucleus. These spores can A. nidulans, which is like S. macrospora a
be used to generate haploid mycelial isolates homothallic species, was used extensively to
(Scheckhuber and Osiewacz 2008; Peraza- study genetic recombination and fruiting-
Reyes and Malagnac 2016). body formation. The sexual cycle starts with
the formation of ascogonia and later dikaryotic
hyphae, a process, which is probably very simi-
C. Sordaria macrospora lar to the life cycle of S. macrospora. Within
cleistothecia, spherical asci are generated con-
S. macrospora is a coprophilic fungus that is taining eight ascospores. These octades are
taxonomically closely related to the above- unordered and thus distinguished from the
described species N. crassa and P. anserina. ordered asci of the abovementioned species.
The life cycles of all these ascomycetes are In recent years, several factors controlling cleis-
very similar, although S. macrospora has a tothecia formation were studied extensively,
homothallic mating system. In contrast to N. such as the velvet complex (Bayram and Braus
crassa however, S. macrospora does not gener- 2012) (see Sect. IV.C).
E. Emerging Model Systems 3. Trichoderma reesei
Here we mention fungal genera or species, Trichoderma reesei, the anamorph of Hypocrea
which were used recently to investigate jecorina, is a major industrial enzyme pro-
fruiting-body development. ducer, particularly of cellulases and hemicellu-
lases, which are used for applications in food,
feed, and biorefinery businesses. Until recently,
1. Aspergillus fumigatus, A. flavus, and it was believed that industrial strains propagate
A. parasiticus exclusively asexually. The industrial strains are
derived from a single isolate that carries the
The genus Aspergillus comprises about 340 spe- MAT1-2 mating-type locus. Crossing experi-
cies and was traditionally believed to contain ments of T. reesei with H. jecorina MAT1-1
species that generally propagate only asexually. wild-type isolates led to fertilized fruiting bod-
Species of this genus having a sexual life cycle ies and the production of mature ascospores.
were grouped in the teleomorphic genera Euro- However, the industrial T. reesei strain, which
tium, Emericella, Neosartorya, or Petromyces. was used for mating experiments, can be used
Recently evidence emerged that heterothallic only as male partner in crossings but is unable
Aspergillus species, which were for long to produce fruiting bodies and thus has to be
believed to propagate exclusively asexually, considered to be female sterile (Seidl et al. 2009;
have also the potential to undergo sexual repro- Linke et al. 2015; Schmoll and Wang 2016).
duction. These are, for example, the human
pathogen A. fumigatus and the mycotoxin-
producing fungi A. flavus and A. parasiticus 4. Penicillium species
(Dyer and O’Gorman 2011, 2012; Kück and Like Aspergillus species, Penicillium species
Pöggeler 2009; Dyer and Kück 2017). belong to the order of Eurotiales. Most species
were for long believed to represent the asexual
teleomorphs of the genera Eupenicillium or
2. Botrytis cinerea Talaromyces. However, the recent discovery of
The gray mold Botrytis cinerea Pers. Fr. (tele- a sexual cycle with cleistothecia in the indus-
omorph Botryotinia fuckeliana) is a notorious trial penicillin producer P. chrysogenum was
plant pathogenic fungus with a wide host range further evidence that supposedly asexual fungi
and has become an important model in molec- can undergo a heterothallic life cycle. Genetic
ular plant pathology. This fungus has two dis- analysis has provided evidence that even indus-
similar mating types (MAT1-1 and MAT1-2) trial strains have conserved their potential to
and is therefore heterothallic (Amselem et al. undergo a recombinant genetic cycle (Böhm
2011). Asexual propagation occurs through et al. 2013, 2015). Recently, also the cheese
microconidia in the light or sclerotia in the fungus Penicillium roqueforti was shown to
dark. The latter represent survival structures, have a sexual cycle with ascogonia, cleistothe-
which also serve the fertilization process during cia, and ascospores (Ropars et al. 2014). These
the sexual life cycle. When microconidia of the investigations suggest that the life cycle of sex-
opposite mating type are available, the sclerotia ually propagating Penicillium species is very
are fertilized and generate apothecia. This pro- similar to the one of heterothallic Aspergillus
cess is induced by light, and fruiting bodies species.
grow in the early stages toward the light source
(Schumacher 2017). However, in the labora- 5. Fusarium graminearum and F. verticillioides
tory, induction of the sexual life cycle is a rather
time-consuming process that takes 4 to 6 Fusarium species belong to the Sordariomy-
months until fully developed apothecia are cetes, which are characterized by perithecia for-
obtained (Faretra and Antonacci 1987). mation. F. graminearum (syn. Gibberella zeae)
and F. verticillioides (syn. Gibberella monilifor- ing of how these factors integrate into develop-
mis) are plant pathogenic fungi, which are mental signaling.
responsible for high losses in the harvest of
cereals. F. graminearum is homothallic, and
its sexual spores are responsible for disease 1. Light
initiation. In contrast, F. verticillioides is het- Fungi respond to light in various ways, includ-
erothallic and produces only modest numbers ing phototropic growth of reproductive struc-
of fruiting bodies. Both fungi have been used tures, modification of (circadian) rhythms, and
for comparative expression studies, using six changes in gene expression as well as primary
developmental stages of perithecia (Sikhakolli and secondary metabolism, among others
et al. 2012; Geng et al. 2014). Recently, F. gra- (Casas-Flores and Herrera-Estrella 2016).
minearum served as source in several genome- Here, we will focus on the influence of light on
wide large-scale functional analyses to decipher sexual reproduction of ascomycetes.
developmental genes and events related to Light-dependent fruiting-body formation,
sexual development. These attempts identified phototropism of perithecial necks, and light-
targets of the mating-type locus-encoded TFs, dependent ascospore discharge have been
RNA interference (RNAi) mechanisms, and described early on for a number of ascomycetes
perithecium-specific RNA-editing events (Kim (reviewed in Moore-Landecker 1992). For
et al. 2015; Liu et al. 2016; Son et al. 2017) example, light is required for apothecia forma-
tion in Ascobolus magnificus, Pyronema con-
fluens, and P. domesticum and for perithecia
6. Pyronema confluens formation in Pleurage setosa (syn. Podospora
P. confluens is a homothallic soil-living sapro- setosa) (Yu 1954; Carlile and Friend 1956; Call-
phytic Pezizomycete, which forms apothecia in aghan 1962). In T. reesei, formation of stro-
a light-dependent manner. The eight-spored mata, harboring the perithecia, occurs only in
asci are generated under laboratory conditions the presence of light (Seidl et al. 2009), while
within 6 days. In the early twentieth century, light inhibits formation of B. cinerea sclerotia,
this fungus was one of the first examples to which serve as survival structures and female
elucidate the dikaryotic phase during sexual mating partners (Schumacher 2017). Light-
development in filamentous ascomycetes. dependent positioning of the perithecial neck
Extensive transcriptome analysis was recently has been described, e.g., for Neurospora crassa
performed to investigate the light-dependent and P. setosa (Callaghan 1962; Harding and
fruiting-body formation (Traeger et al. 2013). Melles 1983). In Aspergillus glaucus and A.
nidulans, light favors asexual reproduction,
while darkness favors sexual reproduction
(Chona 1932; Mooney and Yager 1990; Blumen-
III. Factors Influencing Fruiting-Body stein et al. 2005).
Development Ascomycetes possess several photorecep-
tors, sensing near-UV/blue, green, and red
A. Environmental Factors light (Idnurm and Heitman 2005; Casas-Flores
and Herrera-Estrella 2016). All of these photo-
