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THE MYCOTA 5 A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research Series Editors: Dee Carter · Anuradha Chowdhary Joseph Heitman · Ulrich Kück Barry Scott Carl Mesarich Editors Plant Relationships Fungal-Plant Interactions Third Edition The Mycota A Comprehensive Treatise on Fungi as Experimental Systems for Basic and Applied Research Volume 5 SeriesEditors DeeCarter,SchoolofLifeandEnvironmentalSciences,TheUniversityof Sydney,Sydney,NSW,Australia AnuradhaChowdhary,DepartmentofMedicalMycology,VPChestInstitute, UniversityofDelhi,Delhi,India JosephHeitman,DukeUniversityMedicalCenter,Durham,NC,USA UlrichKück,GeneralandMolecularBotany,Ruhr-UniversityBochum, Bochum,Germany Thefungirepresentaheterogenousassemblageofeukaryoticmicroorganisms andhavebecomefavoredorganismsforresearchatthecellularandmolecular level. Such research involvement has been stimulated by interest in the biotechnologicalapplicationoffungiinprocessesrelatedtoindustry,agricul- ture and ecology. Considering both yeasts and mycelial fungi, THE MYCOTA highlights developments in both basic and applied research and presents an overview of fungal systematics and cell structure. Foremost authoritiesinresearchonmycologyhavebeenassembledtoeditandcontrib- utetothevolumes. (cid:129) Barry Scott Carl Mesarich Editors Plant Relationships Fungal-Plant Interactions Third Edition Editors BarryScott CarlMesarich SchoolofNaturalSciences SchoolofAgricultureandEnvironment MasseyUniversity MasseyUniversity PalmerstonNorth,NewZealand PalmerstonNorth,NewZealand ISSN2945-8048 ISSN2945-8056 (electronic) TheMycota ISBN978-3-031-16502-3 ISBN978-3-031-16503-0 (eBook) https://doi.org/10.1007/978-3-031-16503-0 Thefirsteditionwaspublishedintwovolumes #TheEditor(s)(ifapplicable)andTheAuthor(s),underexclusivelicensetoSpringerNature SwitzerlandAG1997,2009,2023 Thisworkissubjecttocopyright.AllrightsaresolelyandexclusivelylicensedbythePublisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting,reuseofillustrations,recitation,broadcasting,reproductiononmicrofilmsorinany otherphysicalway,andtransmissionorinformationstorageandretrieval,electronicadaptation, computersoftware,orbysimilarordissimilarmethodologynowknownorhereafterdeveloped. Theuseofgeneraldescriptivenames,registerednames,trademarks,servicemarks,etc.inthis publication does not imply, even in the absence of a specific statement, that such names are exemptfromtherelevantprotectivelawsandregulationsandthereforefreeforgeneraluse. Thepublisher,theauthors,andtheeditorsaresafetoassumethattheadviceandinformationin thisbookarebelievedtobetrueandaccurateatthedateofpublication.Neitherthepublishernor the authors or the editors give a warranty, expressed or implied, with respect to the material containedhereinorforanyerrorsoromissionsthatmayhavebeenmade.Thepublisherremains neutralwithregardtojurisdictionalclaimsinpublishedmapsandinstitutionalaffiliations. ThisSpringerimprintispublishedbytheregisteredcompanySpringerNatureSwitzerlandAG Theregisteredcompanyaddressis:Gewerbestrasse11,6330Cham,Switzerland Series Preface The Mycota Series—2022 The Mycota is an encyclopaedic book series published by Springer as a comprehensive treatise spanning the Fungal Kingdom through a focus on fungiasexperimentalsystemsforbasicandappliedresearch.Articlesinthis seriespresentintroductoryinformationinacomprehensivemannertobroaden ourgeneralknowledge,alongwiththestate-of-the-artinformationonselected topics in fungal biology. Thus, these articles serve as a timeless reference source for historically relevant discoveries in fungal biology to drive further advancesinthefield. The Mycota was founded in 1994 by Karl Esser and Paul Lemke. Since then15Volumeshavebeenpublished,withseveralnowintheirthirdedition. ThesteadilygrowinginterestinTheMycota,reflectedinparticularbythehigh numberofe-book downloads,encouragedSpringertocontinue this publica- tionwithanewinternationalSeriesEditorialboardbeginningin2022. Historically,thestudyoffungioriginatedasasub-disciplineofbotanyand was a largely descriptive discipline until the early 19th century. Subsequent experimentalresearchhaspropelledthefieldofmycology,andachievements in the genetics and molecular biology of fungi have benefited studies in the relatedfieldsoffungalbiochemistryandbiophysics,plantpathology,medical mycology, and systematics. Ground-breaking research has been carried out using fungi as recognized by the receipt of nine Nobel Prizes, including the revolutionary1945NobelPrizeinPhysiologyandMedicineforthediscovery of penicillin and the foundational 2006 Nobel Prize in Chemistry for