Morels are one of the most highly prized edible and medicinal mushrooms worldwide. Therefore, historically, there has been a large international interest in their cultivation. Numerous ecological, physiological, genetic, taxonomic, and mycochemical studies have been previously developed. At the beginning of this century, China finally achieved artificial cultivation and started a high-scale commercial development in 2012. Due to its international interest, its cultivation scale and area expanded rapidly in this country. However, along with the massive industrial scale, a number of challenges, including the maintenance of steady economic profits, arise. In order to contribute to the solution of these challenges, formal research studying selection, species recognition, strain aging, mating type structure, life cycle, nutrient metabolism, growth and development, and multi-omics has recently been boosted. This paper focuses on discussing current morel cultivation technologies, the industrial status of cultivation in China, and the relevance of basic biological research, including, e.g., the study of strain characteristics, species breeding, mating type structure, and microbial interactions. The main challenges related to the morel cultivation industry on a large scale are also analyzed. It is expected that this review will promote a steady global development of the morel industry based on permanent and robust basic scientific knowledge.

Morels, which belong to the Ascomycete genus Morchella, are highly valued edible fungi treasured by gourmet chefs worldwide. Some species are saprotrophic and others are able to form facultative mycorrhizal-like associations with plant roots without establishing true ectomycorrhizal symbioses. In general, it is considered that the formation of asexual spores, or mitospores, is an important step in the life cycle of morels. However, ultrastructure characterization and physiological attributes of morel mitospores have received little attention. In this contribution, the mitospores of M. sextelata were successfully induced under laboratory conditions and their ultrastructure, occurrence, germination, physiological characteristics and mating type gene structure were studied. Mitospore production was closely related to aeration, nutrition and humidity conditions. The average germination rate of mitospores on different media and under various induction stimuli was very low, with an average of 1/100,000. Based on the ultrastructure characterization, low germination rate, growth rate decline, rapid aging and mating genotyping, it was concluded that the mitospores of M. sextelata had lost their conventional function as conidia and might act more as mate sperm-like (gamete) structures. Thus, this study contributed to a deeper understanding of the life cycle of the economically and ecologically important morel fungal group.

Continuous cropping obstacle has been becoming the bottleneck for the stable development of morel cultivation. The allelopathic effect of soil allelochemicals may play an instrumental role in the morel soil sickness. In this study, the allelochemicals were identified by gas chromatography-mass spectrometry (GC-MS) combined with in vitro bioassay. A total of 61 chemical substances were identified through the GC-MS analysis of 12 replanting and control soil samples, comprising 10 phenolic acids, 36 acids, 3 aldehydes, etc. Among which, 15 compounds with values of variable importance for the projection (VIP) greater than 1 in the orthogonal partial least squares-discriminant analysis (OPLS-DA) were selected as the differential metabolites between soil samples of continuous cropping and control. The bioassay showed that 4-coumaric acid and vanillic acid exhibited inhibitory effect on mycelial growth of three cultivated Morchella mushrooms (M. sextelata, M. eximia and M. importuna) under soil native concentrations. Analysis of potential biosynthetic pathways of phenolic acids found that the 3 cultivable Morchella mushrooms are unable to synthesize phenolic acid allelochemicals. Therefore, although they were detected in trace amounts in static culture broth of M. sextelata, the two phenolic acids can only be defined as morel allelochemicals rather than autotoxins. Taken together, we found two morel allelochemicals that may derive from morel related microbes and plants, which will be helpful for further fundamental study and application in morel artificial cultivation.