Effect of different nutritional materials on laccase activity and biomass accumulation of the Auricularia cornea var. Li strain

Xiong, X., Li, P., Xiang, Z., He, Y.-W., Zheng, X., and Liu, Z.-X. (2025). "Effect of different nutritional materials on laccase activity and biomass accumulation of the Auricularia cornea var. Li strain," BioResources 20(1), 1713–1724.

Abstract

The laccase activity changes of the Auricularia cornea var. Li strain were studied over a continuous 9-day period using different carbon sources, nitrogen sources, and lignocellulose as liquid fermentation inducers. The results showed that the addition of carbon sources, nitrogen sources, and alkaline lignin all stimulated Auricularia cornea var. Li to secrete laccase and promoted the accumulation of mycelial biomass. Both carbon and nitrogen deficiencies could stimulate Auricularia cornea var. Li to produce more laccase, but they were detrimental to the accumulation of mycelial biomass. Maltose and peptone should be prioritized as materials for cultivating high-laccase-producing Auricularia cornea var. Li strains through liquid fermentation. Lignocellulosic biomass could significantly enhance laccase activity in Auricularia cornea var. Li. During the cultivation of Auricularia cornea var. Li., wheat bran and cottonseed hulls that produced high levels of laccase should be recommended. This study partially revealed the laccase production characteristics of Auricularia cornea var. Li and identified culture substances that were beneficial for laccase secretion during different growth stages of the mushroom. The results provide a foundation for improving the yield and quality of Auricularia cornea var. Li.

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**Effect of Different Nutritional Materials on Laccase Activity and Biomass Accumulation of the Auricularia cornea var. Li Strain**

Xue Xiong , Peng Li , Zhun Xiang,* Yao-Wei He , Xuan Zheng, and Zhong-Xuan Liu

DOI: 10.15376/biores.20.1.1713-1724

Keywords: Auricularia cornea var. Li; Laccase activity; Carbon sources; Nitrogen sources; Mycelial biomass; Lignocellulose

Contact information: Guizhou Institute of Biology, Guiyang, Guizhou 550009 China; * Corresponding author: 564375391@qq.com

INTRODUCTION

Edible fungi have garnered significant attention due to their fruiting bodies being rich in nutritional, medicinal, and economic value (Jacinto-Azevedo et al. 2021; Zhang et al. 2021; Dimopoulou et al. 2022). Various edible fungi have been isolated from nature and cultivated on a large scale for market supply (Jacinto-Azevedo et al. 2021; Zhang et al. 2021; Dimopoulou et al. 2022; Juárez-Hernández et al. 2023). During the process of artificial domestication and cultivation, different carbon and nitrogen sources often have been used to explore the characteristics of edible mushroom strains (Li et al. 2021b). Various lignocellulosic biomasses are commonly used in the large-scale cultivation of edible fungi to provide nutrients for mycelial growth and fruiting body development (Luo and Chen 2010; Kumla et al. 2020; Viriato et al. 2022). Lignocellulosic biomass is primarily composed of long-chain cellulose and hemicellulose, which are interconnected by heterogeneous polymerization of lignin units (Nunes and Kunamneni 2018; Govil et al. 2022; Jindal et al. 2023). Edible fungi can obtain sufficient nutrients from lignocellulose, due to their strong secretion capacity of lignin-degrading enzymes (Kumla et al. 2020; Su et al. 2023). Lignin-degrading enzymes can break down lignin, thereby disrupting the microfibrillar structure and allowing cellulose to be efficiently hydrolyzed into sugars (Nunes and Kunamneni 2018; Govil et al. 2022; Jindal et al. 2023). Some studies have suggested a correlation among the yield of edible fungi, the production of lignin-degrading enzymes, and the degradation of lignin (Fang et al. 2018; Colla et al. 2023; Su et al. 2023).

