Lin Yang
Current Organizations / Institutes
The organizations/institutes from the scholar's latest article
Lin Yang has a total of 21 co-authors. Here are the top co-authors by number of shared publications:
- Yan Zhao
Institute of Edible Fungi, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China
7 shared publications - Changxia Yu
Institute of Edible Fungi, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China
6 shared publications - Qin Dong
Institute of Edible Fungi, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China
6 shared publications - Lei Zha
Institute of Edible Fungi, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China
5 shared publications - Qian Guo
Institute of Edible Fungi, Shanghai Academy of Agricultural Sciences, Shanghai 201403, China
4 shared publications - Pattana Kakumyan
School of Science, Mae Fah Luang University, Chiang Rai, Thailand
2 shared publications
Latest Publications
- Open Access
- 1 November 2026
- Published by MDPI AG in Microorganisms
- Vol. 14 (11) , 2484
Effects of a Composite Additive on Yield, Microbial Community, and Carbon–Nitrogen Transformation During Volvariella volvacea Cultivation on Spent Pleurotus eryngii Substrate
The addition of nutritional additives to the cultivation substrate is an effective way to increase edible fungus yields; however, the mechanisms by which these additives regulate the structure and function of the substrate microbial community to promote nutrient transformation remain unclear. In this study, a patented composite additive composed of soybean meal, corn protein, and wheat bran was added to a Volvariella volvacea cultivation system based on spent Pleurotus eryngii substrate after secondary fermentation, and the additive-treated (T) and control (ck) groups were compared across different cultivation stages. The additive increased V. volvacea yield by 26.4% (4.02 vs. 3.18 kg/m2) and increased fruiting body amino acid content by 16.43%, in addition to shifting carbon and nitrogen metabolism toward amino acid accumulation and biomass. Throughout cultivation, the additive sustained higher cellulase (CL) and alkaline protease (AKP) activities, increased cellulose degradation and organic nitrogen mineralization, and enlarged the pool of available carbon and nitrogen in the substrate. Co-abundance group (CAG) network analysis of 626 ASVs (577 bacterial and 49 fungal ASVs) revealed that the additive was associated with three potential bacterial–fungal functional modules in a stage-specific manner: a cellulose-degrading and nitrogen-accumulating module, a carbon-assimilating module, and a nitrogen-mineralizing module. The core taxa potentially associated with carbon and nitrogen transformation included the bacteria Pseudoxanthomonas and Thermopolyspora and the fungus Mycothermus, together with the additive-enriched genera Schlegelella, Niabella, and Chelatococcus. ASV-level resolution further revealed an intragenus functional division of Mycothermus, in which two ASV clusters were associated with cellulose degradation and nitrogen mineralization. These findings elucidate the mechanism whereby additives enrich functional microbial guilds, accelerate coupled carbon–nitrogen turnover, and ultimately increase the yield and quality of V. volvacea, providing a theoretical basis for the precise management of spent-substrate-based edible fungus cultivation.
- 1 November 2026
- Published by MDPI AG in Microorganisms
- Vol. 14 (11) , 2484
Open AccessEffects of a Composite Additive on Yield, Microbial Community, and Carbon–Nitrogen Transformation During Volvariella volvacea Cultivation on Spent Pleurotus eryngii Substrate
The addition of nutritional additives to the cultivation substrate is an effective way to increase edible fungus yields; however, the mechanisms by which these additives regulate the structure and function of the substrate microbial community to promote nutrient transformation remain unclear. In this study, a patented composite additive composed of soybean meal, corn protein, and wheat bran was added to a Volvariella volvacea cultivation system based on spent Pleurotus eryngii substrate after secondary fermentation, and the additive-treated (T) and control (ck) groups were compared across different cultivation stages. The additive increased V. volvacea yield by 26.4% (4.02 vs. 3.18 kg/m2) and increased fruiting body amino acid content by 16.43%, in addition to shifting carbon and nitrogen metabolism toward amino acid accumulation and biomass. Throughout cultivation, the additive sustained higher cellulase (CL) and alkaline protease (AKP) activities, increased cellulose degradation and organic nitrogen mineralization, and enlarged the pool of available carbon and nitrogen in the substrate. Co-abundance group (CAG) network analysis of 626 ASVs (577 bacterial and 49 fungal ASVs) revealed that the additive was associated with three potential bacterial–fungal functional modules in a stage-specific manner: a cellulose-degrading and nitrogen-accumulating module, a carbon-assimilating module, and a nitrogen-mineralizing module. The core taxa potentially associated with carbon and nitrogen transformation included the bacteria Pseudoxanthomonas and Thermopolyspora and the fungus Mycothermus, together with the additive-enriched genera Schlegelella, Niabella, and Chelatococcus. ASV-level resolution further revealed an intragenus functional division of Mycothermus, in which two ASV clusters were associated with cellulose degradation and nitrogen mineralization. These findings elucidate the mechanism whereby additives enrich functional microbial guilds, accelerate coupled carbon–nitrogen turnover, and ultimately increase the yield and quality of V. volvacea, providing a theoretical basis for the precise management of spent-substrate-based edible fungus cultivation.