Ascomycetes generate fruiting bodies in certain receptors sense light by physical interaction
environmental conditions that are species-spe- with a chromophore: flavin for near-UV/blue
cific. Among others, light, nutrients, tempera- and blue-light receptors, retinal for green-light
ture, physical properties of growth substrates, receptors, and linear tetrapyrroles for red-light
and atmospheric conditions are relevant for receptors. Upon photon absorption, structural
fruiting-body formation. In this review, we will changes in the chromophore induce conforma-
focus on the influence of light and nutrients, tional changes in the photoreceptors, leading
highlighting recent advances in our understand- to changes in protein-protein interactions or
signaling via certain output domains (see photosensory domain combines PAS, GAF (for
below). The repertoire of photoreceptors sens- vertebrate cGMP-specific phosphodiesterases,
ing diverse light qualities differs between spe- cyanobacterial adenylate cyclases, transcription
cies, and, e.g., A. nidulans and B. cinerea activator FhlA), and PHY (phytochrome-
encode 3 and 11 photoreceptors, respectively specific PAS-related) domains. Absorption of
(Schumacher 2017). red light leads to reversible conformational
Blue-light receptors include crypto- changes of the chromophore, shifting the
chromes and LOV (light oxygen voltage) absorption maximum to far-red light. Thus,
domain proteins. Cryptochromes sense near- phytochromes are able to sense red/far-red
UV/blue light by binding flavin dinucleotide ratios (Rockwell and Lagarias 2010).
(FAD) and pterins/folates. Most cryptochromes Blue- and red-light photoreceptors have
show strongly reduced or no photolyase activ- roles in sexual development in various fungi
ity, one exception being A. nidulans (Bayram (for a review, see Casas-Flores and Herrera-
et al. 2008a). The LOV domain is a specialized Estrella 2016; Dasgupta et al. 2016; Fischer
PAS (found in Per, Arnt, Sim) domain, which et al. 2016). Besides photoreceptors, chromatin
binds the flavin chromophore. LOV domain modifiers (see Sect. IV.C.3), MAPK pathways
proteins may contain additional output (see Sect. IV.A.1), and transcription factors
domains, e.g., zinc finger or RGS (regulator of (see Sect. IV.C.1) are required for light regula-
G-protein signaling) domains. The white collar tion of sexual development. Light perception
1 homologs are light-activated transcription and light signal transduction have been ana-
factors with three PAS domains, one of which lyzed in great detail in few model systems,
is a FAD-binding LOV domain, and a GATA- including A. nidulans, N. crassa, and T. reesei
type zinc finger DNA-binding domain. WC-1 of (Seidl et al. 2009; Bayram et al. 2010; Schmoll
N. crassa interacts with a second white collar et al. 2010a; Dasgupta et al. 2016). An overview
protein, WC-2, via its PAS domains, and both of photoreceptors of these three model fungi
control expression of light-regulated genes by and their role in sexual development is shown
binding to promoter sequences as a heterodi- in Fig. 5.
mer (Cheng et al. 2002; Froehlich et al. 2002; He As mentioned above, A. nidulans repro-
et al. 2002; Smith et al. 2010). An example for a duces mainly sexually in the dark and mainly
LOV domain protein without further output asexually in light, and blue together with red
domains is N. crassa Vivid (VVD-1) involved light is the responsible light quality. Molecular
in photoadaptation (Chen et al. 2010). genetic analysis has shown that cryptochrome
Green-light receptors have not been exten- CryA, the white collar proteins LreA (light
sively characterized in filamentous ascomycetes response A) and LreB, as well as the phyto-
so far. These receptors are related to rhodop- chrome FphA are required for this light-
sins, composed of a seven-transmembrane dependent behavior (Blumenstein et al. 2005;
domain opsin bound to retinal via a conserved Bayram et al. 2008a; Purschwitz et al. 2008). The
lysine. Retinal binding in opsins leads to the cryptochrome CryA still has photolyase activity
formation of green-light responsive ion pumps and functions in DNA repair. Notably, A. nidu-
as demonstrated for the CarO protein of F. lans lacks a vivid and a functional, i.e., retinal-
fujikuroi (Garcia-Martinez et al. 2015). How- binding opsin homolog and has only one phy-
ever, other opsins like N. crassa NOP-1 lack tochrome, FphA, which binds biliverdin and
proton pump activity, indicating a putative reg- absorbs red as well as far-red light (Bayram
ulatory role (Bieszke et al. 1999; Brown et al. et al. 2008a; Idnurm et al. 2010; Fischer et al.
2001). 2016). CryA and FphA repress sexual develop-
Fungal phytochromes are red-light sensors ment in light conditions, while LreA and LreB
and contain multiple domains, with an N- stimulate sexual development, and lreA as well
terminal photosensory domain binding to a as lreB deletion mutants generate no perithecia
bilin-type linear tetrapyrrole, a histidine kinase in the light (Blumenstein et al. 2005; Bayram
domain, and a response regulator domain. The et al. 2008a; Purschwitz et al. 2008).
Fig. 5 Regulatory role of photoreceptors in sexual encircled P symbolizes phosphorylation. Bold orange
development of model ascomycetes. Data are summar- lines indicate light-induced interruption of signal
ized from recent reviews and research articles (Casas- transduction. For details, see main text. A, ATPase
Flores and Herrera-Estrella 2016; Dasgupta et al. 2016; domain; HK, histidine kinase; LOV, light oxygen volt-
Fischer et al. 2016; Schmoll and Wang 2016; Bazafkan age domain; PAS, found in Per, Arnt, Sim; PGP, photo-
et al. 2017). The upper part of the figure summarizes sensory domain of phytochromes combining PAS, GAF
photoreceptor types found in fungi and their respective (for vertebrate cGMP-specific phosphodiesterases, cya-
chromophors. The lower part of the figure summarizes nobacterial adenylate cyclases, transcription activator
the occurrence of photoreceptors and current knowl- FhlA), and PHY (phytochrome-specific PAS-related)
edge on signal transduction in light conditions in the domains; PL, photolyase; RR, response regulator
three model fungi A. nidulans, N. crassa, and T. reesei. domain; ZF, GATA-type zinc finger DNA-binding
Dashed lines indicate transcriptional regulation, while domain
continuous lines indicate physical interaction. An
Another random document with
no related content on Scribd:
All women did not run at the approach of the foe. A marked trait of
the settlers’ wives was their courage; and, indeed, opportunities
were plentiful for them to show their daring, their fortitude, and their
ready ingenuity. Hannah Bradley, of Haverhill, Mass., killed one
Indian by throwing boiling soap upon him. This same domestic
weapon was also used by some Swedish women near Philadelphia
to telling, indeed to killing advantage. A young girl in the Minot
House in Dorchester, Mass., shovelled live coals on an Indian
invader, and drove him off. A girl, almost a child, in Maine, shut a
door, barred, and held it while thirteen women and children escaped
to a neighboring block-house before the door and its brave defender
were chopped down. Anthony Bracket and his wife, captured by
savages, escaped through the wife’s skill with the needle. She
literally sewed together a broken birch-bark canoe which they found,
and in which they got safely away. Most famous and fierce of all
women fighters was Hannah Dustin, who, in 1697, with another
woman and a boy, killed ten Indians at midnight, and started for
home; but, calling to mind a thought that no one at home, without
corroborative evidence, would believe this extraordinary tale, they
returned, scalped their victims, and brought home the bloody
trophies safely to Haverhill.
Some Englishwomen were forced to marry their captors, forced by
torture or dire distress. Some, when captured in childhood, learned
to love their savage husbands. Eunice Williams, daughter of the
Deerfield minister, a Puritan who hated the Indians and the church of