Auricularia cornea var. Li is a novel white variety from Auricularia cornea Ehrenb (Wang et al. 2018; Lin et al. 2024). This variant is rich in protein, dietary fiber, polysaccharides, unsaturated fatty acids, and various trace elements, offering significance in both nutritional and medical contexts (Wang et al. 2018; Lin et al. 2024). Laccase is a representative enzyme of lignin-degrading enzymes, capable of completely degrading lignin into carbon dioxide and water (Janusz et al. 2013). Laccase is the most ideal green catalyst for biological degradation (Janusz et al. 2013). The mycelial growth and fruiting body development of edible fungi are closely related to their laccase activity (Sun et al. 2011; Fang et al. 2018; Cesur et al. 2022). Fungal strains with high laccase activity often exhibit faster mycelial growth and shorter growth cycles (Sun et al. 2011). Volvariella brumalis strains could grow rapidly when cultivated on lignocellulosic substrates that promote high laccase production (Xiong et al. 2024). The Auricularia cornea var. Li strains with the highest laccase activity also yielded the highest fruiting body production (Fang et al. 2018). Laccase expression is influenced by multiple factors, including the developmental stage and cultivation environment (Rivera-Hoyos et al. 2013; Reddy and Kanwal 2022). Factors, such as cultivation methods, complexity and concentration ratios of carbon and nitrogen sources, metal ions, secondary metabolites, temperature, and pH levels, can impact laccase activity (Rivera-Hoyos et al. 2013; Durán-Sequeda et al. 2021; Razavi et al. 2021; Reddy and Kanwal 2022; Colla et al. 2023; González-González et al. 2023). Carbon sources, nitrogen sources, and other inducers often play the major role in influencing laccase activity (Rivera-Hoyos et al. 2013; Reddy and Kanwal 2022). Laccase production can be triggered by the depletion of carbon or nitrogen sources and is influenced by the type and concentration of these sources (Janusz et al. 2013, 2015). The inducers typically refer to water-soluble lignocellulosic materials and small molecular aromatic compounds with structures similar to lignin, or metabolites derived from lignin degradation, such as alkaline lignin, ferulic acid, and lignin-derived phenols (An et al. 2018; Swatek and Staszczak 2020; Reddy and Kanwal 2022). These substances promote laccase secretion (An et al. 2018; Swatek and Staszczak 2020; Reddy and Kanwal 2022). Various lignocellulosic biomasses not only act as inducers for laccase production (Atilano-Camino et al. 2020; Li et al. 2021a; Xiong et al. 2021, 2024; Colla et al. 2023), but also serve as the primary nutrient source for edible fungi during the cultivation process (Luo and Chen 2010; Kumla et al. 2020; Viriato et al. 2022).

Therefore, alkaline lignin was used as an inducer, while different carbon sources, nitrogen sources, or lignocelluloses were utilized for the liquid fermentation of Auricularia cornea var. Li strains. The changes in laccase activity under each induction medium were continuously monitored for 10 days, while the biomass of mycelium accumulated under carbon-limiting or nitrogen-limiting conditions was also measured. The aim was to study the secretion patterns of laccase and the accumulation of biomass in A. cornea var. Li under conditions of limited carbon or nitrogen sources. Results showed that the laccase expression characteristics of A. cornea var. Li under liquid fermentation with different nutrient sources. The cultivating materials for high laccase production in A. cornea var. Li are recommended, based on this work. These conclusions may lay a theoretical foundation for accelerating the growth of A. cornea var. Li mycelium and improving the yield and quality of fruiting bodies.

EXPERIMENTAL

Materials

The tested Auricularia cornea var. Li strain was provided by the Edible Fungi Engineering Center of Guizhou Province, and the DNA testing of the strain was conducted in the Guizhou Institute of Biology, China (Li et al. 2021b).

Induction media

The Malt Extract Agar (MEA) Media used was comprised of the following: 10 g/L glucose, 20 g/L malt extract, 3 g/L potassium dihydrogen phosphate, and 20 g/L agar. All biological reagents were dissolved in deionized water, and the volume was adjusted to 1 L. The MEA media were sterilized in the autoclave at 121 °C for 20 min in an autoclave.

Liquid-state fermentation media was comprised of the following: 20 g/L glucose, 2 g/L peptone, 2 g/L yeast extract, 0.5 g/L MgSO₄·7H₂O, 1 g/L K₂HPO₄·3H₂O, and 0.46 g/L KH₂PO₄. All biological reagents were dissolved in deionized water, and the volume was adjusted to 1 L. The media were sterilized in the autoclave at 121 °C for 20 min in an autoclave.

Based on the description of fungal culture media for the genus Auricularia in the book Encyclopedia of Mushroom Industry in China (Luo and Chen 2010), several lignocellulosic materials were selected to cultivate Auricularia fungi to induce Auricularia cornea var. Li.

