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Morphological variations at different stages of development of Morchella importuna (photos taken by Z.L. Yang).
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... However, due to morphological plasticity and intraspecific variability [4][5][6], the microscopic characteristics of Morchella at different developmental stages differ. The morphology and color of Morchella species are highly variable [8], which complicates their delimitation and characterization [9][10][11][12][13]. It is quite difficult to categorize and identify Morchella species on the basis of traditional morphological classification. ...
True morels (Morchella) are a well-known edible fungi, with economically and medicinally important values. However, molecular identification and species taxonomy of the genus Morchella have long been controversial, due to numerous intermediate morphologies among species. In this study, we determined the identification efficiency of DNA barcoding and species classification of 260 individuals from 45 Morchella species, on the basis of multiple nuclear DNA markers. DNA barcoding analysis showed that the individual DNA fragment has a lower resolution of species identification than that of combined multiple DNA markers. ITS showed the highest level of species discrimination among the individual genetic markers. Interestingly, the combined DNA markers significantly increased the resolution of species identification. A combination of four DNA genes (EF1-α, RPB1, RPB2 and ITS) showed a higher species delimitation than that any combination of two or three markers. Phylogenetic analysis suggested that the species in genus Morchella could have been divided into two large genetic clades, the Elata Clade and Esculenta Clade lineages. The two lineages divided approximately 133.11 Mya [95% HPD interval: 82.77–197.95] in the early Cretaceous period. However, some phylogenetic species of Morchella showed inconsistent evolutionary relationships with the traditional morphological classifications, which may have resulted from incomplete lineage sorting and/or introgressive hybridization among species. These findings demonstrate that the interspecific gene introgression may have affected the species identification of true morels, and that the combined DNA markers significantly improve the resolution of species discrimination.
... M.importuna, as one of the few species of Morchella, has been successfully domesticated by mushroom farmers in China in recent years, and its cultivation area accounts for over 95% of the artificial cultivation area (Du et al., 2012;Wen et al., 2021). Morchella dried products are divided into four grades: Grade 1, Grade 2, Export grade and non-graded dried products (Du et al., 2014) [Supplemental content 1]. The fruiting bodies of M. importuna contain rich dietary fiber (Xu et al., 2016). ...
Soluble dietary fiber was extracted from spent Morchella importuna by optimizing the extraction conditions using the response surface method. Response surface analysis results indicated that the extraction yield of MI-SDF reached up to 15.38 ± 0.16% under the optimized conditions with optimal lye concentration of 2.9%, extraction temperature of 71°C and extraction time of 2 h. The results of scanning electron microscope (SEM), Fourier transform infrared spectroscopy (FT-IR), and X-ray diffraction analyses suggested that M. importuna-derived soluble dietary fiber (MI-SDF) exhibited the typical porous fiber structures, polysaccharide functional groups, and cellulose crystal structure. And MI-SDF was applicable to food processing at <250°C. During in vitro fecal fermentation, MI-SDF significantly increased the abundance and diversity of beneficial bacteria and decreased those of harmful bacteria. The levels of total and individual short-chain fatty acids were significantly increased. This study will provide valuable information for further investigation of MI-SDF.
... However, within the group, Morchella exhibit uncertainties in taxonomy due to highly polymorphic fruiting bodies, even molecular phylogenetic techniques give rise to new controversies in taxonomy (Loizides 2017). Due to insufficient microscopic characteristics and high levels of variability in form and color of ascocarps (Du et al. 2014), affected by ecological and climate factors, the species number in Morchella varies from 3 to 50 or more, which has caused confusing use of homonyms and synonyms (Du et al. 2015). Most of them were described from Europe and the USA, with only a few described in Asia. ...
... China tops the list of the world's largest producers of mushrooms and truffles with an annual production of about four million tons. The Yunnan province is the leading producer of wild mushrooms, including morals in China (Cunningham 2012;Du et al. 2014Du et al. , 2015. A large quantity of it is harvested commercially for internal consumption that is sold in large mushroom markets and exported extensively (Du et al. 2015;Figs. ...