- Open Access
- 28 April 2026
- Published by MDPI AG in Microorganisms
- Vol. 14 (5) , 985
Spent Mushroom Substrate Reused as Organic Fertilizer Enhances Lettuce (Lactuca sativa L.) Quality and Soil Nutrients: Insights from Physicochemical and Microbiome Analyses
Returning spent mushroom substrate (SMS) to the field is an effective way to dispose of it. However, given the substantial nutrient consumption associated with Volvariella volvacea SMS, their effects on soil properties and crop performance warrant further investigation. By analyzing the effects of three different application rates of SMS on soil nutrients and lettuce (Lactuca sativa L.) quality, the results showed that the group with 1.5 kg m−2 SMS addition improved the total nitrogen (+21.2%), and organic content (+27.9%) in soil, and it demonstrated particularly outstanding performance in enhancing the survival rate (+21.9%), average weight (+71.7%), chlorophyll content (+45.6%), and total phenolic content (+25.2%) of lettuce. By comparing the soil microbial communities in the control group, the SMS (1.5 kg m−2) treatment group, and the organic fertilizer treatment group, it was found that they were mainly composed of Group S1, S2, and S3 microorganisms, respectively. The microbial community evenness in the treatment groups was greater than that in the control group. Furthermore, the results also revealed that the microbial conversion efficiency of nitrogen and phosphorus in the SMS treatment group was higher than the control group, which promoted nutrient cycling and improved the quality of lettuce. Our analysis provides an environmentally friendly way for Volvariella volvacea SMS disposal.
- 28 April 2026
- Published by MDPI AG in Microorganisms
- Vol. 14 (5) , 985
Open AccessSpent Mushroom Substrate Reused as Organic Fertilizer Enhances Lettuce (Lactuca sativa L.) Quality and Soil Nutrients: Insights from Physicochemical and Microbiome Analyses
Returning spent mushroom substrate (SMS) to the field is an effective way to dispose of it. However, given the substantial nutrient consumption associated with Volvariella volvacea SMS, their effects on soil properties and crop performance warrant further investigation. By analyzing the effects of three different application rates of SMS on soil nutrients and lettuce (Lactuca sativa L.) quality, the results showed that the group with 1.5 kg m−2 SMS addition improved the total nitrogen (+21.2%), and organic content (+27.9%) in soil, and it demonstrated particularly outstanding performance in enhancing the survival rate (+21.9%), average weight (+71.7%), chlorophyll content (+45.6%), and total phenolic content (+25.2%) of lettuce. By comparing the soil microbial communities in the control group, the SMS (1.5 kg m−2) treatment group, and the organic fertilizer treatment group, it was found that they were mainly composed of Group S1, S2, and S3 microorganisms, respectively. The microbial community evenness in the treatment groups was greater than that in the control group. Furthermore, the results also revealed that the microbial conversion efficiency of nitrogen and phosphorus in the SMS treatment group was higher than the control group, which promoted nutrient cycling and improved the quality of lettuce. Our analysis provides an environmentally friendly way for Volvariella volvacea SMS disposal.