Rome worse than he hated Satan, came home to her Puritan
kinsfolk wearing two abhorred symbols, a blanket and crucifix, and
after a short visit, not liking a civilized life, returned to her Indian
brave, her wigwam, and her priest.
I have always been glad that it was my far-away grandfather, John
Hoar, who left his Concord home, and risked his life as ambassador
to the Indians to rescue one of these poor “captivated” English
wives, Mrs. Mary Rowlandson, after her many and heart-rending
“savage removes.” I am proud of his “very forward spirit” which made
him dare attempt this bold rescue, as I am proud of his humanity and
his intelligent desire to treat the red men as human beings,
furnishing about sixty of them with a home and decent civilizing
employment. I picture him “stoutly not afraid,” as he entered the
camp, and met the poor captive, and treated successfully with her
savage and avaricious master, and then I see him tenderly leading
her, ragged, half-starved, and exhausted, through the lonely forests
home—home to the “doleful solemn sight” of despoiled Lancaster.
And I am proud, too, of the noble “Boston gentlewomen” who raised
twenty pounds as a ransom for Mary Rowlandson, “the price of her
redemption,” and tenderly welcomed her to their homes and hearts,
so warmly that she could write of them as “pitiful, tender-hearted,
and compassionate Christians,” whose love was so bountiful that
she could not declare it. If any one to-day marvels that English wives
did not “much desire the new and doleful land,” let them read this
graphic and thrilling story of the Captivity, Removes, and
Restauration of Mary Rowlandson, and he will marvel that the ships
were not crowded with disheartened settlers returning to their “faire
English homes.”
A very exciting and singular experience befell four dignified
Virginian wives in Bacon’s Rebellion, not through the Indians but at
the hands of their erstwhile friends. It is evident that the women of
that colony were universally and deeply stirred by the romance of
this insurrection and war. We hear of their dramatic protests against
the tyranny of the government. Sarah Drummond vowed she feared
the power of England no more than a broken straw, and
contemptuously broke a stick of wood to illustrate her words. Major
Chriesman’s wife, “the honor of her sex,” when her husband was
about to be put to death as a rebel, begged Governor Berkeley to kill
her instead, as he had joined Bacon wholly at her solicitation. One
Ann Cotton was moved by the war to drop into literary composition,
an extraordinary ebullition for a woman in her day, and to write an
account of the Rebellion, as she deemed “too wordishly,” but which
does not read now very wordishly to us. But for these four dames,
the wives of men prominent in the army under Governor Berkeley—
prime men, Ann Cotton calls them—was decreed a more stirring
participation in the excitements of war. The brilliant and erratic young
rebel, Bacon, pressed them into active service. He sent out
companies of horsemen and tore the gentlewomen from their
homes, though they remonstrated with much simplicity that they
were “indisposed” to leave; and he brought them to the scene of
battle, and heartlessly placed them—with still further and more acute
indisposition—on the “fore-front” of the breastworks as a shield
against the attacks of the four distracted husbands with their
soldiers. We read that “the poor Gentlewomen were mightily
astonished at this project; neather were their husbands void of
amazements at this subtill invention.” The four dames were
“exhibited to the view of their husbands and ffriends in the towne
upon the top of the smalle worke he had cast up in the night where
he caused them to tarey till he had finished his defence against the
enemy’s shott.” There stood these four innocent and harmless wives,
—“guardian angells—the white gardes of the Divell,” shivering
through the chill September night till the glimmering dawn saw
completed the rampart of earth and logs, or the leaguer, as it was
called by the writers with that exactness and absolute fitness of
expression which, in these old chronicles, gives such delight to the
lover of good old English. One dame was also sent to her husband’s
camp as a “white-aproned hostage” to parley with the Governor. And
this hiding of soldiers behind women was done by the order of one
who was called the most accomplished gentleman in Virginia, but
whom we might dub otherwise if we wished, to quote the
contemporary account, to “oppose him further with pertinances and
violent perstringes.”
I wish I could truthfully say that one most odious and degrading
eighteenth century English custom was wholly unknown in America
—the custom of wife-trading, the selling by a husband of his wife to
another man. I found, for a long time, no traces or hints of the
existence of such a custom in the colonies, save in two doubtful
cases. I did not wholly like the aspect of Governor Winthrop’s note of
the suggestion of some members of the church in Providence, that if
Goodman Verin would not give his wife full liberty to go to meeting
on Sunday and weekly lectures as often as she wished, “the church
should dispose her to some other man who would use her better.” I
regarded this suggestion of the Providence Christians with shocked
suspicion, but calmed myself with the decision that it merely
indicated the disposition of Goodwife Verin as a servant. And again,
in the records of the “Pticuler Court” of Hartford, Conn., in 1645, I
discovered this entry: “Baggett Egleston for bequething his wyfe to a
young man is fyned 20 shillings.” Now, any reader can draw his
conclusions as to exactly what this “bequething” was, and I cannot
see that any of us can know positively. So, though I was aware that
Baggett was not a very reputable fellow, I chose to try to persuade
myself that this exceedingly low-priced bequeathing did not really
mean wife-selling. But just as I was “setting down satysfyed” at the
superiority in social ethics and morality of our New England
ancestors, I chanced, while searching in the Boston Evening Post of
March 15, 1736, for the advertisement of a sermon on the virtues of
our forbears, entitled New England Tears and Fears of Englands
Dolours and Horrours, to find instead, by a malicious and contrary
fate, this bit of unwelcome and mortifying news not about old
England but about New England’s “dolours and horrours.”
Boston. The beginning of last Week a pretty odd and uncommon
Adventure happened in this Town, between 2 Men about a certain
woman, each one claiming her as his Wife, but so it was, that one of
them had actually disposed of his Right in her to the other for Fifteen
Shillings this Currency, who had only paid ten of it in part, and
refus’d to pay the other Five, inclining rather to quit the Woman and
lose his Earnest; but two Gentlemen happening to be present, who
were Friends to Peace, charitably gave him half a Crown a piece, to
enable him to fulfil his Agreement, which the Creditor readily took,
and gave the Woman a modest Salute, wishing her well, and his
Brother Sterling much Joy of his Bargain.
The meagre sale-money, fifteen shillings, was the usual sum
which changed hands in England at similar transactions, though one
dame of high degree was sold for a hundred guineas. In 1858 the
Stamford Mercury gave an account of a contemporary wife-sale in
England, which was announced through the town by a bellman. The
wife was led to the sale with a halter round her neck, and was “to be
taken with all her faults.” I am glad to say that this base British
husband was sharply punished for his misdemeanor.
It seems scarcely credible that the custom still exists in England,
but in 1882 a husband sold his wife in Alfreton, Derbyshire; and as
late as the 13th July, 1887, Abraham Boothroyd, may his name be
Anathema maranatha, sold his wife Clara at Sheffield, England, for
five shillings.
A most marked feature of social life in colonial times was the
belleship of widows. They were literally the queens of society. Fair
maids had so little chance against them, swains were so plentiful for