establishing the molecular basis of eukaryotic transcription. All such studies expand our knowledge of fungi and of fungal processes and improve our abilitytounderstand,utilize,andcontrolfungiforthebenefitofhumankind. Fungi have invaded every conceivable ecological niche. Saprobic forms abound,especiallyinthedecayoforganicdebris.Pathogenicformsexistwith bothplantandanimalhosts.Fungievengrowonotherfungi.Theyarefound inaquaticaswellassoilenvironments,andtheirsporeshavebeenfoundinthe v vi SeriesPreface airofextremeenvironments.Fungicanbevariouslyassociatedwithplantsas symbiontsintheformoflichens,mycorrhizae,andendophytesandalsooccur asovertpathogens.Associationwithanimalsystemsvaries;examplesinclude the predaceous fungi that trap nematodes, the micro-fungi that grow in the anaerobic environment of the rumen, and medically important pathogens afflictinghumans.Currently,taxonomistshaveidentifiedabout150,000spe- cies, but it is thought that over 90% of fungal species remain undescribed, with conservative estimates ranging from 2 to 5 million fungal species on earth. From this perspective, there are new topics in fungal biology to explore, which will expand the current volumes and take them in new directions. Further, there will be new volumes in areas (e.g., Cryptomycota) that were notcoveredbefore. Ourunderstandingoftheevolutionoffungiisstillincompleteandmainly basedonspeciesthatcanbegrowninculture.ButrecentenvironmentalDNA analyseshaverevealedahighlydiverseformofeukaryoticlifethatbranches withtheFungi,andthustheresulting andhighlydiversecladeswerenamed the Cryptomycota. The discovery of novel intermediate forms will redefine the fungal tree of life. Other topics will consider the adaptation of fungi to climatechanges,theoccurrenceoffungalpathogensintheenvironment,and thedispersaloffungiduringglobalpandemics. Fungal constituents of the microbiome have received much less attention thusfar,yetrecentfindingsfromclinicalandanimalstudiesclearlyestablish fungiasasignificantcomponentoftheoral,gastrointestinal,pulmonary,and skin microbiomes. Finally, the relevance of fungal biology to society is reflectedbytheincreasingnumberoffungalrelatedhumandiseases. Inthehistoryofpharmacology,fungihavealwaysbeensourcesofuseful molecules for humans, but the diversity of molecules that can be obtained fromfungiisstilllargelyunderexplored.However,effortsinfungalgenomics provide resources for data-driven genome mining and large-scale comparisons to explore the molecular repertoires produced by fungi. The result will be new compounds with applications in the pharmaceutical and agriculturalscienceindustries. Fungicanalsohelptheentireplanetandmay,forexample,berelevantin specificsectors,suchasthatlinkedtopollutionfromplastic.Fungiproducea wide range of enzymes that have the potential to break down the chemical bonds of plastic polymers, and in this context the potential role of marine fungi in plastic degradation may be of major relevance. Finally, new biomaterials from fungal species may openthedoortoalternativestofossil- basedmaterials,andthusreduceenvironmentalpollution. Forconsistencythroughout thisseriesofvolumes,thenames adoptedfor major groups of fungi should be followed according to the following paper, which gives an overview of all of the orders in the fungal kingdom: https:// pubmed.ncbi.nlm.nih.gov/32660385/ SeriesPreface vii We are grateful to Springer for continuing The Mycota series and are especially thankful to all of the Volume Editors in selecting topics and assemblingexpertsfromdiversefieldsoffungalbiology. Sydney,Australia DeeCarter Delhi,India AnuradhaChowdhary Durham,USA JosephHeitman Bochum,Germany UlrichKück July15,2022 Volume Preface to the Third Edition Filamentousfungicanformarangeofdifferentassociationswithplantsthat vary from antagonistic through to mutualistic. However, the outcome of an interaction is very dependent on the stage of development of both plant and fungus, the genetic background of both organisms, and the ecological and environmental conditions under which they interact. Consequently, a given species may have a relatively benign interaction under one set of conditions but be severely pathogenic under a different set of conditions or stage of development. Despite this variability, some fungi are more generally recognizedasplantpathogensandothersasplantsymbionts. Itisnowover10yearssincethesecondeditionofTheMycota,VolumeV (Plant Relationships) was published. In the two earlier editions, the volume editors commented on the large number of fungi and vascular plants in the Tree of Life and the enormous number and complexity of the different interactionsthatoccurinnature.Itisthereforechallenginginasinglevolume to capture this range of diversity. However, we hope that through the examples presented in this third edition the reader can gain a sense of the very significant new discoveries that have been made, what key biological questions are being addressed, and where the new frontiers in this field of researchlie.Tohelpthereaderexplorethehugerangeofresearchtopicsand diversityoffungistudied,wehavedividedthevolumeintosixsectionswhich we think capture the most important research themes within this very active andfast-movingareaofresearch. The theme of Chaps. 