Morchella esculenta(Fr.) I.R. Hazll: Phallus esculentus L.; Helvella esculenta (L.) Sowerby; Morellus esculentus (L.) Eaton; Morilla esculenta (L.) Quél.; Phalloboletus esculentus (L.) Kuntze; Morchella rotunda var. esculenta (L.) Jacquet.; Morchella prunarii Schulzer; Morchella esculenta var. prunarii (Schulzer & Hazsl.) Sacc.; Phallus esculentus var. albus Bull.; Morchella vulgaris var. alba (Bull.) Clowez; Phallus esculentus var. cinereus Bull.; Phallus esculentus var. fuscus Bull.; Phallus tremelloides Vent.; Morchella tremelloides (Vent.) Pers.; Morilla tremelloides (Vent.) Quél.; Morchella vulgaris var. tremelloides (Vent.) Boud. [as ‘tremelloïdes’]; Phallus esculentus var. rotundus Pers.; Morchella esculenta var. rotunda (Pers.) Sacc.; Morchella rotunda (Pers.) Boud.; Morchella esculenta f. rotunda (Pers.) Reichert; Morchella conica Pers.; Morchella esculenta var. conica (Pers.) Fr.; Morilla conica (Pers.) Quél.; Morchella esculenta var. alba Mérat; Morchella rotunda var. alba (Mérat) Sacc.; Morchella esculenta var. cinerea Mérat; Morchella esculenta var. fulva Fr.; Morchella esculenta subsp. pubescens Pers.; Morchella pubescens (Pers.) Krombh.; Morchella rotunda var. pubescens (Pers.) Boud.; Morchella esculenta var. pubescens (Pers.) Sacc. & Traverso; Morchella esculenta var. grisea Pers.; Morchella conica var. metheformis Pers.; Morchella esculenta var. stipitata Lenz; Morchella conica var. pusilla Krombh.; Morchella conica var. rigida Krombh.; Morchella rigida (Krombh.) Boud.; Morchella rotunda var. rigida (Krombh.) Jacquet.; Morchella esculenta var. rigida (Krombh.) I.R. Hall; Morchella conica var. ceracea Krombh.; Morchella esculenta var. violacea Lév.; Morchella distans Fr.; Morchella conica var. distans (Fr.) Clowez; Morchella esculenta var. corrugata Sacc.; Morchella esculenta var. ovalis Fr. ex Sacc.; Morchella viridis Leuba; Morchella esculenta var. viridis (Leuba) Sacc. & D. Sacc.; Morchella abietina Leuba; Morchella esculenta var. abietina (Leuba) Sacc. & Trotter; Morchella lutescens Leuba; Morchella esculenta var. lutescens (Leuba) Sacc. & Traverso; Morchella conica var. elata Henn.; Morchella rotunda var. cinerea Boud.; Morchella umbrina Boud.; Morchella esculenta var. umbrina (Boud.) S. Imai; Morchella vulgaris var. albida Boud.; Morchella esculenta var. albida (Boud.) Sacc.; Morchella vulgaris var. cinerascens Boud.; Morchella rotunda var. alba Boud.; Morchella vulgaris var. alba Boud.; Morchella conica var. serotina Peck; Morchella conica var. angusticeps Peck; Morchella cylindrica Velen.; Morchella conica f. cylindrica (Velen.) Svrček; Morchella rotunda var. cinerea Grelet; Morchella rotunda var. fulva Grelet; Morchella vulgaris var. parvula Bánhegyi; Morchella esculenta var. atrotomentosa M.M. Moser; Morchella distans f. longissima Jacquet.; Morchella distans f. spathulata Jacquet.; Morchella umbrina f. macroalveola Jacquet.; Morchella esculenta f. alba Galli; Morchella rotunda var. pallida Jacquet.; Morchella rotunda var. crassipes Jacquet.; Morchella rotunda var. minutula Jacquet.; Morchella rotunda var. pallida Jacquet.; Morchella ovalis f. pallida (Jacquet.) Clowez & Luc Martin; Morchella esculenta f. dunensis Castañera; Morchella dunensis (Castañera, J.L. Alonso & G. Moreno) Clowez; Morchella esculenta var. dunensis (Castañera, J.L. Alonso & G. Moreno) Blanco-Dios; Morchella dunensis f. sterilis Picón [as ‘sterile’]; Morchella esculenta f. sterilis (Picón) Blanco-Dios; Morchella vulgaris var. aucupariae Clowez & J.-M. Moingeon; Morchella esculenta var. aurantiaca Clowez; Morchella esculenta var. brunnea Clowez; Morchella esculenta var. mahoniae Clowez & R. Durand bis; Morchella esculenta var. roseostraminea Clowez; Morchella esculenta var. rubroris Clowez & Luc Martin; Morchella esculenta var. theobromichroa Clowez & Vanhille; Morchella esculenta var. umbrinoides Clowez; Morchella vulgaris var. atrogrisea Clowez; Morchella vulgaris var. griseosordida Clowez & Franç. Petit; Morchella conica var. cilicicae Clowez, Baş Serm. & Işıloğlu; Morchella conica var. crassa Clowez; Morchella conica var. flexuosa Clowez & Luc Martin; Morchella conica var. meandriformis Clowez & Moyne; Morchella conica var. nigra Clowez & Moyne; Morchella conica var. pygmaea Clowez & Delaunoy; Morchella conica var. violeipes Clowez & Y. Mourgues; Morchella vulgaris var. parvipilea Clowez
... However, within the group, Morchella exhibit uncertainties in taxonomy due to highly polymorphic fruiting bodies, even molecular phylogenetic techniques give rise to new controversies in taxonomy (Loizides 2017). Due to insufficient microscopic characteristics and high levels of variability in form and color of ascocarps (Du et al. 2014), affected by ecological and climate factors, the species number in Morchella varies from 3 to 50 or more, which has caused confusing use of homonyms and synonyms (Du et al. 2015). Most of them were described from Europe and the USA, with only a few described in Asia. ...