Table 1. The Formulations of Different Induction Media

The materials included corn cobs, Quercus acutissima sawdust, cottonseed hulls, wheat bran, and fermented soybean meal powder. Fermented soybean meal powder and biological reagents, lignin alkali, were purchased from Beijing Solarbio Science & Technology Co., Ltd. (Beijing, China). The other lignocellulosic materials were collected from Guiyang City, Guizhou Province, China. The substrates were chopped into small fragments, sun-dried, and then oven-dried at 60 °C. Some fragments were pulverized to a mesh size ranging between 40-mesh and 60-mesh using an electric grinder. All reagents and materials were dried to constant weight before using. Different induction media were prepared according to the compositions listed in Table 1. Then, the 250-mL Erlenmeyer flasks containing 100 mL of different induction media were sterilized at 121 °C for 30 min in an autoclave.

Methods

Culture method

The Auricularia cornea var. Li strain was activated using MEA media and incubated at 25 °C for 8 days.

The best-growing colonies were selected for inoculum preparation. Five 5-mm diameter mycelial discs were excised from the activated agar plates and transferred into a 250-mL Erlenmeyer flask, which was filled with 100 mL of liquid-state fermentation media. All Erlenmeyer flasks were incubated at 25 °C and shaken at 150 rpm for 10 days. After incubation, the mycelial pellets in the Erlenmeyer flask were homogenized using a modular homogenizer HFJ-10 (Tianjin HengAo Technology Co., Ltd., China) at 5,000 rpm for 180 s. The mycelial pellets were used as inoculums. Then, 3 mL of well-mixed Auricularia cornea var. Li homogenate were added to each of the 250-mL Erlenmeyer flasks containing 100 mL of different induction media. All flasks were incubated in the dark at 25 °C and 150 r/min for 10 days on a shaking incubator, with three replicates for each treatment.

Assay of laccase activity

Starting from day 2 of cultivation, 2 mL of crude enzyme extract was collected daily from each treatment. The samples were centrifuged at a temperature of 4 ℃ with a rotational speed of 12,000 rpm for 20 min. Following centrifugation, the supernatants were stored at a temperature of -80 ℃ for subsequent analysis of the laccase activity.

The laccase activity was assayed with 2,2′-azinobis-[3-ethyltiazoline-6-sulfonate) (ABTS) (Xiong et al. 2021). The amount of enzyme required to convert 1 μmol of ABTS per minute was defined as one unit of enzyme activity. The molar extinction coefficient of ABTS at 420 nm is 3.6 × 10⁴L/(mol⋅cm).

Mycelial biomass measurement

Starting from day 2 of cultivation, mycelial biomass was measured every two days, with three replicates for each treatment. After filtering the mycelial mats through double-layer filter paper, they were rinsed multiple times with distilled water, dried at 70 °C to a constant weight, and the dry weight recorded.

Data statistics

Analyses of variance (ANOVA) between the test groups were performed with the PASW Statistics 18.0 (International Business Machines Corporation, version 18.0, New York, NY, USA). The figures were created by WPS Office (Kingsoft, V12.1.0.16388, Beijing, China).

RESULTS AND DISCUSSION

**Effects of Different Carbon and Nitrogen Sources on Laccase Activity and Mycelial Biomass Production in Liquid Fermentation of Auricularia cornea var. Li**

As shown in Table 1, different carbon and nitrogen sources had a highly significant effect on laccase activity during the liquid fermentation of Auricularia cornea var. Li throughout the entire cultivation period (P < 0.001). On day 5 of cultivation, different carbon and nitrogen sources had the most pronounced effect on laccase activity (Table 1). Except for day 2 after cultivation, different carbon and nitrogen sources significantly affected the accumulation of mycelial biomass in A. cornea var. Li (P < 0.001). After 8 days of cultivation, the rate of increase in mycelial biomass of A. cornea var. Li began to slow down (Table 1).

