U'Ren Jana M, Arnold A Elizabeth
School of Plant Sciences, University of Arizona, Tucson, AZ, United States of America; Department of Agricultural and Biosystems Engineering, University of Arizona, Tucson, AZ, United States of America.
School of Plant Sciences, University of Arizona, Tucson, AZ, United States of America; Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ, United States of America.
PeerJ. 2016 Dec 13;4:e2768. doi: 10.7717/peerj.2768. eCollection 2016.
Fungal endophytes inhabit symptomless, living tissues of all major plant lineages to form one of earth's most prevalent groups of symbionts. Many reproduce from senesced and/or decomposing leaves and can produce extracellular leaf-degrading enzymes, blurring the line between symbiotrophy and saprotrophy. To better understand the endophyte-saprotroph continuum we compared fungal communities and functional traits of focal strains isolated from living leaves to those isolated from leaves after senescence and decomposition, with a focus on foliage of woody plants in five biogeographic provinces ranging from tundra to subtropical scrub forest.
We cultured fungi from the interior of surface-sterilized leaves that were living at the time of sampling (i.e., endophytes), leaves that were dead and were retained in plant canopies (dead leaf fungi, DLF), and fallen leaves (leaf litter fungi, LLF) from 3-4 species of woody plants in each of five sites in North America. Our sampling encompassed 18 plant species representing two families of Pinophyta and five families of Angiospermae. Diversity and composition of fungal communities within and among leaf life stages, hosts, and sites were compared using ITS-partial LSU rDNA data. We evaluated substrate use and enzyme activity by a subset of fungi isolated only from living tissues vs. fungi isolated only from non-living leaves.
Across the diverse biomes and plant taxa surveyed here, culturable fungi from living leaves were isolated less frequently and were less diverse than those isolated from non-living leaves. Fungal communities in living leaves also differed detectably in composition from communities in dead leaves and leaf litter within focal sites and host taxa, regardless of differential weighting of rare and abundant fungi. All focal isolates grew on cellulose, lignin, and pectin as sole carbon sources, but none displayed ligninolytic or pectinolytic activity . Cellulolytic activity differed among fungal classes. Within Dothideomycetes, activity differed significantly between fungi from living vs. non-living leaves, but such differences were not observed in Sordariomycetes.
Although some fungi with endophytic life stages clearly persist for periods of time in leaves after senescence and incorporation into leaf litter, our sampling across diverse biomes and host lineages detected consistent differences between fungal assemblages in living vs. non-living leaves, reflecting incursion by fungi from the leaf exterior after leaf death and as leaves begin to decompose. However, fungi found only in living leaves do not differ consistently in cellulolytic activity from those fungi detected thus far only in dead leaves. Future analyses should consider Basidiomycota in addition to the Ascomycota fungi evaluated here, and should explore more dimensions of functional traits and persistence to further define the endophytism-to-saprotrophy continuum.
真菌内生菌栖息于所有主要植物谱系无症状的活组织中,形成地球上最普遍的共生生物群体之一。许多真菌内生菌从衰老和/或正在分解的叶片中繁殖,并能产生细胞外叶片降解酶,模糊了共生营养和腐生营养之间的界限。为了更好地理解内生菌 - 腐生菌连续体,我们比较了从活叶中分离出的重点菌株与从衰老和分解后的叶片中分离出的真菌群落及其功能特性,重点关注从苔原到亚热带灌丛森林的五个生物地理省份木本植物的叶片。
我们从采样时仍存活的经表面消毒叶片内部(即内生菌)、已死亡但仍保留在植物冠层中的叶片(死叶真菌,DLF)以及北美五个地点中每个地点3 - 4种木本植物的落叶(落叶真菌,LLF)中培养真菌。我们的采样涵盖了代表松杉纲两个科和木兰纲五个科的18种植物。利用ITS - 部分LSU rDNA数据比较了叶片生命阶段、宿主和地点内及之间真菌群落的多样性和组成。我们评估了仅从活组织中分离出的一部分真菌与仅从非活叶中分离出的真菌的底物利用情况和酶活性。
在此调查的不同生物群落和植物类群中,从活叶中分离出的可培养真菌比从非活叶中分离出的频率更低且多样性更低。无论对稀有和丰富真菌的加权差异如何,活叶中的真菌群落在组成上也与重点地点和宿主类群中的死叶及落叶中的群落有明显差异。所有重点分离菌株都能以纤维素、木质素和果胶作为唯一碳源生长,但均未表现出木质素分解或果胶分解活性。纤维素分解活性在真菌类别之间存在差异。在座囊菌纲中,来自活叶与非活叶的真菌之间活性差异显著,但在粪壳菌纲中未观察到此类差异。
尽管一些具有内生菌生活阶段的真菌在叶片衰老并融入落叶层后显然能在叶片中持续存在一段时间,但我们在不同生物群落和宿主谱系中的采样发现,活叶与非活叶中的真菌组合存在一致差异,这反映了叶片死亡后以及叶片开始分解时真菌从叶片外部的侵入。然而,仅在活叶中发现的真菌在纤维素分解活性方面与迄今仅在死叶中检测到的真菌并无一致差异。未来的分析除了本文评估的子囊菌门真菌外,还应考虑担子菌门,并且应探索功能特性和持久性的更多维度,以进一步界定从内生菌到腐生菌的连续体。