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Cytospora ceratosperma on Juglans regia (BJFC-S774). a-b. Habit of conidiomata on branch; c. transverse section through conidioma; d. longitudinal section through conidioma; e. conidiogenous cells with attached conidia; f. conidia. -Scale bars: a = 1 mm; b -d = 0.5 mm; e -f = 10 μm.
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Members of the genus Cytospora are often reported as endophytes, saprobes or phytopathogens, primarily causing canker diseases of woody host plants. They occur on a wide range of hosts and have a worldwide
distribution. Although several species have in the past been reported from China, the vast majority are not known
from culture or DNA phylogeny....
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Citations
... Under the current nomenclatural code, the teleomorphic genus Valsa is currently treated as a synonym of the anamorphic genus Cytospora, which is the oldest name among Cytospora and its related teleomorphic genera Rossman et al., 2015). The genus Cytospora is currently undergoing taxonomic revision and many new species have been described worldwide (e.g., Adams et al., 2002Adams et al., , 2005Adams et al., , 2006Fan et al., 2014Fan et al., , 2020Fotouhifar et al., 2010;Gao et al., 2021;Ilyukhin et al., 2023;Jami et al., 2018;Lawrence et al., 2018;Norphanphoun et al., 2018;Pan et al., 2020;Travadon et al., 2022;Úrbez-Torres et al., 2020). On the other hand, although they are important pathogens, the taxonomic positions of Valsa and Cytospora species originally described in Japan, such as V. paulowniae, have not been fully clarified and need reappraisal. ...
Paulownia tree canker is a major disease of Paulowniae tomentosa in Japan. The pathogen was described as Valsa paulowniae in 1916 by Hemmi and Miyabe. However, its current taxonomic status and phylogenetic position are uncertain. In this study, we reviewed the protologue of this species and rediscovered the syntypes maintained at the Hokkaido University Museum (SAPA). From these specimens, a lectotype was selected. The molecular phylogenetic position of this species was examined with newly collected samples. Based on the result of phylogeny and morphology, an epitype of this species was designated and transferred to the genus Cytospora.
... Members of diaporthalean fungi are associated with several diseases including canker and dieback in economically and ecologically important woody plants such as Quercus species (Luque et al., 2000;Lynch et al., 2014;Fan et al., 2018;Jiang et al., 2018Jiang et al., , 2019Zhu et al., 2019Zhu et al., , 2021. Among the diaporthalean fungi, Cytospora with approximately 700 species listed in Index Fungorum (November 2023; www.indexfungorum.org) is the most common and widespread genus associated with a wide variety of woody plants around the world, which causes various disease symptoms such as canker and dieback or found as endophyte and saprobe (Adams et al., 2005;Lawrence et al., 2018;Azizi et al., 2020;Fan et al., 2020;Hanifeh et al., 2022;Ilyukhin et al., 2023). Cytospora species are found in association with canker and dieback diseases on various Quercus species (Kowalski, 1991;Adams et al., 2005;Lynch et al., 2014;Fan et al., 2018;Lawrence et al., 2018;Pinna et al., 2019;Pan et al., 2021). ...
Oak decline is a complex disorder that seriously threatens the survival of Zagros forests. In an extensive study on taxonomy and pathology of fungi associated with oak decline in the central and northern part of Zagros forests, 462 fungal isolates were obtained from oak trees showing canker, gummosis, dieback, defoliation, and partial or total death symptoms. Based on inter-simple sequence repeat (ISSR) fingerprinting patterns, morphological characteristics, and sequences of ribosomal DNA (28S rDNA and ITS) and protein coding loci (acl1, act1, caM, tef-1α, rpb1, rpb2, and tub2), 24 fungal species corresponding to 19 genera were characterized. Forty percent of the isolates were placed in eight coelomycetous species from seven genera, namely, Alloeutypa, Botryosphaeria, Cytospora, Didymella, Gnomoniopsis, Kalmusia, and Neoscytalidium. Of these, four species are new to science, which are introduced here as taxonomic novelties: Alloeutypa iranensis sp. nov., Cytospora hedjaroudei sp. nov., Cytospora zagrosensis sp. nov., and Gnomoniopsis quercicola sp. nov. According to pathogenicity trials on leaves and stems of 2-year-old Persian oak (Quercus brantii) seedlings, Alternaria spp. (A. alternata, A. atra, and A. contlous), Chaetomium globosum, and Parachaetomium perlucidum were recognized as nonpathogenic. All coelomycetous species were determined as pathogenic in both pathogenicity trials on leaves and seedling stems, of which Gnomoniopsis quercicola sp. nov., Botryosphaeria dothidea, and Neoscytalidium dimidiatum were recognized as the most virulent species followed by Biscogniauxia rosacearum.