Laccase activities of A. cornea var. Li in the induction media X1 to X9 showed a trend of initially increasing and then decreasing with extended cultivation time (Fig. 1A), while the mycelial biomass exhibited an upward trend (Fig. 1B). Compared to the induction medium X1, the addition of alkaline lignin (X2) was beneficial for A. cornea var. Li to secrete laccase and accumulate mycelial biomass (Fig. 1). Alkaline lignin consistently demonstrated a notable induction capacity for laccase production in previous fungal laccase induction experiments (An et al. 2018; Xiong et al. 2024). Compared to the induction media X3, the increasing of carbon sources (X4, X6, X7), nitrogen sources (X5, X8, X9), or the combination of carbon and nitrogen sources (X2) could stimulate A. cornea var. Li to produce more laccase and accumulate greater biomass (Fig. 1). Compared to the induction of media X2, both nitrogen deficiency (X4, X6, X7) and carbon deficiency (X5, X8, X9) hindered the accumulation of mycelial biomass in A. cornea var. Li. The carbon deficiency had a greater impact on the mycelial biomass of A. cornea var. Li than the nitrogen deficiency (Fig. 1B). However, compared to the induction of medium X2, the laccase activity of A. cornea var. Li under carbon deficiency (X5) was higher than that under nitrogen deficiency (X4). This was likely related to the inhibitory effect of glucose on laccase expression, which had also been observed in other edible mushroom strains (Galhaup et al. 2002; Mikiashvili et al. 2004; Rivera-Hoyos et al. 2013; An et al. 2018). Researchers have stated that laccase expression can be influenced by adjusting glucose concentration (Galhaup et al. 2002; Mikiashvili et al. 2004; Rivera-Hoyos et al. 2013; An et al. 2018). At certain concentrations, glucose could also promote the secretion of laccase in fungi (An et al. 2018). At least under nitrogen deficiency, the addition of glucose promotes both laccase activity expression and biomass accumulation in A. cornea var. Li (Fig. 1). Both carbon and nitrogen limitation could stimulate A. cornea var. Li to produce more laccase (Fig. 1A). The specific activity of laccase also changed depending on the choice of carbon or nitrogen sources (Rivera-Hoyos et al. 2013). Compared to sucrose (X6) or glucose (X4), the carbon source maltose (X7) showed greater advantages in both laccase induction and mycelial biomass accumulation (Fig. 1). The nitrogen source peptone (X5) significantly stimulated laccase production in A. cornea var. Li more than yeast extract (X8) and beef extract (X9) (Fig. 1A), while its effect on mycelial biomass accumulation was second only to yeast extract (Fig. 1B). This suggested that the nitrogen source most favorable for A. cornea var. Li mycelial biomass accumulation was not necessarily the best laccase inducer. Therefore, to obtain the high-yielding laccase-producing liquid strains of A. cornea var. Li, maltose as the carbon source and peptone as the nitrogen source should be recommended.

Table 2. Effects of Different Carbon and Nitrogen Sources on Laccase Activity and Mycelial Biomass Production in Liquid Fermentation of Auricularia cornea var. Li by One-way ANOVA

Fig. 1. The effects of different carbon and nitrogen sources on Laccase activity and mycelial biomass production in liquid fermentation of Auricularia cornea var. Li; A: Laccase activity, B: Mycelial biomass

**Effect of Different Cultivation Substrates on Laccase Activity in Liquid Fermentation of Auricularia cornea var. Li**

The carbon-to-nitrogen ratio of different lignocellulosic materials varies greatly, significantly affecting the mycelial growth, mushroom yield, and quality of edible fungi (Hoa et al. 2015; Kumla et al. 2020; Colla et al. 2023). In the history of edible mushroom cultivation, low-nitrogen substrates have often been chosen as the primary materials, with organic or inorganic nitrogen added as supplementary materials (Luo and Chen 2010; Kumla et al. 2020). The nitrogen content in the cultivation supplements is often higher than that in the primary substrate; however, the primary substrate provides the main nutrients for the fungi (Luo and Chen 2010; Kumla et al. 2020). Based on the characteristics of the cultivation materials, this study employed different primary substrates (corn cobs, Quercus acutissima sawdust, and cottonseed hulls) and supplementary materials (wheat bran and fermented soybean meal powder) to induce laccase secretion in Auricularia cornea var. Li (Luo and Chen 2010; Kumla et al. 2020). Throughout the entire cultivation period, different substrates had a highly significant impact on the laccase activity produced by A. cornea var. Li in liquid fermentation (P < 0.001) (Table 3). On the 6^th day of cultivation, different substrates had the most significant effect on laccase activity of A. cornea var. Li (Table 3, Fig. 2). The enzyme activity detected under different substrates was consistently higher than that observed under nitrogen-limited or carbon-limited conditions, showing a more significant and sustained effect on laccase activity (Tables 2 and 3, Figs. 1 and 2).