... China tops the list of the world's largest producers of mushrooms and truffles with an annual production of about four million tons. The Yunnan province is the leading producer of wild mushrooms, including morals in China (Cunningham 2012;Du et al. 2014Du et al. , 2015. A large quantity of it is harvested commercially for internal consumption that is sold in large mushroom markets and exported extensively (Du et al. 2015;Figs. ...
... However, within the group, Morchella exhibit uncertainties in taxonomy due to highly polymorphic fruiting bodies, even molecular phylogenetic techniques give rise to new controversies in taxonomy (Loizides 2017). Due to insufficient microscopic characteristics and high levels of variability in form and color of ascocarps (Du et al. 2014), affected by ecological and climate factors, the species number in Morchella varies from 3 to 50 or more, which has caused confusing use of homonyms and synonyms (Du et al. 2015). Most of them were described from Europe and the USA, with only a few described in Asia. ...
... China tops the list of the world's largest producers of mushrooms and truffles with an annual production of about four million tons. The Yunnan province is the leading producer of wild mushrooms, including morals in China (Cunningham 2012;Du et al. 2014Du et al. , 2015. A large quantity of it is harvested commercially for internal consumption that is sold in large mushroom markets and exported extensively (Du et al. 2015;Figs. ...
... However, within the group, Morchella exhibit uncertainties in taxonomy due to highly polymorphic fruiting bodies, even molecular phylogenetic techniques give rise to new controversies in taxonomy (Loizides 2017). Due to insufficient microscopic characteristics and high levels of variability in form and color of ascocarps (Du et al. 2014), affected by ecological and climate factors, the species number in Morchella varies from 3 to 50 or more, which has caused confusing use of homonyms and synonyms (Du et al. 2015). Most of them were described from Europe and the USA, with only a few described in Asia. ...
... China tops the list of the world's largest producers of mushrooms and truffles with an annual production of about four million tons. The Yunnan province is the leading producer of wild mushrooms, including morals in China (Cunningham 2012;Du et al. 2014Du et al. , 2015. A large quantity of it is harvested commercially for internal consumption that is sold in large mushroom markets and exported extensively (Du et al. 2015;Figs. ...
... As a food source, the morel is considered a nutritious food of high nutritional values as fish or meat in Asia. As a medicinal mushroom, the morel is recorded as "sweet, cold and non-toxic, beneficial to the stomach, reducing phlegm and promoting the circulation of qi" in "Ben-Cao-Gang-Mu," a Chinese outline treatise of medical herbs [2]. Currently, morels are mainly harvested from the wild in China, India, Turkey, Mexico, and the USA [3]. ...
Morels (Morchella spp.) are valuable medicinal and edible mushrooms. In this study, chemical profiles of aromas and lipophilic extracts of black morel (Morchella importuna) grown in China were analyzed by gas chromatography/mass spectroscopy, along with the evaluation of antioxidant and antimicrobial activities for the lipophilic extracts. Sixty-five compounds in total were identified from the aromas, and 1-octen-3-ol was the main component for aromas of fresh (34.40%) and freeze-dried (68.61%) black morels, while the most abundant compound for the aroma of the oven-dried sample was 2(5H)-furanone (13.95%). From the lipophilic extracts, 29 compounds were identified with linoleic acid as the main compound for fresh (77.37%) and freeze-dried (56.46%) black morels and steroids (92.41%) as the main constituent for an oven-dried sample. All three lipophilic extracts showed moderate antioxidant activities against 2,2-diphenyl-1-picrylhydrazyl and 2,2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic acid) with the IC50 values ranging 7.56 ∼ 17.52 mg/mL and 5.75 ∼ 9.73 mg/mL, respectively, and no obvious antimicrobial activity was observed for lipophilic extracts. The drying methods affect the chemical profile of black morel, and freeze-drying was favorable for retaining nutrients and morel smell. This is the first report on the aroma and lipophilic extracts of M. importuna grown in China.