widows, that I often wonder whence came the willing men who
married the girls the first time, thus offering themselves as the
sacrifice at the matrimonial altar through which the girls could attain
the exalted state of widowhood. Men sighed sometimes in their
callow days for the girl friends of their own age, but as soon as their
regards were cast upon a widow, the girls at once disappear from
history, and the triumphant widow wins the prize.
Another marked aspect of this condition of society was the vast
number of widows in early days. In the South this was accounted for
by one of their own historians as being through the universally
intemperate habits of the husbands, and consequently their frequent
early death. In all the colonies life was hard, exposure was great to
carry on any active business, and the excessive drinking of
intoxicating liquors was not peculiar to the Southern husbands any
more than were widows. In 1698 Boston was said to be “full of
widows and orphans, and many of them very helpless creatures.” It
was counted that one sixth of the communicants of Cotton Mather’s
church were widows. It is easy for us to believe this when we read of
the array of relicts among which that aged but actively amorous
gentleman, Judge Sewall, found so much difficulty in choosing a
marriage partner, whose personal and financial charms he recounted
with so much pleasurable minuteness in his diary.
A glowing tribute to one of these Boston widows was paid by that
gossiping traveller, John Dunton, with so much evidence of deep
interest, and even sentiment, that I fancy Madam Dunton could not
have been wholly pleased with the writing and the printing thereof.
He called this Widow Breck the “flower of Boston,” the “Chosen
exemplar of what a Widow is.” He extols her high character, beauty,
and resignation, and then bridles with satisfaction while he says,
“Some have been pleas’d to say That were I in a single state they do
believe she wou’d not be displeas’d with my addresses.” He rode on
horseback on a long journey with his fair widow on a pillion behind
him, and if his conversation on “Platonicks and the blisses of
Matrimony” was half as tedious as his recounting of it, the road must
indeed have seemed long. He says her love for her dead husband is
as strong as death, but Widow Breck proved the strength of her
constancy by speedily marrying a second husband, Michael Perry.
As an instance of the complicated family relations which might
arise in marrying widows, let me cite the familiar case of the rich
merchant, Peter Sergeant, the builder of the famous Province House
in Boston. I will use Mr. Shurtleff’s explanation of this bewildering
gallimaufrey of widows and widowers:—
He was as remarkable in his marriages as his wealth; for he
had three wives, the second having been a widow twice
before her third venture; and his third also a widow, and even
becoming his widow, and lastly the widow of her third
husband.
To this I may add that this last husband, Simon Stoddart, also had
three wives, that his father had four, of whom the last three were
widows,—but all this goes beyond the modern brain to comprehend,
and reminds us most unpleasantly of the wife of Bath.
These frequent and speedy marriages were not wholly owing to
the exigencies of colonial life, but were the custom of the times in
Europe as well. I read in the diary of the Puritan John Rous, in
January, 1638, of this somewhat hasty wooing:—
A gentleman carried his wife to London last week and died
about eight o’clock at night, leaving her five hundred pounds a
year in land. The next day before twelve she was married to
the journeyman woolen-draper that came to sell mourning to
her.
I do not believe John Rous made special note of this marriage
simply because it was so speedy, but because it was unsuitable; as
a landed widow was, in social standing, far above a journeyman
draper.
As we approach Revolutionary days, the reign of widows is still
absolute.
Washington loved at fifteen a fair unknown, supposed to be Lucy
Grimes, afterward mother of Gen. Henry Lee. To her he wrote
sentimental poems, from which we gather (as might be expected at
that age) that he was too bashful to reveal his love. A year later he
writes:—
I might, was my heart disengaged, pass my time very
pleasantly as there’s a very agreeable Young Lady Lives in
the same house; but as thats only adding fuel to the fire it
makes me more uneasy; for by often and unavoidably being
in Company with her revives my former Passion for your
Lowland Beauty; whereas was I to live more retired from
young women, I might in some measure eliviate my sorrows
by burying that chast and troublesome passion in the grave of
oblivion or eternal forgetfulness.
The amorous boy of sixteen managed to “bury this chast and
troublesome passion,” to find the “Young Lady in the house” worth
looking at, and when he was twenty years old, to write to William
Fantleroy thus of his daughter, Miss Bettie Fantleroy:—
I purpose as soon as I recover my strength (from the
pleurisy) to wait on Miss Bettie in hopes of a reconsideration
of the former cruel sentence, and to see if I cannot obtain a
decision in my favor. I enclose a letter to her.
Later he fell in love with Mary Phillipse, who, though beautiful,
spirited, and rich, did not win him. This love affair is somewhat
shadowy in outline. Washington Irving thinks that the spirit of the
alert soldier overcame the passion of the lover, and that Washington
left the lists of love for those of battle, leaving the field to his
successful rival, Colonel Morris. The inevitable widow in the shape of
Madam Custis, with two pretty children and a fortune of fifteen
thousand pounds sterling, became at last what he called his
“agreeable partner for life,” and Irving thinks she was wooed with
much despatch on account of the reverses in the Phillipse episode.
Thomas Jefferson was another example of a President who
outlived his love-affair with a young girl, and married in serenity a
more experienced dame. In his early correspondence he reveals his
really tumultuous passion for one Miss Becca Burwell. He sighs like
a furnace, and bemoans his stammering words of love, but fair
Widow Martha Skelton made him eloquent. Many lovers sighed at
her feet; two of them lingered in her drawing-room one evening to
hear her sing a thrilling love-song to the accompaniment of
Jefferson’s violin. The love-song and music were so expressive that
the two disconsolate swains plainly read the story of their fate, and
left the house in defeat.
James Madison, supposed to be an irreclaimable old bachelor,
succumbed at first sight to the charms of fair Widow Dorothy Todd,
twenty years his junior, wooed her with warmth, and made her, as
Dolly Madison, another Mrs. President. Benjamin Franklin also
married a widow.
The characteristic glamour which hung round every widow
encircled Widow Sarah Syms, and Colonel Byrd gives a spirited
sketch of her in 1732:—
In the evening Tinsley conducted me to Widow Syms’
house where I intended to take up my quarters. This lady at
first suspecting I was some lover put on a gravity that
becomes a weed, but as soon as she learned who I was
brightened up with an unusual cheerfulness and serenity. She
was a portly handsome dame, of the family of Esau, and
seemed not to pine too much for the death of her husband.
This widow is a person of lively and cheerful conversation
with much less reserve than most of her country women. It
becomes her very well and sets off her other agreeable
qualities to advantage. We tossed off a bottle of honest port
which we relished with a broiled chicken. At nine I retired to
my devotions, and then slept so sound that fancy itself was
stupefied, else I should have dreamed of my most obliging
land-lady.
This “weed” who did not pine too much for her husband, soon
married again, and became the mother of Patrick Henry; and the
testimony of Colonel Byrd as to her lively and cheerful conversation
shows the heredity of Patrick Henry’s “gift of tongues.”
Hie! Betty Martin! tiptoe fine,