1–3 is Pathogen Fungus–Plant Interactions. These chaptersprovideagoodoverviewofsomeofthekeyadvancesthathavebeen made in understanding the molecular dialogue between pathogen and host, especially in the field of effector biology. Understanding the biological function of the protein effectors has been particularly challenging as most share little homology with functionally characterized proteins. Despite this challenge,verysignificantadvanceshavebeenmadeinidentifyingthefunc- tion of many effectors, including in some cases the host target proteins with which they interact. One of the most exciting emerging fields has been the demonstration that small RNAs play a crucial role in the dialogue between fungusandplant.Thisis,andwillremain,averyactivefieldofdiscovery. ix x VolumePrefacetotheThirdEdition Chapters4–6(MutualisticFungus–PlantInteractions)provideasnapshot ofthesymbioticinteractionsthatoccurbetweenfungiandplantsbothbelow ground (the arbuscular mycorrhiza) and above ground (leaf endophytes). A hugechallengewiththeformeristheinabilitytoculturemostofthesefungi separatefromtheirhosttherebylimitingthemethodologiesthatcanbeusedto dissect these interesting associations. Recent advances in long-read DNA sequencing have enabled the complete assembly of a number of the large and complex arbuscular mycorrhizal fungal genomes thereby facilitating for thefirsttimenewinsightsintocomparativeandevolutionarygeneticsofthis importantgroup of fungi. The mutualistic interaction ofEpichloë fungi with grasses is now the most well-understood above-ground fungal-plant mutual- isticassociation.Comparativegenomestudiescombinedwithgeneticanalysis and new advances in metabolite chemistry reveal a remarkable diversity of alkaloids synthesized by these seed endophytes. Among the more complex fungal-plant mutualistic symbioses are the lichens—associations between nutritionally specialized fungi and green algae and/or cyanobacteria photobionts.Theseassociationsareverydiversewithapproximately 17%of extant fungi species lichenized. The obligate nature of these associations makesitverychallengingtostudytheindividualcontributionsofthepartners but advances in microscopy and genome analysis have provided significant newinsightsintotheseassociations.Genomeanalysisofthemycobiontshas provided important new insights into the lifestyle of these fungi, and in particular their potential to synthesize a remarkable range of secondary metabolites. The use of heterologous expression systems combined with mass spectrometry analysis has led to the identification of a range of novel lichenmetabolites.Afuturechallengewillbedeterminingtheirbiologicalrole innature. Howfungisensesignalsfromtheirhostandrespondthroughtransduction of those signalsinternally isthetheme ofChaps.8–10(Sensing and Signal- linginFungus–PlantInteractions).Amongthemostwell-characterizedsignal transduction systems are the Mitogen-Activated Protein kinase (MAPK) pathways.Geneticanalysescombinedwithprotein–proteininteractionshave revealed how fungi respond to external stimuli and regulate infection and developmentalprocesses.Thesestudiesdemonstratethatthereareconserved, aswellasspecies-specific,functionsassociatedwiththeMAPKpathways.In addition, there is considerable crosstalk among the pathways, a result that highlights the complexity of signalling associated with plant host coloniza- tion.AmbientpHisoneofthekeyenvironmentalsignalsthatfungiencounter in their environment. Fungi have an exquisitely tuned system to regulate cytosolic pH in response to changes in external pH. Signalling through MAPK pathways control this system and lead to changes in fungal growth, development, and pathogenicity. Plant volatiles are key signals that control fungal invasion of their hosts. Using the Trichoderma system, key volatiles havebeenidentifiedthatcontrolplantcolonizationaswellasplantimmunity anddevelopment. ThethemeofChaps.11–14isRegulationofFungalGeneExpressionand Development.

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