- Open Access
- 28 November 2025
- Published by MDPI AG in Journal of Fungi
- Vol. 11 (12) , 845
Comparative Analysis of Volatile Aroma Compounds, Fatty Acids, and LOX Pathway Gene Expression of Two Lentinula edodes Mycelia
Although the aroma profile of Lentinula edodes has been extensively studied in fruiting bodies, the mycelial stage provides a distinct context for elucidating the fundamental metabolic pathways, free from the complexities of organismal development. To elucidate the mechanism underlying aroma differences between L. edodes strain 808 (the control strain) and its mutant strain ww808 (with almost no shiitake aroma), this study employed GC-IMS combined with PCA and OPLS-DA to identify key aroma biomarkers during the mycelial stage. All analyses were performed with three biological replicates. Furthermore, fatty acids composition, key enzyme activities of the LOX pathway, and their gene expression levels were systematically compared. The results indicated significant differences in the content of volatile aroma compounds in the mycelia of the two strains, primarily stemming from fundamental restructuring of gene expression and enzyme activity in the LOX pathway. The LOX gene expression and LOX activity of 808 mycelium were relatively high, facilitating the accumulation of key aroma compounds such as phenylethanal, benzaldehyde, and ethyl acetate, which constitute its distinctive aromatic profile. However, although the mycelium of ww808 possessed richer fatty acid precursor (C18:2), its lower LOX gene expression restricted the flux of this pathway. The significantly increased expression of ADH2, ADH3, and ADH5 genes and higher ADH activity enhanced the conversion capacity of aldehydes to alcohols and ketones. Given the generally higher odor thresholds of alcohols and ketones compared to aldehydes, distinct aroma profiles emerged between the two strains. Pearson correlation analysis further confirmed the significant correlations between the aroma biomarkers, fatty acids, key genes, and enzyme activities. This study revealed the formation mechanism of aroma differences in the mycelia of the two strains from the perspective of metabolic pathways, providing a theoretical foundation and candidate targets for the directed genetic improvement of L. edodes aroma quality.
- 28 November 2025
- Published by MDPI AG in Journal of Fungi
- Vol. 11 (12) , 845
Open AccessComparative Analysis of Volatile Aroma Compounds, Fatty Acids, and LOX Pathway Gene Expression of Two Lentinula edodes Mycelia
Although the aroma profile of Lentinula edodes has been extensively studied in fruiting bodies, the mycelial stage provides a distinct context for elucidating the fundamental metabolic pathways, free from the complexities of organismal development. To elucidate the mechanism underlying aroma differences between L. edodes strain 808 (the control strain) and its mutant strain ww808 (with almost no shiitake aroma), this study employed GC-IMS combined with PCA and OPLS-DA to identify key aroma biomarkers during the mycelial stage. All analyses were performed with three biological replicates. Furthermore, fatty acids composition, key enzyme activities of the LOX pathway, and their gene expression levels were systematically compared. The results indicated significant differences in the content of volatile aroma compounds in the mycelia of the two strains, primarily stemming from fundamental restructuring of gene expression and enzyme activity in the LOX pathway. The LOX gene expression and LOX activity of 808 mycelium were relatively high, facilitating the accumulation of key aroma compounds such as phenylethanal, benzaldehyde, and ethyl acetate, which constitute its distinctive aromatic profile. However, although the mycelium of ww808 possessed richer fatty acid precursor (C18:2), its lower LOX gene expression restricted the flux of this pathway. The significantly increased expression of ADH2, ADH3, and ADH5 genes and higher ADH activity enhanced the conversion capacity of aldehydes to alcohols and ketones. Given the generally higher odor thresholds of alcohols and ketones compared to aldehydes, distinct aroma profiles emerged between the two strains. Pearson correlation analysis further confirmed the significant correlations between the aroma biomarkers, fatty acids, key genes, and enzyme activities. This study revealed the formation mechanism of aroma differences in the mycelia of the two strains from the perspective of metabolic pathways, providing a theoretical foundation and candidate targets for the directed genetic improvement of L. edodes aroma quality.