... Cytospora has priority over Leucocytospora, Leucostoma, Valsa, Valsella and Valseutypella based on the dual-nomenclature criterion (Adams et al. 2005;Rossman et al. 2015). Members of Cytospora are characterised by the single or labyrinthine, loculate stromata, filamentous conidiophores or asci and allantoid hyaline conidia or ascospores (Spielman 1985;Adams et al. 2005;Norphanphoun et al. 2017Norphanphoun et al. , 2018Fan et al. 2020;Shang et al. 2020). Species identification in Cytospora was previously largely based on the host affiliation and morphological descriptions; however, molecular phylogeny combined with morphology and host affiliation have became the main approaches recently Shang et al. 2020;Zhu et al. 2020). ...
... 2023). However, most of these were regarded as synonyms and most descriptions were unable to identify them accurately (Adams et al. 2005;Fan et al. 2020;Pan et al. 2020). ...
... Species of Cytospora are known as opportunistic pathogens mainly infecting woody hosts and some of the species occur on a wide host range (Adams et al. 2005;Fan et al. 2020). The Cytospora species and their host association have been revealed in this study; however, further studies are required to confirm the fungal pathogenicity. ...
During our biodiversity investigations in Tibet, China, typical Cytospora canker symptoms were observed on branches of hosts Myricaria paniculate, Prunus cerasifera and Sibiraea angustata. Samples were studied, based on morphological features coupled with multigene phylogenetic analyses of ITS, act, rpb2, tef1 and tub2 sequence data, which revealed two new species (Cytospora myricicolasp. nov. and C. sibiraeicolasp. nov.) and a known species (C. populina). In addition, Cytospora populina is newly discovered on the host Prunus cerasifera and in Tibet.
... The results of our phylogenetic analyses using ITS-rDNA sequences placed C. balanejica together with C. albodisca, C. corylina, and C. olivacea in an unresolved clade, confirmed the poor utility of this genomic region in the differentiation of Cytospora species 21,38,61,62 ( Supplementary Fig. S1). Recent studies using a polyphasic approach, morphology, and multi-gene phylogeny, have revealed hidden fungal diversity, and led to the description of several new cryptic Cytospora species 21,23,26,38,41 . Based on our multi-gene phylogenetic analyses, C. balanejica formed a well-defined monophyletic lineage distinct from all other strains with close affinity to C. albodisca and C. corylina, two recently described Cytospora species (Fig. 2). ...
Apple is the most important fruit tree in West Azarbaijan province of Iran. In a survey of apple orchards, a disease with crown and collar canker and necrosis symptoms was observed in three young apple orchards in Urmia, affecting 15% and 1% of ‘Red Delicious’ and ‘Golden Delicious’ cultivars, respectively. A fungus with typical characteristics of the asexual morph of Cytospora was regularly isolated from the diseased tissues. Morphological characteristics and phylogenetic analyses inferred from the combined dataset of the ITS-rDNA, parts of LSU, tef1-α, rpb2, and act1 genes revealed that the isolates represent a new species of Cytospora, described herein as Cytospora balanejica sp. nov.. The pathogenicity of all isolates was confirmed on apple cv. ‘Red Delicious’ based on Koch’s postulates. Also, the reaction of 12 other apple cultivars was assessed against five selected isolates with the highest virulence. The results showed that except for cv. ‘Braeburn’, which did not produce any symptoms of the disease, the other 11 cultivars showed characteristic disease symptoms including sunken and discolored bark and wood. The mean length of the discolored area was different among the 11 so-called susceptible cultivars, hence cvs. ‘M4’ and ‘Golden Delicious’ showed the highest and the lowest lesion length, respectively. Moreover, the aggressiveness of the five tested isolates was different, and the isolates BA 2-4 and BA 3-1 had the highest and lowest aggressiveness, respectively. Based on our observations on the potential ability of the fungus to cause disease on young and actively growing apple trees, it will be a serious threat to apple cultivation and industry.
