. 2024 Dec 28;14:31225. doi: 10.1038/s41598-024-82600-7

Effects of mycelium post-ripening time on the yield, quality, and physicochemical properties of Pleurotus geesteranus

Jiling Song ^1,^#, Lin Ma ^3,^#, Tingxuan Zong ², Zhiying Zhang ³, Qing Chen ^2,^✉, Weidong Yuan ^1,^✉

PMCID: PMC11682351 PMID: 39732904

Abstract

This study determined the effects of the mycelium post-ripening time on the growth of Pleurotus geesteranus and the substrate metabolism. The characteristic indexes and timing reflecting the physiological maturity of P. geesteranus mycelium were identified to facilitate precise cultivation in factories. The effects of seven mycelium post-ripening times (20–50 d) on the characteristics, yield, and nutrients of P. geesteranus and the substrate physicochemical properties were investigated using the “Jinxiu” strain. Prolonging the mycelium post-ripening time initially increased and then decreased the yield, high-quality mushroom rate, hardness, and elasticity. Mycelium post-ripening time and yield were positively correlated with the high-quality fruiting rate, total sugar content, and chromatic value, and negatively correlated with the lightness value. Moreover, the mycelium post-ripening time was positively correlated with ergosterol content and catalase activity and negatively correlated with C/N, cellulose content, and superoxide dismutase activity. The most suitable mycelium post-ripening time of P. geesteranus was 35–45 days. The pH and acid protease activities can be indicators for the end of P. geesteranus substrate colonization.

Supplementary Information

The online version contains supplementary material available at 10.1038/s41598-024-82600-7.

Keywords: Pleurotus Geesteranus, Mycelium post-ripening time, Agronomic traits, Nutrient content, Physicochemical properties

Subject terms: Environmental sciences, Plant breeding

Pleurotus geesteranus, scientifically known as P. pulmonarius (Fr.) Quél., is a species of rare edible and medicinal mushroom belonging to Basidiomycota, Agaricales, Pleurotaceae¹. P. geesteranus is tender, tasty, rich in protein, vitamins, amino acids, and other nutrients², and is thus popular among consumers. P. geesteranus also contains polysaccharides, flavonoids, phenolics, and other bioactive substances, which possess anti-tumor^3,4 and immune enhancement⁵ abilities and can aid in preventing alcoholic liver disease¹.

As a rare edible fungus cultivated mainly in Zhejiang Province, China, P. geesteranus has witnessed a steady increase in its cultivation scale. By 2021, the cultivation scale had reached 112 million bags, the output of fresh products was 40190 tons, and the output value was $ 72 million⁶. The production of P. geesteranus is generally conducted under summer facility-based cultivation⁷. This type of cultivation produces P. geesteranus with high fruiting uniformity and several economic benefits. The average yield of the first flush ranged from 89.71 g/bag to 111.71 g/bag, and the biological efficiency was 61.27–81.16%⁸. However, the cultivation cycle of the P. geesteranus is lengthy (eight months, from February to October each year) and the number of mushroom flushes varies six to eight. Consequently, the occurrence of pests and diseases at the late stage of production is prevalent⁹, resulting in a significant reduction in yield and quality (Fig. S1), particularly in light of the rising cost of labor. These issues significantly impede the healthy and sustainable development of China’s P. geesteranus industry¹⁰, underscoring the urgent need to accelerate the transformation and upgrading of this sector.

Despite the initiation of P. geesteranus commercial cultivation in numerous regions since 2022, the technology remains in its infancy. The relatively brief history of this effort has resulted in several significant challenges that require further attention. Determining how to effectively improve the yield and quality of P. geesteranus has been a topic of great interest among scholars engaged in related research, as well as among practitioners involved in its production. Current research into the commercial cultivation of P. geesteranus is focused on three key areas: the development of new varieties¹¹; the determination of new substrate formulations¹²; and the optimization of cultivation techniques¹³. The mycelium post-ripening culture period represents a crucial phase in the transition of the mycelium from a state of substrate colonization to fructification. This phase exerts a significant influence on the quality and yield of edible mushrooms. As demonstrated by Guan¹⁴, an insufficient mycelium post-ripening time often leads to reduced yield, diminished quality, an elevated deformity rate, and, in some cases, the complete absence of fruit bodies. Dedicating too much time to mycelium post-ripening cultivation is likely to result in the aging of the mycelium, which can have adverse effects on fruit body production, lead to a significant waste of resources, and increase production costs. Employing an appropriate mycelium post-ripening time often results in increased yield and quality, and represents a critical node in the factory production process¹⁵. This study examines the indicators important for mushroom cultivation including yield, quality, and the physicochemical properties of the substrate under different times of mycelium post-maturation period. The objective of the study is to determine the timing of the physiological maturity of P. geesteranus mycelium and the evaluation criteria for the maturity of the bag. This work provides a solid foundation for the improvement of P. geesteranus cultivation sector.

