Genetics and Molecular Biology of Entomopathogenic Fungi

498 Pages·2016·20.894 MB·English

by  Brian Lovett and Raymond J. St. Leger (Eds.)

VOLUMENINETYFOUR
A
DVANCES IN
GENETICS
Genetics and Molecular Biology
of Entomopathogenic Fungi
ADVANCES IN GENETICS, VOLUME 94
Serial Editors
Theodore Friedmann
Department of Pediatrics, University of California at San Diego,
School of Medicine, CA, USA
Jay C. Dunlap
DepartmentofGenetics,TheGeiselSchoolofMedicineatDartmouth,
Hanover, NH, USA
Stephen F. Goodwin
Department of Physiology, Anatomy and Genetics,
University of Oxford, Oxford, UK
VOLUME NINETY FOUR
A
DVANCES IN
GENETICS
Genetics and Molecular Biology
of Entomopathogenic Fungi
Edited by
BRIAN LOVETT
University of Maryland, College Park, MD,
United States
RAYMOND J. St. LEGER
University of Maryland, College Park, MD,
United States
AMSTERDAM(cid:129)BOSTON(cid:129)HEIDELBERG(cid:129)LONDON
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SANFRANCISCO(cid:129)SINGAPORE(cid:129)SYDNEY(cid:129)TOKYO
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DEDICATION
The editors dedicate this book to the doyen of insect pathology, Donald
Roberts, and to his versatile little friend Metarhizium robertsii.
j
v
CONTRIBUTORS
J.P.M.Araújo
Pennsylvania State University, University Park, PA,UnitedStates
L.Barelli
Brock University, St.Catharines, ON, Canada
M.J. Bidochka
Brock University, St.Catharines, ON, Canada
T.M.Butt
Swansea University, Swansea,Wales,UnitedKingdom
C.J. Coates
Swansea University, Swansea,Wales,UnitedKingdom
C.de Bekker
Ludwig-Maximilians-University Munich,Munich, Germany
H.H. De FineLicht
University of Copenhagen, Frederiksberg, Denmark
B.G.G. Donzelli
Cornell University, Ithaca, NY,UnitedStates
I.M.Dubovskiy
SBRussian Academy of Sciences, Novosibirsk, Russia
J. Eilenberg
University of Copenhagen, Frederiksberg, Denmark
H.C. Evans
CAB International, Surrey,UnitedKingdom
W.Fang
Zhejiang University,Hangzhou, Zhejiang, China
A.E. Hajek
Cornell University, Ithaca, NY,UnitedStates
D.P. Hughes
Pennsylvania State University, University Park, PA,UnitedStates
A.B. Jensen
University of Copenhagen, Frederiksberg, Denmark
N.O. Keyhani
University of Florida, Gainesville, FL,UnitedStates
S.B. Krasnoff
USDA-ARS, Ithaca,NY,United States
j
xi
xii Contributors
R.G.Loreto
Pennsylvania State University,University Park, PA, UnitedStates; CAPES Foundation,
Ministryof Education of Brazil, Brasília,DF,Brazil
B.Lovett
University of Maryland, College Park, MD,UnitedStates
H.-L.Lu
University of Maryland, College Park, MD,UnitedStates
S.Moonjely
Brock University,St. Catharines, ON, Canada
A.Ortiz-Urquiza
University of Florida, Gainesville,FL, UnitedStates
L.Quevillon
Pennsylvania State University,University Park, PA, UnitedStates
N.A. Ratcliffe
SwanseaUniversity,Swansea,Wales,UnitedKingdom;UniversidadeFederalFluminense,
Niteroi,Rio deJaneiro, Brazil
R.J. St.Leger
University of Maryland, College Park, MD,UnitedStates
C.Wang
Chinese Academy of Sciences, Shanghai,China
J.B.Wang
University of Maryland, College Park, MD,UnitedStates
H.Zhao
Zhejiang University,Hangzhou, Zhejiang, China
PREFACE
Insects are the most species-rich group of eukaryotes, and this diversity
grants these animals profound power to influence ecosystems. The great
research interest in insects arises from their ability to negatively impact
human society: as vectors for disease in humans and our livestock and as
destroyers of crops and stored products. It is estimated that insects destroy
approximately18%oftheworldannualcropproduction,andvectorborne
diseaseskillmillionseveryyear.Thus,thediseases,pathogens,andimmune
responses of insects have been a long-standing research interest. Early
interest grew mostly from economic concerns. For example, Beauveria
bassiana was described by Agostino Bassi in 1835 as the cause of the devas-
tating muscardine disease of silkworm, and it was instrumental in his
development of the germ theory of disease (Steinhaus, 1956). In 1880, the
pioneer immunologist Elie Metchnikoff was among the first to propose
practical methods of microbial biological control of an insect crop pest,
initiating trials of the fungus Metarhizium anisopliae against grain beetles
(Lord, 2005; Mechnikoff, 1879, Fig. 1).
Suchappliedinterestcontinuestoday,asweallstillhaveastakeineither
lengthening the life expectancy of useful insects or shortening the life
expectancy of pestiferous ones. Since fungi are the commonest disease-