... The genus Cytospora was established by Ehrenberg (1818) and classified in Cytosporaceae, Diaporthales, Sordariomycetes (Wijayawardene et al. 2018;Fan et al. 2020). It includes numerous important pathogens associated with canker and dieback diseases of woody plants, with a worldwide distribution and broad host range (Sinclair et al. 1987;Adams et al. 2005Adams et al. , 2006Lawrence et al. 2018;Fan et al. 2020;Lin et al. 2023a, b). ...
... The genus Cytospora was established by Ehrenberg (1818) and classified in Cytosporaceae, Diaporthales, Sordariomycetes (Wijayawardene et al. 2018;Fan et al. 2020). It includes numerous important pathogens associated with canker and dieback diseases of woody plants, with a worldwide distribution and broad host range (Sinclair et al. 1987;Adams et al. 2005Adams et al. , 2006Lawrence et al. 2018;Fan et al. 2020;Lin et al. 2023a, b). Dieback and stem canker caused by Cytospora lead to the growth weakness or death of host plants, thereby causing significant economic and ecological losses (Sinclair et al. 1987;Adams et al. 2005). ...
... Currently, use of polyphasic approaches, such as morphological and phylogenetic analyses to define species of Cytospora has been proposed (Norphanphoun et al. 2017;Fan et al. 2020). In morphology, presence or absence of conceptacle, quantity and arrangement of locule(s), shape and size of conidiophores and conidial size are significantly taxonomic. ...
Members of Cytospora include saprobes, endophytes and important plant pathogens, which are widely distributed on various wood hosts and have a wide global distribution. In this study, the species definitions were conducted, based on multigene phylogeny (ITS, act , rpb2 , tef1-α and tub2 genes) and comparisons of morphological characters. A total of 22 representative isolates obtained from 21 specimens in Fengtai District of Beijing City were identified as seven species of Cytospora , including four known species ( C. albodisca , C. ailanthicola , C. euonymina , C. haidianensis ) and three novel species ( C. fengtaiensis , C. pinea , C. sorbariae ). The results provide an understanding of the taxonomy of Cytospora species associated with canker and dieback diseases in Fengtai District, Beijing, China.
... Particularly, it is worth noting that the host plants of most well-studied phytopathogenic fungi are crops. Cytospora chrysosperma, the causal agent of poplar canker disease, results in significant economic and ecological losses in China every year (Fan et al. 2020). C. chrysosperma infects woody plants through micro-wounds and various symptoms are observed on different host species and at different stages of disease development (Biggs et al. 1983;Fan et al. 2020;Kepley et al. 2015). ...
... Cytospora chrysosperma, the causal agent of poplar canker disease, results in significant economic and ecological losses in China every year (Fan et al. 2020). C. chrysosperma infects woody plants through micro-wounds and various symptoms are observed on different host species and at different stages of disease development (Biggs et al. 1983;Fan et al. 2020;Kepley et al. 2015). Despite the high economic impact of poplar canker disease, efficient strategies for poplar canker disease management have not been well established. ...
The pathogen Cytospora chrysosperma is the causal agent of poplar canker disease and causes
considerable eco-
nomic losses in China. Mitogen-activated protein kinase (MAPK) cascades play a crucial role in
mediating cellular responses and Pmk1-MAPKs are indispensable for pathogenic related processes in
plant pathogenic fungi. In previous studies, we demonstrated that the CcPmk1 acts as a core
regulator of fungal pathogenicity by modulating a small number of master downstream targets, such
as CcSte12. In this study, we identified and characterized two upstream components of CcPmk1:
MAPKKK CcSte11 and MAPKK CcSte7. Deletion of CcSte11 and CcSte7, resulted in slowed growth, loss of
sporulation and virulence, similar to the defects observed in the CcPmk1 deletion mutant. In addi-
tion, CcSte11, CcSte7 and CcPmk1 interact with each other, and the upstream adaptor protein CcSte50
interact
with CcSte11 and CcSte7. Moreover, we explored the global regulation network of CcSte12 by
transcriptional analysis between CcSte12 deletion mutants and wild-type during the simulated
infection process. Two hydrolase activity GO terms (GO:0004553 and GO:0016798) and starch and
sucrose metabolism (mgr00500) KEGG pathway were signifi- cantly enriched in the down-regulated
genes of CcSte12 deletion mutants. In addition, a subset of glycosyl hydrolase genes and putative
effector genes were significantly down-regulated in the CcSte12 deletion mutant, which might be
important for fungal pathogenicity. Especially, CcSte12 bound to the CcSp84 promoter region
containing the TGAAAC A motif. Moreover, comparison of CcSte12-regulated genes with
CcPmk1-regulated genes revealed 116 overlapping regulated genes in both CcSte12 and CcPmk1,
including some virulence-associated genes. Taken together, the protein complexes
CcSte11-CcSte7-CcPmk1 receive signals transmitted by upstream CcSte50 and transmit signals to down-
stream CcSte12, which regulates hydrolase, effectors and other genes to promote virulence. Overall,
these results indicate that the CcPmk1-MAPK signaling pathway of C. chrysosperma plays a key role
in the pathogenicity.