Results

Effects of mycelium post-ripening time on the fructification of P. geesteranus

Figure 1 and Table S1 reveal the effect of the variation of the mycelium post-ripening time on P. geesteranus fruit body formation. Compared with T1 and T2, T3–T7 exhibited fruiting uniformly, cap fan-shaped and spread. The harvest period of T7 was the longest of all treatments. The prolongation of mycelium post-ripening time resulted in the corresponding slower fruiting body growth, a longer harvesting period, and mushrooms with uniform appearance and a fan-shaped and spread cap.

Fig. 1 — Effects of mycelium post-ripening time on the fructification of *P. geesteranus*. (A) The average primordia shape for different treatments; (B) The average fruit body shape for different treatments.

Effects of mycelium post-ripening time on the yield and fruiting traits of P. geesteranus

There were significant differences between the different mycelium post-ripening treatments on the yield and substrate traits of P. geesteranus (Fig. 2 and Table S1). The yield and single sub-entity weight tended to increase with the decrease of the mycelium post-ripening time (Fig. 2A and D, respectively). Treatment T4 exhibited the highest value for both parameters, with an average of 258.27 ± 1.68 g/bag and 7.40 ± 0.21 g, respectively. The high-quality fruiting rate and the sub-entity number were observed to initially increase and subsequently decline (Fig. 2B and C, respectively). Treatment T6 exhibited the highest efficacy (93.62 ± 0.92%). For a single fruiting body, the cap length and width demonstrated a biphasic trend, initially decreasing and subsequently increasing (Fig. 2E and F, respectively), with treatment T1 exhibiting the highest value for both parameters (45.87 ± 4.11 mm and 56.12 ± 6.90 mm, respectively). The cap and stipe thickness showed a similar trend that initially increased and subsequently decreased (Fig. 2G and I, respectively). The thickest cap and stipe were observed for T4 (12.72 ± 1.23 mm) and T5 (12.35 ± 1.55 mm). The stipe length followed a trend distinct from the aforementioned traits (Fig. 2H), with T6 exhibiting a significantly higher value than the other treatments (52.72 ± 5.50 mm). The results revealed significant differences in the yield and all fruiting traits across the various mycelium post-ripening times for P. geesteranus. Treatments T4–T6 demonstrated the highest overall yield, heavier individual fruit bodies, small and thick caps, and thick and sturdy stipes.

Fig. 2 — Effects of various mycelium post-ripening times on the yield and fruiting traits of *P. geesteranus*. (A) Yield; (B) Quality mushroom rate; (C) Sub-entity number; (D) Single sub-entity weight; (E) Cap length; (F) Cap width; (G) Cap thickness; (H) Stipe length; (I) Stipe thickness. Lowercase letters indicate that values are significantly different (P < 0.05).

Effects of mycelium post-ripening time on the color and texture of P. geesteranus

The impact of the different mycelium post-ripening time treatments on the hue of the cap of P. geesteranus exhibited notable variations, as illustrated in Fig. 3A–D and Table S2. The L-values demonstrated a declining and subsequently ascending trend with the prolongation of the mycelium post-ripening incubation period (Fig. 3A). The L-value for treatment T4 was significantly lower (46.21 ± 5.66) than that of the other treatments. T4 also exhibited the darkest color. The chromatic values demonstrated an initial increase, followed by a subsequent decline (Fig. 3D), with T4 obtaining the highest value at 5187.41 ± 857.37. The results demonstrate that the fruiting bodies of T4 were dark, commercially viable, and had a glossy appearance.