- Open Access
- 12 November 2025
- Published by MDPI AG in Horticulturae
- Vol. 11 (11) , 1361
Effects of Cadmium Stress on Mycelial Growth and Antioxidant Systems in Agaricus subrufescens Peck
Agaricus subrufescens Peck is a nutrient-rich edible fungi with a distinctive flavor, but most varieties are sensitive to cadmium (Cd), making cadmium contamination common during cultivation. Currently, excessive fertilizer uses and increased solid waste are exacerbating cadmium contamination in soils. Since A. subrufescens utilize agricultural residues like straw and livestock manure as cultivation substrates, Cd can be adsorbed readily, leading to secondary accumulation. In this study, the toxic effects of and response mechanisms to different Cd concentrations with respect to mycelial growth, heavy metal accumulation, and antioxidant systems of A. subrufescens were systematically investigated. The results indicated that the mycelia exhibited Cd accumulation capacity, with accumulation levels positively correlated with stress concentration. At a Cd concentration of 5 mg/L, the intracellular Cd concentration in the mycelia reached approximately 800 mg/kg. As the Cd concentration increased, the efficiency of Cd uptake by mycelia correspondingly decreased. Cadmium stress (≥0.5 mg/L) significantly inhibited mycelial growth and induced morphological abnormalities, with the mycelia exhibiting yellowing. Furthermore, Cd induced dose-dependent oxidative stress. Hydrogen peroxide and MDA levels peaked at a Cd concentration of 2 mg/L, reaching 2.26 μmol/g and 8.98 nmol/g, respectively, indicating heightened lipid peroxidation. Low concentrations of Cd (≤2 mg/L) promoted increases in ASA and GSH activity. SOD, POD, GR, and APX activities significantly increased, with the ASA-GSH cycle synergistically scavenging ROS. CAT activity remained persistently inhibited, APX/GR activity was suppressed, and total sugar metabolism was disrupted, leading to the collapse of antioxidant defenses. In summary, depending on the Cd concentration, A. subrufescens mycelia exhibit markedly different responses at low versus high concentrations. This study provides a foundation for further research into the application of edible fungi in heavy metal-resistant cultivation.
- 12 November 2025
- Published by MDPI AG in Horticulturae
- Vol. 11 (11) , 1361
Open AccessEffects of Cadmium Stress on Mycelial Growth and Antioxidant Systems in Agaricus subrufescens Peck
Agaricus subrufescens Peck is a nutrient-rich edible fungi with a distinctive flavor, but most varieties are sensitive to cadmium (Cd), making cadmium contamination common during cultivation. Currently, excessive fertilizer uses and increased solid waste are exacerbating cadmium contamination in soils. Since A. subrufescens utilize agricultural residues like straw and livestock manure as cultivation substrates, Cd can be adsorbed readily, leading to secondary accumulation. In this study, the toxic effects of and response mechanisms to different Cd concentrations with respect to mycelial growth, heavy metal accumulation, and antioxidant systems of A. subrufescens were systematically investigated. The results indicated that the mycelia exhibited Cd accumulation capacity, with accumulation levels positively correlated with stress concentration. At a Cd concentration of 5 mg/L, the intracellular Cd concentration in the mycelia reached approximately 800 mg/kg. As the Cd concentration increased, the efficiency of Cd uptake by mycelia correspondingly decreased. Cadmium stress (≥0.5 mg/L) significantly inhibited mycelial growth and induced morphological abnormalities, with the mycelia exhibiting yellowing. Furthermore, Cd induced dose-dependent oxidative stress. Hydrogen peroxide and MDA levels peaked at a Cd concentration of 2 mg/L, reaching 2.26 μmol/g and 8.98 nmol/g, respectively, indicating heightened lipid peroxidation. Low concentrations of Cd (≤2 mg/L) promoted increases in ASA and GSH activity. SOD, POD, GR, and APX activities significantly increased, with the ASA-GSH cycle synergistically scavenging ROS. CAT activity remained persistently inhibited, APX/GR activity was suppressed, and total sugar metabolism was disrupted, leading to the collapse of antioxidant defenses. In summary, depending on the Cd concentration, A. subrufescens mycelia exhibit markedly different responses at low versus high concentrations. This study provides a foundation for further research into the application of edible fungi in heavy metal-resistant cultivation.