As shown in Fig. 3, the laccase activity of A. cornea var. Li induced by different substrates initially increased and then decreased over the cultivation period. The cultivation auxiliary material wheat bran stimulated the secretion of laccase in A. cornea var. Li more effectively than fermented soybean meal powder (Fig. 2). In induction culture experiments with various edible fungi, wheat bran was considered a cultivation auxiliary material that yielded high laccase production (Atilano-Camino et al. 2020; Kumla et al. 2020; Li et al. 2021; Xiong et al. 2021, 2024). Compared to Q. acutissima sawdust and corn cobs, cottonseed hulls more effectively stimulated the secretion of laccase by A. cornea var. Li (Fig. 2). Moreover, all tested cultivation additives demonstrated a stronger ability to stimulate laccase production in A. cornea var. Li compared to the tested primary substrates (Fig. 2). High nitrogen environments could stimulate fungi to produce laccase (D’Agostini et al. 2011; Colla et al. 2023; Pradeep Kumar et al. 2023). This indicated that both carbon or nitrogen limitation and excess could cause variations in laccase activity of the same fungal strain (Figs. 1 and 2). Both the addition of alkaline lignin and lignocellulosic biomass could significantly stimulate the production of more laccase by Auricularia cornea var. Li (Figs. 1, 2). This phenomenon was also observed on the other edible fungi (Xiong et al. 2024). However, the ability of single lignin addition to induce laccase secretion in the A. cornea var. Li strain was lower than that of complex lignocellulosic biomass (Fig. 2). Additionally, due to its single nutrient composition and high cost, pure lignin is not considered in edible mushroom cultivation. Some scholars believe that strains of A. cornea var. Li with high laccase activity tend to have higher fruiting body yields (Fang et al. 2018). Substrate formulations that yield high laccase production could promote the growth of fungal mycelia (Xiong et al. 2024). Therefore, the laccase activity of A. cornea var. Li could be enhanced by adjusting the substrate formulation, which could lead to increased yield. Therefore, the tested lignocellulosic biomass could be used to cultivate A. cornea var. Li, and cottonseed hulls and wheat bran should be prioritized (Luo and Chen 2010).

Table 3. Effect of Different Cultivation Substrates on Laccase Activity in Liquid Fermentation of Auricularia cornea var. Li by One-way ANOVA

Fig. 2. The effect of different cultivation substrates on laccase activity in liquid fermentation of Auricularia cornea var. Li

CONCLUSIONS

The addition of alkaline lignin was beneficial for Auricularia cornea var. Li to secrete laccase and accumulate mycelial biomass. Based on the inducer alkaline lignin, the addition of carbon sources, nitrogen sources, or a combination of both could stimulate Auricularia cornea var. Li to produce more laccase and accumulate greater biomass. Both carbon and nitrogen deficiencies could stimulate Auricularia cornea var. Li to produce more laccase, but they were detrimental to the accumulation of mycelial biomass. Carbon deficiency had a more significant impact on the biomass accumulation of Auricularia cornea var. Li than nitrogen deficiency.
Compared to sucrose and glucose, the carbon source maltose demonstrated greater advantages in both inducing laccase production and accumulating mycelial biomass of Auricularia cornea var. Li. The nitrogen source peptone significantly stimulated laccase production in Auricularia cornea var. Li outperforming yeast extract and beef extract, while being second only to yeast extract in promoting mycelial biomass accumulation. Therefore, maltose and peptone should be used with priority for cultivating high laccase-producing Auricularia cornea var. Li strains through liquid fermentation.
The cultivation supplement wheat bran was more effective than fermented soybean meal in stimulating Auricularia cornea var. Li to secrete laccase. The cultivation substrate cottonseed hulls were more effective than Q. acutissima sawdust and corn cob in stimulating Auricularia cornea var. Li to secrete laccase. The addition of single lignin (alkaline lignin) induced lower laccase secretion in Auricularia cornea var. Li strains compared to complex lignocellulosic biomass. Wheat bran and cottonseed hulls should be recommended in cultivation to develop an optimal substrate formulation for high laccase production, thereby enhancing the growth rate and yield of Auricularia cornea var. Li.

ACKNOWLEDGMENTS

The work was supported by the Science and Technology Plan Project issued by the Department of Science and Technology of Guizhou Province (QKHZYD[2024]026) and Task Book for the Construction of Functional Laboratories for the Edible Mushroom Industry System in Guizhou Province (2025 Annual Project) (GZSYJCYJSTX-03). The authors are grateful to the Guizhou Institute of Biology for providing the experimental platform and to Professor Dequn Zhou for his guidance and encouragement in writing of this article.

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Article submitted: October 17, 2024; Peer review completed: December 9, 2024; Revised version received: December 10, 2024; Accepted: December 13, 2024; Published: January 6, 2025.

DOI: 10.15376/biores.20.1.1713-1724