Keywords Cytospora chrysosperma, MAPK, Transcription factor Ste12, Pathogenicity, Transcriptional
analysis
... Particularly, it is worth noting that the host plants of most well-studied phytopathogenic fungi are crops. Cytospora chrysosperma, the causal agent of poplar canker disease, results in significant economic and ecological losses in China every year (Fan et al. 2020). C. chrysosperma infects woody plants through micro-wounds and various symptoms are observed on different host species and at different stages of disease development (Biggs et al. 1983;Fan et al. 2020;Kepley et al. 2015). ...
... Cytospora chrysosperma, the causal agent of poplar canker disease, results in significant economic and ecological losses in China every year (Fan et al. 2020). C. chrysosperma infects woody plants through micro-wounds and various symptoms are observed on different host species and at different stages of disease development (Biggs et al. 1983;Fan et al. 2020;Kepley et al. 2015). Despite the high economic impact of poplar canker disease, efficient strategies for poplar canker disease management have not been well established. ...
The pathogen Cytospora chrysosperma is the causal agent of poplar canker disease and causes considerable economic losses in China. Mitogen-activated protein kinase (MAPK) cascades play a crucial role in mediating cellular responses and Pmk1-MAPKs are indispensable for pathogenic related processes in plant pathogenic fungi. In previous studies, we demonstrated that the CcPmk1 acts as a core regulator of fungal pathogenicity by modulating a small number of master downstream targets, such as CcSte12. In this study, we identified and characterized two upstream components of CcPmk1: MAPKKK CcSte11 and MAPKK CcSte7. Deletion of CcSte11 and CcSte7, resulted in slowed growth, loss of sporulation and virulence, similar to the defects observed in the CcPmk1 deletion mutant. In addition, CcSte11, CcSte7 and CcPmk1 interact with each other, and the upstream adaptor protein CcSte50 interact with CcSte11 and CcSte7. Moreover, we explored the global regulation network of CcSte12 by transcriptional analysis between CcSte12 deletion mutants and wild-type during the simulated infection process. Two hydrolase activity GO terms (GO:0004553 and GO:0016798) and starch and sucrose metabolism (mgr00500) KEGG pathway were significantly enriched in the down-regulated genes of CcSte12 deletion mutants. In addition, a subset of glycosyl hydrolase genes and putative effector genes were significantly down-regulated in the CcSte12 deletion mutant, which might be important for fungal pathogenicity. Especially, CcSte12 bound to the CcSp84 promoter region containing the TGAAACA motif. Moreover, comparison of CcSte12-regulated genes with CcPmk1-regulated genes revealed 116 overlapping regulated genes in both CcSte12 and CcPmk1, including some virulence-associated genes. Taken together, the protein complexes CcSte11-CcSte7-CcPmk1 receive signals transmitted by upstream CcSte50 and transmit signals to downstream CcSte12, which regulates hydrolase, effectors and other genes to promote virulence. Overall, these results indicate that the CcPmk1-MAPK signaling pathway of C. chrysosperma plays a key role in the pathogenicity.
Supplementary Information
The online version contains supplementary material available at 10.1007/s44154-023-00142-w.
... Of these, Cytospora is common on the cankered branches of woody plants, with C. chrysosperma (Pers.) Fr. widely causing polar and willow diseases in China [9]. ...
... The obtained sequences for ITS, act, rpb2, tef1 and tub2 from the present study and related sequence data from recent publications (Supplementary file) [9,24,25] were aligned using the MAFFT v.7 online web server (http://mafft.cbrc.jp/alignment/server/index.html, accessed on 1 November 2023) under the default settings. ...
... Culture characteristics: Colonies on PDA are flat, spreading, with sparse to moderate aerial mycelium, initially white, becoming umber after 15 days, reaching a 90 mm diameter after 15 days and forming abundant conidiomata after 30 days at 25 • C in the dark. Notes: Cytospora euonymina was initially described from diseased branches and twigs of Euonymus kiautschovicus collected from Shanxi Province [9], and was subsequently discovered on leaf spots of E. japonicus in Beijing City [25]. This fungal species has only been reported from the host genus Euonymus and in China. ...
