15The balance and interplay between sexual and asexual reproduction is one of the most 16 attractive mysteries in fungi. The choice of developmental strategy reflects the ability of 17 fungi to adapt to the changing environment. However, the evolution of developmental paths 18 and the metabolic regulation during differentiation and morphogenesis are poorly understood. 19 Here, we monitor the carbohydrate metabolism and gene expression regulation during the 20 early differentiation process from the "fungal stem cell", vegetative mycelium, to the highly 21 differentiated tissue/cells, fruiting body, oidia or sclerotia, of a homokaryotic fruiting 22 Coprinopsis cinerea strain A43mut B43mut pab1-1 #326, uncovering the systematic changes 23 during morphogenesis and the evolutionary process of developmental strategies. Conversion 24 between glucose and glycogen and conversion between glucose and beta-glucan are the main 25 carbon flows in the differentiation processes. Genes related to carbohydrate transport and 26 metabolism are significantly differentially expressed among paths. RNA editing, a novel 27 layer of gene expression regulation, occurs in all four developmental paths and enriched in 28 cytoskeleton and carbohydrate metabolic genes. It is developmentally regulated and 29 evolutionarily conserved in basidiomycetes. Evolutionary transcriptomic analysis on four 30 developmental paths showed that all transcriptomes are under purifying selection, and the 31 more stressful the environment, the younger the transcriptome age. Oidiation has the lowest 32 value of transcriptome age index (TAI) and transcriptome divergence index (TDI), while 33 fruiting process has the highest of both indexes. These findings provide new insight to the 34 regulations of carbon metabolism and gene expressions during fungal developmental paths 35 differentiation. 36 37 3 Importance 38 Fungi is a group of species with high diversity and plays essential roles to the ecosystem. The 39 life cycle of fungi is complex in structure and delicate in function. Choice of developmental 40 strategies and internal changes within the organism are both important for the fungus to fulfill 41 their ecological functions, reflecting the relationship between environment and the 42 population. This study put the developmental process of vegetative growth, sexual and 43 asexual reproduction, resistant structure formation of a classical model basidiomycetes 44 fungus, C. cinerea, together for the first time to view the developmental paths differentiation 45 process with physiology, transcriptomics and evolutionary prospects. Carbohydrate assays 46 and RNA-seq showed the changes of the fungus. Our results fill the gaps on gene expression 47 regulation during the early stage of developmental paths differentiation, and expand our 48 understanding of the evolutionary process of life history and reproductive strategy in fungi. 49 Keywords 50 Developmental paths, carbon metabolic flux, transcriptome, RNA editing, 51 phylotranscriptomic 52 53 4 Introduction 54 Th...