... The fruiting bodies of morels are arguably the most highly prized and sought after, not only for their delicious taste, unique appearance, and nutritional value, but also for their medicinal properties (Cai et al. 2018;Tietel and Masaphy 2018a). In recent years, significant progress has been made in understanding their nutritional benefits (Shameem et al. 2017;Tietel and Masaphy 2018b), biological characteristics (Xie et al. 2014;He et al. 2017), species diversity (Richard et al. 2015;Loizides et al. 2016), and evolutionary history (Du et al. 2014;Liu et al. 2018b). Morel cultivation has been a research focus worldwide for more than 100 years, and the artificial cultivation of M. importuna, M. sextelata, and M. eximia was achieved in China as of 2012 (He et al. 2018;Liu et al. 2018a). ...
True morels (Morchella spp.) are a group of edible fungi that are in high demand worldwide. However, this group of fungi remains poorly understood in terms of its genetic structure, life cycle, and mating system. In the present study, we cloned the MAT locus in Morchella sp. Mes-20 using long-range polymerase chain reaction (PCR) amplification. Our results showed that the MAT1-2 idiomorph in the single-ascospore isolate YAASMCB-3 was 7.5 kb in length, harboring a single MAT1-2-1 gene, whereas the MAT1-1 idiomorph in the isolate YAASMCB-15 was 7.8 kb, carrying both MAT1-1-1 and MAT1-1-10 genes. Mating-type diagnostic assays of single-ascospore populations confirmed that Mes-20 is heterothallic. In addition, 42 collections belonging to 17 morel species (7 black morel species [Elata clade], 9 yellow morel species [Esculenta clade], and 1 species from the Rufobrunnea clade) were used to clone and characterize the MAT1-1-1 and MAT1-2-1 genes for development of the mating gene genealogies. In concordance with the multilocus phylogenetic trees, collections for the respective species were readily distinguished by well-supported lineages in mating gene genealogies. The topologies were consistent within the Elata clade, whereas the branching order and sister-group relationships slightly varied within the Esculenta clade. Our results show that species in the genus Morchella could be delimited by mating gene genealogies.
... Nowadays, M. esculenta is mainly cultivated through submerged fermentation, as it is very impractical to grow this rare and wild mushroom in traditional methods. Recent years has seen a boom in the artificial cultivation of the mushroom in China (Du et al., 2014). Some of the polysaccharides extracted from the fluid medium of M. esculenta are potentially bioactive (Wasser, 2002;Lei et al., 2016). ...
This paper aims to disclose the hyperglycaemic activity of the polysaccharides extracted from Morchella esculenta (M. esculenta) mycelia. To this end, M. esculenta mycelia were cultured submerged fermentation, and the acidic polysaccharides were extracted and purified by anion exchange chromatography gel permeation chromatography. Then, a group of healthy rats was prepared, together with several groups of diabetes mellitus (DM) rats intervened with different doses of M. esculenta acidic polysaccharides. Next, several Morris water maze (MWM) tests were performed to reveal the regulation effect of these polysaccharides on the mRNA expression of the nerve growth factor (NGF) of the rats. The main results are as follows. The fasting blood glucose (FBG) and serum insulin (INS) of Group D (intervened by 600 μg∙mL⁻¹ of polysaccharide) plunged to 8.12mmol/L and 31.67mU/L, respectively, compared to those of the control group (P<0.01), indicating that the polysaccharides can promote INS secretion. Meanwhile, the relative mRNA expression level of Group D grew by from the level of the control group (p<0.01). Thus, the acidic M. esculenta polysaccharide must have triggered the expression of NGF protein. The research findings lay a solid basis for the development of anti-hyperglycaemia food based on M. esculenta polysaccharides.
... The edible, medicinal Morchella fungi (morels), mainly M. importuna and M. sextelata, have been commercially cultivated in fields in China since 2012 (Du et al. 2014;He et al. 2015). In the 2015-2016 production season, the total cultivation area exceeded 1200 ha in more than 15 provinces, municipalities and autonomous regions of China. ...
... Morchella fungi fructify at low temperature (6-16 °C) and moderate relative humidity (around 85%) (Du et al. 2014). D. longispora might be able to switch from endophytic to pathogenic under favorable conditions, particularly at temperatures above 25 °C and high relative humidity (> 90%), thereby causing the observed malformations on Morchella fruitbodies. ...
In March 2013, pileus rot disease was first observed on cultivated Morchella importuna. Infected ascomata were covered by white, velvety mycelia mainly on the pileus, and the infection resulted in malformed fruiting bodies. The causal pathogen was identified as Diploöspora longispora based on its morphology and the internal transcribed spacer of its ribosomal DNA sequences. After inoculation of young ascomata with the isolates, the original symptoms were reproduced, and the same pathogen was reisolated from diseased ascomata. This is the first report of pileus rot disease on morels caused by D. longisporain China.