Couldn’t get a husband for to suit her mind!
was a famous Maryland belle, to whom came a-courting two friends,

young lawyers, named Dallam and Winston. It was a day of much
masculine finery and the two impecunious but amicable friends
possessed but one ruffled shirt between them, which each wore on
courting-day. Such amiability deserved the reward it obtained, for,
strange to say, both suitors won Betty Martin. Dallam was the first
husband,—the sacrifice,—and left her a widow with three sons and a
daughter. Winston did likewise, even to the exact number of children.
Daughter Dallam’s son was Richard Caswell, governor of South
Carolina, and member of Congress. Daughter Winston’s son was
William Paca, governor of Maryland, and member of the Continental
Congress. Both grandsons on their way to and from Congress
always visited their spirited old grandmother, who lived to be some
say one hundred and twenty years old.
There must have been afforded a certain satisfaction to a dying
husband—of colonial times—through the confidence that, by
unwavering rule, his widow would soon be cared for and cherished
by another. There was no uncertainty as to her ultimate settlement in
life, and even should she be unfortunate enough to lose her second
partner, he still had every reason to believe that a third would
speedily present himself. The Reverend Jonathan Burr when almost
moribund, piously expressed himself to “that vertuous gentlewoman
his wife with confidence” that she would soon be well provided for;
and she was, for “she was very shortly after very honourably and
comfortably married unto a gentleman of good estate,” a magistrate,
Richard Dummer, and lived with him nearly forty years. Provisions
were always made by a man in his will in case his wife married
again; scarcely ever to remove the property from her, but simply to
re-adjust the division or conditions. And men often signed ante-
nuptial contracts promising not to “meddle” with their wives’ property.
One curious law should be noted in Pennsylvania, in 1690, that a
widow could not marry till a year after her husband’s death.
There seem to have been many advantages in marrying a widow
—she might prove a valuable inheritance. The second husband
appeared to take a real pride in demanding and receiving all that
was due to the defunct partner. As an example let me give this
extract from a court record. On May 31st, 1692, the governor and
council of Maryland were thus petitioned:—
James Brown of St Marys who married the widow and relict
of Thomas Pew deceased, by his petition humbly prays
allowance for Two Years Sallary due to his Predecessor as
Publick Post employed by the Courts, as also for the use of a
Horse, and the loss of a Servant wholly, by the said Pew
deputed in his sickness to Officiate; and ran clear away with
his Horse, some Clothes &c., and for several months after not
heard of.
Now we must not be over-critical, nor hasty in judgment of the
manners and motives of two centuries ago, but those days are held
up to us as days of vast submissiveness and modesty, of patient
long-suffering, of ignorance of extortion; yet I think we would search
far, in these degenerate days, for a man who, having married a relict,
would, two years after his “Predecessor’s” death, have the colossal
effrontery to demand of the government not only the back salary of
said “Predecessor,” but pay for the use of a horse stolen by the
Predecessor’s own servant—nay, more, for the value of the said
servant who elected to run away. Truly James Brown builded well
when he chose a wife whose departing partner had, like a receding
wave, deposited much lucrative silt on the matrimonial shore, to be
thriftily gathered in and utilized as a bridal dower by his not-too-
sensitive successor.
In fact it may plainly be seen that widows were life-saving stations
in colonial social economy; one colonist expressed his attitude
towards widows and their Providential function as economic aids,
thus:—
Our uncle is not at present able to pay you or any other he
owes money to. If he was able to pay he would; they must
have patience till God enable him. As his wife died in mercy
near twelve months since, it may be he may light of some rich
widow that may make him capable to pay; except God in this
way raise him he cannot pay you or any one else.
It certainly must have been some satisfaction to every woman to
feel within herself the possibility of becoming such a celestial agent
of material salvation.
I wish to state, in passing, that it is sometimes difficult to judge as
to the marital estate of some dames, to know whether they were
widows at the time of the second marriage or not, for the prefixed
Mrs. was used indifferently for married and single women, and even
for young girls. Cotton Mather wrote of “Mrs. Sarah Gerrish, a very
beautiful and ingenious damsel seven years of age.” Rev. Mr.
Tompson wrote a funeral tribute to a little girl of six, which is entitled
and begins thus:—
A Neighbors Tears dropt on ye grave of an Amiable Virgin,
a pleasant Plant cut down in the blooming of her Spring viz;
Mrs Rebecka Sewall Anno Aetatis 6, August ye 4ᵗʰ 1710.
I saw this Pritty Lamb but t’other day