- Open Access
- 15 October 2025
- Published by MDPI AG in Microorganisms
- Vol. 13 (10) , 2372
Sustainable Recycling of Mushroom Residue as an Effective Substitute for Cotton Hull Waste in Volvariella volvacea Cultivation: Evidence from Physicochemical and Microbiome Analyses
Mushroom residue (MR) is extensively produced during the industrialized cultivation of mushrooms, and its utilization is environmentally sustainable. Cotton hull waste (CW) serves as a common raw material for the cultivation of Volvariella volvacea in China. This study compared MR- and CW-based cultivation formulas with respect to their physicochemical characteristics, bacterial communities, and functional dynamics during substrate fermentation (composting). Xylanase production was greater in the MR formula than in the CW formula. Conversely, cellulase (CMCase) was generated at higher levels in the CW formula compared to the MR formula. Interestingly, the biological efficiency of MR was found to be comparable to that of CW, but the cost of MR was much lower. The dynamics of bacterial communities and their associated metabolic functions during substrate fermentation were monitored using 16S rRNA metagenomics techniques. Significant alterations in bacterial community structure were observed within both formulas throughout the preparation phase. Indicator species analysis revealed distinct patterns of bacterial diversity development between MR- and CW-based composts during fermentation. Metabolic function analysis indicated that carbohydrate and amino acid metabolism remained relatively active throughout this process. These results suggest that the MR formula is equally effective as conventional CW compost for supporting V. volvacea cultivation, while also offering a lower raw material cost.
- 15 October 2025
- Published by MDPI AG in Microorganisms
- Vol. 13 (10) , 2372
Open AccessSustainable Recycling of Mushroom Residue as an Effective Substitute for Cotton Hull Waste in Volvariella volvacea Cultivation: Evidence from Physicochemical and Microbiome Analyses
Mushroom residue (MR) is extensively produced during the industrialized cultivation of mushrooms, and its utilization is environmentally sustainable. Cotton hull waste (CW) serves as a common raw material for the cultivation of Volvariella volvacea in China. This study compared MR- and CW-based cultivation formulas with respect to their physicochemical characteristics, bacterial communities, and functional dynamics during substrate fermentation (composting). Xylanase production was greater in the MR formula than in the CW formula. Conversely, cellulase (CMCase) was generated at higher levels in the CW formula compared to the MR formula. Interestingly, the biological efficiency of MR was found to be comparable to that of CW, but the cost of MR was much lower. The dynamics of bacterial communities and their associated metabolic functions during substrate fermentation were monitored using 16S rRNA metagenomics techniques. Significant alterations in bacterial community structure were observed within both formulas throughout the preparation phase. Indicator species analysis revealed distinct patterns of bacterial diversity development between MR- and CW-based composts during fermentation. Metabolic function analysis indicated that carbohydrate and amino acid metabolism remained relatively active throughout this process. These results suggest that the MR formula is equally effective as conventional CW compost for supporting V. volvacea cultivation, while also offering a lower raw material cost.
- Open Access
- 24 June 2025
- Published by MDPI AG in Journal of Fungi
- Vol. 11 (7) , 479
Dynamics of Nutrient Components and Microbial Communities in Substrates During the Development of the Fruiting Bodies of Volvariella volvacea
Cotton waste, a growth medium for Volvariella volvacea, has significant commercial and nutritional value. Under controlled environmental conditions, substrate nutrient composition and microorganisms affect the growth of V. volvacea. In this study, the changes in the nutrient content of the substrate at different stages of fruiting body development were compared based on an 86% waste cotton substrate, and microbial diversity was studied via 16S rRNA analysis. The results indicated that there were significant differences in nutrient content in the substrate at different stages of fruiting body development. The total contents of carbon, nitrogen, and phosphorus initially increased but then decreased due to nutrient absorption and utilization by V. volvacea. It was also found that large amounts of organic nitrogen decomposed into more readily utilizable inorganic nitrogen. The nutritional content and microbial community structure of the substrate during the egg stage significantly differed from those during the other four stages, making the egg stage the most critical period in cultivation. Through correlation analysis between nutrient content and microbial differences, it was found that differential microbial taxa (Beijerinckiaceae, Burkholderiales, Chitinophaga jiangningensis, etc.) with nitrogen fixation, denitrification, and cellulose decomposition functions were significantly related to carbon- and nitrogen-related indicators such as nitrate nitrogen, microbial biomass carbon, and alkali-hydrolyzed nitrogen. These microorganisms play important roles in determining the variation in the nutritional profile of the substrate. This study provides a theoretical basis for promoting the absorption and utilization of nutrients by V. volvacea by altering the structure of the microbial community of the growth substrate.
