Science and technology breakthroughs to advance artificial cultivation of true morels

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Systematic understanding of Morchella and the ecological environment
Many factors affected the growth of Morchella, including nutrient substances, soil, plants, etc. (Liu et al., 2018;Papadaki et al., 2019;Evangelista et al., 2021;Tan et al., 2021a;Xu et al., 2022a).Therefore, for the upgrading of morel cultivation technology, we should consider it from the perspective of eco-environment integration, and thinking from a single factor should be avoided.A comprehensive evaluation of the impacts of water, air, and materials on the entire process of morel cultivation is necessary.Other than the seven elata clade species that are currently cultivated (Du, 2019;Wu et al., 2019), it is important to systematically screen all Morchella species to exploit the stress resistant resources in Morchella.Keeping on collecting wild Morchella resources and expanding the germplasm resource bank through the discovery of new species and strains is important.Many other factors should be taken into account, such as morphology and biogeography, DNA barcoding, molecular systematics, global diversity, etc.In summary, to achieve breakthroughs in morel cultivation technology, we should first establish a systematic understanding of Morchella and the ecological environment.

Microbiome analysis will be contributed to understanding the cultivation process of Morchella
It has been reported that soil microbiome has an important influence on morel cultivation and the α-diversity level and community evenness among soil fungal taxa could affect the production of morels (Tan et al., 2021b;Yu et al., 2022;Zhang et al., 2023).However, the changes in microbial communities during field cultivation and their effects are still not well understood.The microbial community is very sensitive to the interference of abiotic environmental factors, including soil tillage, temperature, heavy metals, and so on (Hu et al., 2022;Nelson et al., 2022).Currently, the changes in the microbiome in morel cultivation soil after the intervention of abiotic factors are little known.
With the deepening of microbiome research (Benucci et al., 2019;Longley et al., 2019), a database of microorganisms related to morel cultivation should be established to better understand the interaction between Morchella and the environmental microbiome.At the same time, the influence of the microbial community on the life history of Morchella can also be explored, and the spawn production process of Morchella can be deeply analyzed (Yu et al., 2023).By researching the functions of various microorganisms in the soil microbial community, some beneficial microorganisms can be found with the potential to create a more productive and sustainable morel cultivation system, which can promote the growth of Morchella, improve the use efficiency of exogenous nutrients, and enhance the resistance of Morchella to environmental stress and pathogenic microorganisms.The establishment of a platform for data collection and communication will be helpful for improving the management of morel cultivation At present, sensor technology has been widely used in the field of agriculture (Singh et al., 2020;Arrubla-Hoyos et al., 2022).The development of big data, artificial intelligence, and other technologies provides more rapid collection, analysis, storage, sharing, and integration of data (Jung et al., 2021;Xu et al., 2022b).The large-scale cultivation of Morchella has been ongoing for more than 10 years in China and many laboratories and companies have collected a large amount of data related to morel cultivation (Guo et al., 2020;Tan et al., 2021c).However, there is a lack of effective tools to analyze and use the existing data.The progress of data science and information technology can greatly improve the ability to solve complex problems and provide an important opportunity for improving the management technology of morel cultivation.Therefore, the establishment of a network platform that can store and open access morel cultivation data sets will be helpful for sharing data.Making full use of the latest information technology to accelerate the unification and standardization of morel cultivation technology is the key to improving the management of morel cultivation.

Focus on basic research and embrace new breeding techniques
In recent years, basic research on Morchella has made some progress in terms of life cycle, sclerotium development mechanisms, exogenous nutrition utilization, characteristic of mitochondrial genome, etc. (Tan et al., 2019;Liu et al., 2020Liu et al., , 2021a;;Du and Yang, 2021;Chai et al., 2022).However, there is still a lack of understanding of conidial production, working mechanisms of exogenous nutrition bags, primordium formation, and fruiting body development mechanisms (Liu et al., 2022), so basic research should be continued.
With the development of gene editing technology (Wang et al., 2019), improvement of Morchella strains by precise breeding is expected to be realized in the next few years.Advances in basic research combining with precise breeding methods can help to accurately and rapidly improve the biological traits that are important for the yield and quality of Morchella.This technology opens the door to the rapid development of superior Morchella strains.
New equipments and materials will be the new driving force for the development of the Morchella industry How to improve the labor efficiency of morel cultivation is also a problem that needs to be addressed.In recent years, automatic external nutrient bag packing machines, automatic strain bag breaking machines, and other small equipments have been applied in morel cultivation.On the other hand, the development of automatic morel fruiting body picking equipment and packaging equipment is of great significance for reducing labor consumption and transportation loss.New equipment-related research and development should be paid more attention.
Exogenous nutrition bags consume a lot of labor and may bring environmental pollution (Liu et al., 2017).Recent experiments have found that filter paper coated with exogenous nutrients can play the same role as polypropylene bags (Liu et al., 2021b), suggesting that it is possible for environmentally friendly materials to replace polypropylene plastics.Based on the basic principle of exogenous nutrient bags delivering nutrients to soil (Tan et al., 2019), more materials should be tried to replace plastics or make more efficient use of exogenous nutrients and reduce environmental pressure.

Conclusions and future outlook
In the next 10 years, we will seek scientific and technological breakthroughs in morel cultivation in five directions, including the microbiome, data science, accurate dynamic perception, new equipments and materials, and gene editing breeding (Figure 1).These are also key technologies that are indispensable in the morels field.It is expected that the scale of morel cultivation will continue to expand, not only in field cultivation but also in the scale of industrial cultivation.Advances in science and technology will lay the foundation for the development of the morels industry.

FIGURE
FIGUREFive directions worth e orts in the development of Morchella cultivation.