With a small flock of Doves just in my way
Ah pitty tis Such Prittiness should die
With rare alliances on every side.
Had Old Physitians liv’d she ne’er had died.
The pious old minister did not really mean by this tribute to the old-
school doctors, that Mrs. Rebecka would have achieved earthly
immortality. He modestly ends his poetic tribute thus:—
Had you given warning ere you pleased to Die
You might have had a Neater Elegy.
These consorts and relicts are now but shadows of the past:—
their bones are dust,

Their souls are with the saints, I trust.
The honest and kindly gentlemen who were their husbands,

sounded their virtues in diaries and letters; godly ministers preached
their piety in labored and dry-as-dust sermons. Their charms were
sung by colonial poets in elegies, anagrams, epicediums, acrostics,
threnodies, and other decorous verse. It was reserved for a man of
war, and not a very godly man of war either, to pæan their good
sense. Cervantes says that “womans counsel is not worth much, yet
he who despises it is no wiser than he should be.” With John
Underhill’s more gallant tribute to the counsel of a consort, we may
fitly end this chapter.
Myself received an arrow through my coat sleeve, a second
against my helmet on the forehead; so as if God in his
Providence had not moved the heart of my wife to persuade
me to carry it along with me (which I was unwilling to do) I had
been slain. Give me leave to observe two things from hence;
first when the hour of death is not yet come, you see God
useth weak means to keep his purpose unviolated; secondly
let no man despise advice and counsel of his wife though she
be a woman. It were strange to nature to think a man should
be bound to fulfil the humour of a woman, what arms he
should carry; but you see God will have it so, that a woman
should overcome a man. What with Delilahs flattery, and with
her mournful tears, they must and will have their desire, when
the hand of God goes along in the matter, and this to
accomplish his own will. Therefore let the clamor be
quenched that I hear daily in my ears, that New England men
usurp over their wives and keep them in servile subjection.
The country is wronged in this matter as in many things else.
Let this precedent satisfy the doubtful, for that comes from the
example of a rude soldier. If they be so courteous to their
wives as to take their advice in warlike matters, how much
more kind is the tender affectionate husband to honor his wife
as the weaker vessel. Yet mistake not. I say not they are
bound to call their wives in council, though they are bound to
take their private advice (so far as they see it make for their
advantage and good). Instance Abraham.
CHAPTER II.
WOMEN OF AFFAIRS.
The early history of Maryland seems singularly peaceful when