- 24 June 2025
- Published by MDPI AG in Journal of Fungi
- Vol. 11 (7) , 479
Open AccessDynamics of Nutrient Components and Microbial Communities in Substrates During the Development of the Fruiting Bodies of Volvariella volvacea
Cotton waste, a growth medium for Volvariella volvacea, has significant commercial and nutritional value. Under controlled environmental conditions, substrate nutrient composition and microorganisms affect the growth of V. volvacea. In this study, the changes in the nutrient content of the substrate at different stages of fruiting body development were compared based on an 86% waste cotton substrate, and microbial diversity was studied via 16S rRNA analysis. The results indicated that there were significant differences in nutrient content in the substrate at different stages of fruiting body development. The total contents of carbon, nitrogen, and phosphorus initially increased but then decreased due to nutrient absorption and utilization by V. volvacea. It was also found that large amounts of organic nitrogen decomposed into more readily utilizable inorganic nitrogen. The nutritional content and microbial community structure of the substrate during the egg stage significantly differed from those during the other four stages, making the egg stage the most critical period in cultivation. Through correlation analysis between nutrient content and microbial differences, it was found that differential microbial taxa (Beijerinckiaceae, Burkholderiales, Chitinophaga jiangningensis, etc.) with nitrogen fixation, denitrification, and cellulose decomposition functions were significantly related to carbon- and nitrogen-related indicators such as nitrate nitrogen, microbial biomass carbon, and alkali-hydrolyzed nitrogen. These microorganisms play important roles in determining the variation in the nutritional profile of the substrate. This study provides a theoretical basis for promoting the absorption and utilization of nutrients by V. volvacea by altering the structure of the microbial community of the growth substrate.
- Open Access
- 17 February 2025
- Published by MDPI AG in Microorganisms
- Vol. 13 (2) , 437
Comparison of the Bacterial and Fungal Communities and Metabolic Functions of Cottonseed Hull Waste Compost Associated with High and Low Yields of Straw Mushroom Volvariella volvacea
Volvariella volvacea was grown on cottonseed hull waste compost and divided into high-yield (HBE) and low-yield (LBE) conditions. Gene sequencing was used to examine bacterial and fungal populations during cottonseed husk waste composting. At the end of fermentation, the dominant bacterial genera in the HBE compost were Chelatococcus and Thermobacillus, while Symbiobacterium and Acinetobacter were more abundant in the LBE compost. Ascomycota and Basidiomycota dominated all the composting phases. The Ascomycota genera Colletotrichum, Pichia, Mycothermus, and Thermomyces dominated in phase II of HBE composting. The LBE compost had higher abundances of the Basidiomycota genera Cystofilobasidium and Cryptococcus than the HBE compost. The predicted pathotroph and saprotroph-symbiotroph abundances were more positively linked to HBE composting phase II than to LBE composting. High-biological-efficiency microbial communities are characterized by high pH, carbon, and nitrogen levels. Changes in physiochemical traits, microbial diversity, and metabolism affect the V. volvacea yield.
- 17 February 2025
- Published by MDPI AG in Microorganisms
- Vol. 13 (2) , 437
Open AccessComparison of the Bacterial and Fungal Communities and Metabolic Functions of Cottonseed Hull Waste Compost Associated with High and Low Yields of Straw Mushroom Volvariella volvacea
Volvariella volvacea was grown on cottonseed hull waste compost and divided into high-yield (HBE) and low-yield (LBE) conditions. Gene sequencing was used to examine bacterial and fungal populations during cottonseed husk waste composting. At the end of fermentation, the dominant bacterial genera in the HBE compost were Chelatococcus and Thermobacillus, while Symbiobacterium and Acinetobacter were more abundant in the LBE compost. Ascomycota and Basidiomycota dominated all the composting phases. The Ascomycota genera Colletotrichum, Pichia, Mycothermus, and Thermomyces dominated in phase II of HBE composting. The LBE compost had higher abundances of the Basidiomycota genera Cystofilobasidium and Cryptococcus than the HBE compost. The predicted pathotroph and saprotroph-symbiotroph abundances were more positively linked to HBE composting phase II than to LBE composting. High-biological-efficiency microbial communities are characterized by high pH, carbon, and nitrogen levels. Changes in physiochemical traits, microbial diversity, and metabolism affect the V. volvacea yield.