Branch canker diseases are important in forest ecosystems, causing economic and ecological losses. Members of Cytospora are common on cankered branches and associated with the diseases. Investigations on Cytospora cankers were conducted in Tibet, China, during 2022 and 2023. Samples were collected from Euonymus japonicus, Larix gmelinii, Malus pumila, M. spectabilis and Rosa omeiensis f. pteracantha, and cultures were obtained and identified by morphological features and molecular phylogeny of a combination of internal transcribed spacer region rDNA (ITS), the partial actin (act) region, RNA polymerase II second largest subunit (rpb2) gene, the translation elongation factor 1-alpha (tef1) gene and the partial beta-tubulin (tub2) gene. As a result, a new species is proposed herein named Cytospora lhasaensis and four known species are described for the first time from Tibet, viz. C. euonymina, C. gigaspora, C. mali and C. schulzeri. The current research enhances our understanding of the Cytospora species associated with woody host diseases in Tibet, China.
... In the past, species identification in Cytospora was largely based on host affiliation, morphological characters of pycnidia/perithecia, and spore dimensions, with morphological description [26][27][28]. This morphological species approach is confounded by many morphological characters' overlap among species and by the morphological plasticity of pycnidial locules which are affected by the host bark and cambium characteristics (e.g., stromatal arrangement in the host tissues, locular arrangement within pycnidia, locule division into chambers, independent or shared locular walls) [18,19,[29][30][31]. ...
... As plant pathogens, Cytospora species are primarily associated with canker diseases. Symptoms vary with host species and stage of development [26]. Cytospora species mainly impact branches, but they can cause more destructive infections in the trunk and larger scaffolds, consequently limiting the productivity of orchards [38,39]. ...
... The wood below the cambium is stained brown. Later, these fungi quickly girdle and kill branches and twigs, forming several black sporocarps [26]. ...
Olive (Olea europaea L.) is a very important crop grown in the Mediterranean part of Croatia. Olive branch and fruit dieback symptoms were observed in two olive orchards in Istria, Croatia. The samples from symptomatic trees were collected and brought to the laboratory for analysis. Based on their morphological characterization, isolated fungi were identified as Cytospora sp. Two
representative isolates (one per orchard) were taken for molecular analysis, and based on DNA sequence data of the ITS and TUB gene regions, and phylogenetic analysis of the sequences, the isolates were identified as Cytospora pruinosa Défago. To determine pathogenicity, pathogenicity tests were conducted on detached olive branches and two-year-old olive trees in the greenhouse. This is
the first report of C. pruinosa causing olive branch and fruit dieback in Croatia.
... the poplar canker; it is widespread in poplar plantations and results in huge annual economic losses in northern China (10). One strategy used by C. chrysosperma to establish infection is the colonization first restricted to injured or dying bark in healthy poplar, then the hyphae rapidly and expansively spread in poplar stem once the tree vigor becomes weak (9,11). Fungal cell walls play an essential role in plant-pathogen interactions (12). ...
Maintenance of cell wall integrity is important for fungal cell morphology against external stresses and even virulence. Although the transcription factor Rlm1 is known to play major regulatory roles in the maintenance of cell integrity, the underlying mechanism of how Rlm1 contributes to cell wall integrity and virulence in phytopathogenic fungi remains unclear. Here, we demonstrated that CcRlm1 plays important roles in cell wall maintenance and virulence in the poplar canker fungus Cytospora chrysosperma. Among putative downstream targets, CcChs6 (chitin synthase) and CcGna1 (glucosamine 6-phosphate N-acetyltransferase) were found to be direct targets of CcRlm1 and shown to function in chitin synthesis and virulence. Furthermore, we found stronger induction of poplar defense responses when challenged with these gene deletion mutants. Collectively, these results suggest that CcRlm1 plays a critical role in the regulation of cell wall maintenance, stress response, and virulence by directly regulating CcChs6 and CcGna1 in C. chrysosperma.
IMPORTANCE Cytospora chrysosperma causes canker diseases on woody plants, and the molecular basis of its infection is not well understood. This study shows that CcRlm1 is the major regulator of chitin synthesis and virulence of the poplar canker fungus. Our research contributes to further understanding the molecular basis of the interaction between C. chrysosperma and poplar.