contrasted with that of other colonies. There were few Indian horrors,
few bitter quarrels, comparatively few petty offences. In spite of the
influx of convicts, there was a notable absence of the shocking
crimes and equally shocking punishments which appear on the court
records of other provinces; it is also true that there were few schools
and churches, and but scanty intellectual activity. Against that
comparatively peaceful background stands out one of the most
remarkable figures of early colonial life in America—Margaret Brent;
a woman who seemed more fitted for our day than her own. She was
the first woman in America to demand suffrage, a vote, and
representation.
She came to the province in 1638 with her sister Mary (another
shrewd and capable woman), her two brothers, and nine other
colonists. The sisters at once took up land, built manorhouses, and
shortly brought over more colonists; soon the court-baron and court-
leet were held at Mary Brent’s home, St. Gabriel’s Manor, on old
Kent Island. We at once hear of the sisters as active in business
affairs, registering cattle marks, buying and selling property,
attending with success to important matters for their brothers; and
Margaret soon signed herself “Attorney for my brother, &c., &c.,” and
was allowed the right so to act. The Brents were friends and
probably kinsfolk of Lord Baltimore, and intimate friends, also, of the
governor of Maryland, Leonard Calvert. When the latter died in 1647,
he appointed by nuncupation one Thomas Greene as his successor
as governor, and Margaret Brent as his sole executrix, with the
laconic instruction to “Take all and Pay all,” and to give one Mistress
Temperance Pypott a mare colt. His estate was small, and if he had
made Greene executor, and Mistress Margaret governor, he would
have done a much more sensible thing; for Greene was vacillating
and weak, and when an emergency arose, he had to come to
Margaret Brent for help. The soldiers, who had assisted the
government in recent troubles, were still unpaid, and Governor
Calvert had pledged his official word and the property of Lord
Baltimore that they should be paid in full. After his death an
insurrection in the army seemed rising, when Mistress Brent calmly
stepped in, sold cattle belonging to the Proprietary, and paid off the
small but angry army. This was not the only time she quelled an
incipient mutiny. Her kinsman, Lord Baltimore, was inclined to find
bitter fault, and wrote “tartly” when the news of her prompt action and
attendant expenditure reached his ears; but the Assembly sent him a
letter, gallantly upholding Mistress Brent in her “meddling,” saying
with inadvertent humour, that his estate fared better in her hands
than “any man elses.”
Her astonishing stand for woman’s rights was made on January
21, 1647-48, two centuries and a half ago, and was thus recorded:—
Came Mrs Margaret Brent and requested to have vote in
the House for herself and voyce allsoe, for that on the last
Court 3rd January it was ordered that the said Mrs Brent was
to be looked upon and received as his Ldp’s Attorney. The
Governor deny’d that the s’d Mrs Brent should have any vote
in the house. And the s’d Mrs Brent protested against all
proceedings in this present Assembly unlesse she may be
present and have vote as afores’d.
With this protest for representation, and demand for her full rights,
this remarkable woman does not disappear from our ken. We hear of
her in 1651 as an offender, having been accused of killing wild cattle
and selling the beef. She asserted with vigor and dignity that the
cattle were her own, and demanded a trial by jury.
And in 1658 she makes her last curtsey before the Assembly and
ourselves, a living proof of the fallacy of the statement that men do
not like strong-minded women. For at that date, at the fully ripened
age of fifty-seven, she appeared as heir of an estate bequeathed to
her by a Maryland gentleman as a token of his love and affection,
and of his constant wish to marry her. She thus vanishes out of
history, in a thoroughly feminine rôle, that of a mourning sweetheart;
yet standing signally out of colonial days as the most clear-cut,
unusual, and forceful figure of the seventeenth century in Maryland.
Another Maryland woman of force and fearlessness was Verlinda
Stone. A letter from her to Lord Baltimore is still in the Maryland
archives, demanding an investigation of a fight in Anne Arundel
County, in which her husband was wounded. The letter is
businesslike enough, but ends in a fiery postscript in which she uses
some pretty strong terms. Such women as these were not to be
trifled with; as Alsop wrote:—
All Complemental Courtships drest up in critical Rarities are
meer Strangers to them. Plain wit comes nearest to their
Genius, so that he that intends to Court a Maryland girle,
must have something more than the tautologies of a long-
winded speech to carry on his design.
Elizabeth Haddon was another remarkable woman; she founded
Haddonfield, New Jersey. Her father had become possessed of a
tract of land in the New World, and she volunteered to come alone to
the colony, and settle upon the land. She did so in 1701 when she
was but nineteen years old, and conducted herself and her business
with judgment, discretion, and success, and so continued throughout
her long life. She married a young Quaker named Esthaugh, who
may have been one of the attractions of the New World. Her
idealized story has been told by L. Maria Child in her book The
Youthful Emigrant.
John Clayton, writing as early as 1688 of “Observables” in Virginia,
tells of several “acute ingenious gentlewomen” who carried on
thriving tobacco-plantations, draining swamps and raising cattle and
buying slaves. One near Jamestown was a fig-raiser.
In all the Southern colonies we find these acute gentlewomen
taking up tracts of land, clearing them, and cultivating their holdings.
In the settlement of Pennsylvania, Mary Tewee took two thousand
five hundred acres in what is now Lancaster County. She was the
widow of a French Huguenot gentleman, the friend of William Penn,
and had been presented at the court of Queen Anne.
New England magistrates did not encourage such independence.
In the early days of Salem, “maid-lotts” were granted to single
women, but stern Endicott wrote that it was best to abandon the
custom, and “avoid all presedents & evil events of granting lotts vnto
single maidens not disposed of.” The town of Taunton, Mass., had an
“ancient maid” of forty-eight years for its founder, one Elizabeth
Poole; and Winthrop says she endured much hardship. Her
gravestone says she was a “native of old England of good family,
friends and prospects, all of which she left in the prime of her life to
enjoy the religion of her conscience in this distant wilderness. A
great proprietor of the township of Taunton, a chief promoter of its
settlement in 1639. Having employed the opportunity of her virgin
state in piety, liberality and sanctity of manners, she died aged 65.”
Lady Deborah Moody did not receive from the Massachusetts
magistrates an over-cordial or very long-lived welcome. She is
described as a “harassed and lonely widow voluntarily exiling herself
for conscience’ sake.” Perhaps her running in debt for her
Swampscott land and her cattle had quite as much to do with her
unpopularity as her “error of denying infant baptism.” But as she paid
nine hundred or some say eleven hundred pounds for that wild land,
it is no wonder she was “almost undone.” She was dealt with by the
elders, and admonished by the church, but she “persisted” and
finally removed to the Dutch, against the advice of all her friends.
Endicott called her a dangerous woman, but Winthrop termed her a
“wise and anciently religious woman.” Among the Dutch she found a
congenial home, and, unmolested, she planned on her Gravesend
farm a well-laid-out city, but did not live to carry out her project. A
descendant of one of her Dutch neighbors writes of her:—
Tradition says she was buried in the north-west corner of
the Gravesend church yard. Upon the headstone of those
who sleep beside her we read the inscription In der Heere
entslapen—they sleep in the Lord. We may say the same of
this brave true woman, she sleeps in the Lord. Her rest has
been undisturbed in this quiet spot which she hoped to make
a great city.
It seems to be plain that the charge of the affairs of Governor John
Winthrop, Jr., in New Haven was wholly in the hands of Mrs.
Davenport, the wife of the minister, Rev. John Davenport. Many
sentences in her husband’s letters show her cares for her friends’
welfare, the variety of her business duties, and her performance of
them. He wrote thus to the Governor in 1658:—
For your ground; my wife speedily, even the same day she
received your letter, spake with sundry about it, and received
this answer, that there is no Indian corne to be planted in that
quarter this yeare. Brother Boykin was willing to have taken it,
but saith it is overrun with wild sorrell and it will require time to
subdue it, and put it into tillage, being at present unfit to be
improved. Goodman Finch was in our harbour when your
letter came, & my wife went promptly downe, and met with
yong Mr Lamberton to whom she delivered your letter. He
offered some so bad beaver that my wife would not take it. My
wife spake twise to him herself. My wife desireth to add that
she received for you of Mr Goodenhouse 30s worth of beaver
& 4s in wampum. She purposeth to send your beaver to the
Baye when the best time is, to sell it for your advantage and
afterwards to give you an account what it comes to. Your
letter to Sarjiunt Baldwin my wife purposeth to carry to him by
the 1st opportunity. Sister Hobbadge has paid my wife in part
of her debt to you a bushel of winter wheate.
The letters also reveal much loving-kindness, much eagerness to
be of assistance, equal readiness to welcome new-comers, and to
smooth the rough difficulties in pioneer housekeeping. Rev. Mr.
Davenport wrote in August, 1655, from New Haven to Gov. Winthrop
at Pequot:—
Hon’ᵈ Sir,—We did earnestly expect your coming hither
with Mrs. Winthrop and your familie, the last light moone,
having intelligence that a vessel wayted upon you at Pequot
for that end, and were thereby encouraged to provide your
house, that it might be fitted in some measure, for your
comfortable dwelling in it, this winter.
My wife was not wanting in her endeavors to set all
wheeles in going, all hands that she could procure on worke,
that you might find all things to your satisfaction. Though she
could not accomplish her desires to the full, yet she
proceeded as farr as she could; whereby many things are
done viz. the house made warme, the well cleansed, the
pumpe fitted for your use, some provision of wood layed in,
and 20 loades will be ready, whensoever you come; and
sundry, by my wife’s instigation, prepared 30 bush. of wheate
for the present and sister Glover hath 12 lb of candles ready
for you. My wife hath also procured a maid servant for you,
who is reported to be cleanly and saving, her mother is of the
church, and she is kept from a place in Connectacot where
she was much desired, to serve you....
If Mrs. Winthrop knew how wellcome she will be to us she
would I believe neglect whatsoever others doe or may be
forward to suggest for her discouragement. Salute her, with
due respect, in my name and my wife’s, most affectionately.
Madam Davenport also furnished the rooms with tables and
“chayres,” and “took care of yor apples that they may be kept safe
from the frost that Mrs. Winthrop may have the benefit of them,” and
arranged to send horses to meet them; so it is not strange to learn in
a postscript that the hospitable kindly soul, who thus cheerfully
worked to “redd the house,” had a “paine in the soles of her feet,
especially in the evening;” and a little later on to know she was
“valetudinarious, faint, thirsty, of little appetite yet cheerful.”
All these examples, and many others help to correct one very
popular mistake. It seems to be universally believed that the
“business woman” is wholly a product of the nineteenth century.
Most emphatically may it be affirmed that such is not the case. I
have seen advertisements dating from 1720 to 1800, chiefly in New
England newspapers, of women teachers, embroiderers, jelly-
makers, cooks, wax-workers, japanners, mantua-makers,—all truly
feminine employments; and also of women dealers in crockery,
musical instruments, hardware, farm products, groceries, drugs,
wines, and spirits, while Hawthorne noted one colonial dame who
carried on a blacksmith-shop. Peter Faneuil’s account books show
that he had accounts in small English wares with many Boston
tradeswomen, some of whom bought many thousand pounds’ worth
of imported goods in a year. Alice Quick had fifteen hundred pounds
in three months; and I am glad to say that the women were very
prompt in payment, as well as active in business. By Stamp Act
times, the names of five women merchants appear on the Salem list
of traders who banded together to oppose taxation.
It is claimed by many that the “newspaper-woman” is a growth of
modern times. I give examples to prove the fallacy of this statement.
Newspapers of colonial times can scarcely be said to have been
edited, they were simply printed or published, and all that men did as
newspaper-publishers, women did also, and did well. It cannot be
asserted that these women often voluntarily or primarily started a
newspaper; they usually assumed the care after the death of an
editor husband, or brother, or son, or sometimes to assist while a
male relative, through sickness or multiplicity of affairs, could not
attend to his editorial or publishing work.
Perhaps the most remarkable examples of women-publishers may
be found in the Goddard family of Rhode Island. Mrs. Sarah
Goddard was the daughter of Ludowick Updike, of one of the oldest
and most respected families in that State. She received an excellent
education “in both useful and polite learning,” and married Dr. Giles
Goddard, a prominent physician and postmaster of New London.
After becoming a widow, she went into the printing business in
Providence about the year 1765, with her son, who was postmaster
of that town. They published the Providence Gazette and Country
Journal, the only newspaper printed in Providence before 1775.
William Goddard was dissatisfied with his pecuniary profit, and he
went to New York, leaving the business wholly with his mother; she
conducted it with much ability and success under the name Sarah
Goddard & Company. I wish to note that she carried on this business
not under her son’s name, but openly in her own behalf; and when

