Bioscience Evidence 2026, Vol.16, No.4, 316-328 http://bioscipublisher.com/index.php/be 319 Figure 1 Improvement of the nutritional composition of Lentinula edodes using different cultivation substrates (Adapted from Desisa et al., 2024) The utilization of substrate nutrients also changes during different growth stages of shiitake mushrooms. During the mycelial colonization stage, shiitake preferentially utilizes relatively easily degradable components such as hemicellulose. After entering primordium formation and fruiting body development stages, the fungus further utilizes more complex components such as cellulose and lignin (Atila, 2019). 2.5 Moisture content and physical structure of substrates For family farms and small-scale producers, a good substrate should not only provide sufficient nutrients but also create suitable environmental conditions for mycelial growth. Mycelia require adequate moisture, oxygen supply, and appropriate physical structure for effective colonization. In practice, different particle sizes of sawdust are often mixed to ensure that the substrate maintains both water retention capacity and good aeration. Substrate particle size, pore structure, and raw material composition jointly influence mycelial expansion, nutrient utilization, and final production performance, indicating that physical structure is an important factor in substrate design rather than a secondary consideration (Nabhan et al., 2025). The moisture content of substrates should be adjusted according to the characteristics of raw materials. Desisa et al. (2023) controlled the moisture content of agricultural waste-based shiitake substrates at 60%–65% and added gypsum and calcium carbonate to improve substrate stability. Gypsum provides minerals such as calcium and sulfur while improving substrate structure, whereas calcium carbonate helps regulate substrate pH conditions. Different agricultural residues show considerable differences in water retention, air permeability, and mycelial colonization ability due to their structural characteristics. Baktemur et al. (2022) compared several agricultural by-products as shiitake substrate materials and found significant differences in mycelial growth rates among different formulations, with large variation between the fastest and slowest treatments. These differences were not
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