causing agents in insects and populations of most insects are regulated by
density-dependent factors involving pathogens and predators, we need to
understand and be able to manipulate the interactions of pest insects with
fungiandtheirothernaturalenemiesinordertofeedtheworldandprevent
disease.Mostresearchonfungal-insectpathogenshascontinuedtofocuson
hypocrealean Ascomycetes from the genera Beauveria and Metarhizium
(familyCordycipitaceaeandClavicipitaceae,respectively).Thesegeneraare
tractablemodelspecies,arereadilycultivatable,andhaveaparticularlywide
host range allowing them to be applied en masse against vectors of human
disease and multifarious agricultural pests. Numerous registered mycoin-
secticideformulationsarebasedonBeauveriaandMetarhiziumspp.Theyalso
have a worldwide distribution with variants adapted from the arctic to the
tropicsandcolonizinganimpressivearrayofenvironmentsincludingforests,
savannahs, swamps, coastal zones, and deserts.
The majority of chapters in this volume focus on Metarhizium and
Beauveria, reflecting the preponderance of published data that has utilized
j
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xiv Preface
Figure 1 (1) Blood cells and fungal conidia from the blood of dying wheat chafer
larvae.Magnified550times(Eyepiece3+System9alonepicture).(2)Fungalconidia
from the blood of living and apparently healthy larvae. Magnified 860 times (4+9).
(3)Piecesofsubcutaneousmycelialconidiophoresformedonfreshcadavers.Magnified
Preface xv
thesegeneraasmodelsforentomopathogenicity.Yet,asclarifiedbyArau(cid:1)jo
and Hughes in Chapter 1, most major fungal taxonomic groups have
memberspathogenictoinsects,sofungal-insectpathogensrepresentlifestyle
adaptations that have likely evolved numerous times. Arau(cid:1)jo and Hughes
emphasize that, despite their ecological importance and potential applica-
tions, the entomopathogenic fungi are woefully understudied, especially
regardingtheirbiodiversity,whichlikelyharborsoneofthelargestreservoirs
of undocumented fungal species. Their chapter provides a broad overview
of host–pathogen relationships by focusing on the impressive diversity of
morphologies,ecologies,andinteractionsbetweeninsectsandfungi.Ashost
andpathogenaretiedtogetherinaveryintimaterelationship,thepresented
work strongly argues for empirical studies investigating both sides of the
relationship in a much broader range of fungal-insect interactions than has
hitherto been conducted. The authors reveal significant differences in host
range and pathogenic strategies between the major groups of fungal
pathogens.Thebasalgroups,suchasaquaticchytrids,infectmostlyDiptera,
while Microsporidia and Entomophthoromycota infect a wide range of
hosts.BasidiomycotainfectsmostlyHemiptera,whileAscomycota,themost
species-rich group, infects a vast number of insect groups. In terms of
pathogenicstrategies,theascomycetes,forexample,areallhemibiotrophic,
switching from a biotrophic phase (parasitism) to a necrotrophic phase,
growing on the dead host.
Incontrast,theEntomophthoromycotaarecharacteristicallybiotrophics
with no somatic growth after death.
=---------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------------
550times.(4)Conidiafromwithinthesamelarvaeonthedayafterdeath.Magnified
550times.(5)Overgrownconidiaonthethirddayemergingfromthelarvalcadaverina
humidchamber.Magnified400times(3+8).(6)Conidiaandhyphaewithtwoforming
conidia(a)inahumidifiedchamber.Magnified400times.(7)Candelabra-shapedstalks
with sterigmata forming conidia cultured in a humid chamber. Magnified 400 times.
(8) Mature Entomophthora anisopliae spores. Magnified 1340 times (Eyepiece 3+14
System picture). (9) Cross section of larva covered with fungus. a-conidial layer,
b-straighthyphallayer,c-feltedlayer,d-candelabralayerwithsterigmataandconidia.
Magnified 90 times (3+4). (10) Three conidia grown in sugar water. Magnified 550
times. (11) Germinating spores on larval cuticle. a, b- Spore germ tubes producing
yellowspots.c-Bursiformgermliningsonthecuticle.d-Furtherstageofgermination.
e- Spot on larval cuticle with ungerminated and germinating conidia. Magnified 550
times.(12)Spotsonthecuticlewithsproutslikeraysofradiatinghyphae;withnotable
formation of conidia. Magnified 400 times. This figure is from Elie Metchnikoff’s pub-
licationdiscoveringMetarhiziumanisopliaeinOdessain1879;thelegendisatranslation
ofhisoriginalRussianlegendforthefigure.