Textbook Physiology and Genetics Selected Basic and Applied Aspects 2Nd Edition Timm Anke Ebook All Chapter PDF

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Textbook Physiology and Genetics Selected Basic and Applied Aspects 2Nd Edition Timm Anke Ebook All Chapter PDF

Uploaded by

Copyright:

Available Formats

Physiology and Genetics Selected

Basic and Applied Aspects 2nd Edition

Systemic Lupus Erythematosus: Basic, Applied and

Neotropical Social Wasps: Basic and applied aspects

Basic Physiology for Anaesthetists 2nd Edition David

Advancing Frontiers in Mycology amp Mycotechnology

Basic and Applied Bone Biology 2nd Edition David B.

Basic and Applied Phytoplankton Biology Perumal

Marathon Running Physiology Psychology Nutrition and

The Heparins: Basic and Clinical Aspects 1st Edition

Physiology and Genetics

I Growth, Differentiation and Sexuality

Physiology and Genetics

Professor Dr. Timm Anke

Dr. Anja Schüffler

ISBN 978-3-319-71739-5 ISBN 978-3-319-71740-1 (eBook)

Library of Congress Control Number: 2017964284

Printed on acid-free paper

This Springer imprint is published by Springer Nature

chemistry and in the production of antibiotics and other bioactive compounds.

Bochum, Germany KARL ESSER

covers the current molecular targets of antifungal compounds and discusses

Kaiserslautern and Mannheim, Germany Timm Anke

Fruiting-Body Development in Ascomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Fungal Inteins: Distribution, Evolution, and Applications . . . . . . . . . . . . . 57

Yeast Killer Toxins: Fundamentals and Applications . . . . . . . . . . . . . . . . . . . 87

The Fungal MCC/Eisosome Complex: An Unfolding Story . . . . . . . . . . . 119

Volatiles in Communication of Agaricomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Endophytic Fungi, Occurrence, and Metabolites . . . . . . . . . . . . . . . . . . . . . . . . 213

Secondary Metabolites of Basidiomycetes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231

Identification of Fungicide Targets in Pathogenic Fungi . . . . . . . . . . . . . . . 277

Helminth Electron Transport Inhibitors Produced by Fungi . . . . . . . . . . . 297

Cyclic Peptides and Depsipeptides from Fungi . . . . . . . . . . . . . . . . . . . . . . . . . . . 331

Polyketide Synthase–Nonribosomal Peptide Synthetase Hybrid Enzymes

Biosynthesis of Fungal Polyketides . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385

Aspects of the Occurrence, Genetics, and Regulation of Biosynthesis of the

Biosystematic Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 435

Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 447

ANNA MARIA PIRTTILÄ

S. PÖGGELER1, M. NOWROUSIAN2, I. TEICHERT2, A. BEIER2, U. KÜCK2

CONTENTS 2. Transcriptional Changes During

Physiology and Genetics, 2nd Edition

the expression of developmentally regulated DOPA (L-3,4-dihydroxyphenylalanine) path-

E. Emerging Model Systems 3. Trichoderma reesei

Hie! Betty Martin! tiptoe fine,

was a famous Maryland belle, to whom came a-courting two friends,

I saw this Pritty Lamb but t’other day

their bones are dust,

The honest and kindly gentlemen who were their husbands,

The early history of Maryland seems singularly peaceful when

You might also like