Fig. 3 — Effects of mycelium post-ripening times on the color and texture of *P. geesteranus*. (A) L value; (B) a value; (C) b value; (D) Chromatic value; (E) Hardness; (F) Elasticity. Lowercase letters indicate that values are significantly different (P < 0.05).

Notable differences were observed in the texture of the P. geesteranus fruiting bodies subjected to distinct mycelium post-ripening times (Fig. 3E and F and Table S2). The hardness and elasticity both showed a tendency to increase and then decrease with the prolongation of the mycelium post-ripening time. The hardness and elasticity of T5 were significantly higher than those of the other treatments, reaching 37.39 N and 5.99 mm, respectively. Thus, the fruiting bodies of T5 P. geesteranus were hard and elastic.

Effects of mycelium post-ripening time on the nutritional characteristics of P. geesteranus

The nutritional content in P. geesteranus exhibited notable variations across the different mycelium post-ripening time treatments (Fig. 4 and Table S3). The protein (Fig. 4A) and total sugar (Fig. 4B) content demonstrated a gradual increase with the incubation time. Conversely, the ash content (Fig. 4A) exhibited a fluctuating trend, initially decreasing and subsequently increasing. The highest ash content was observed for treatment T1, reaching 0.67 ± 0.014%. The crude fiber, fat, and water contents remained relatively stable (Fig. 4A and B).

The highest essential amino acid and amino acid contents were observed in T5 (Fig. 4C), with values of 1.16% and 2.93%, respectively. Amino acid cluster analysis was performed for the P. geesteranus fruiting bodies under different mycelium post-ripening time treatments (Fig. 4D). Based on the results, the seven treatments could be grouped into three distinct taxa. Group 1 comprises T1–T3, with a low amino acid content; group 2 comprises T6 and T7, with a high amino acid content; and group 3 comprises T4 and T5, with the highest amino acid content. The analysis revealed that with the prolongation of mycelium post-ripening time, the nutrient contents in the fruiting body increased gradually. Nutrition was abundant when the mycelium post-ripening time lasted from day 35 to day 50 (T4–T7).

Effects of mycelium post-ripening time on the physicochemical properties of substrates

The effects of the mycelium post-ripening time on the physicochemical indexes of P. geesteranus substrates exhibited notable variations (Fig. 5 and Table S4). Except for cellulose content and SOD activity, the greatest significant differences in the indexes were observed for the treatments with long mycelium post-ripening times (T4–T7). Figure 5A shows that as the mycelium post-ripening incubation period is extended, water content displayed an initial increase, followed by a decline. Ergosterol content demonstrated a gradual increase, with T7 exhibiting a significantly higher concentration (253.17 ± 0.20 µg/g) compared to the other treatments. Both C/N and pH initially increased and subsequently declined (Fig. 5B). C/N reached a peak in T6 (46.98 ± 2.89), while the pH was the highest in T4 (6.34 ± 0.08). Cellulose content exhibited a gradual decline, with T1 exhibiting the highest concentration (304.99 ± 8.62 mg/g) (Fig. 5C). Cellulase and SOD activities demonstrated an initial increase followed by a decline, with the highest values observed in T5 and T2, reaching 622.66 ± 19.15 U and 385.83 ± 11.38 U/g, respectively. No significant differences were observed for the three indexes in Fig. 5D. PPO and acid protease activities followed an increasing and then decreasing trend. The highest acid protease and PPO activities were observed in T4 and T5, with values of 84.46 ± 1.82 U/g and 89.64 ± 4.74 U/g, respectively. CAT and MDA demonstrated a gradual increase, peaking in T7 (89.53 ± 2.38 U/g and 83.82 ± 3.42 nmol/g). The results reveal that the different mycelium post-ripening time treatments exerted a strong influence on the nutrient composition and associated enzyme activities within the substrate, but not on amylase activity.

Correlation between mycelium post-ripening time and the fruiting traits, yield, and substrate properties of P. geesteranus

The correlation analysis was performed for the mycelium post-ripening times and different traits (Fig. 6). There were no notable correlations between mycelium post-ripening time and yield. The mycelium post-ripening time was positively correlated with ergosterol, total sugar, and protein, with correlation coefficients of 0.897, 0.923, and 0.942, respectively. Conversely, there was a negative correlation between the mycelium post-ripening time and the cellulose content, C/N, and water content of the substrates, with correlation coefficients of − 0.94, − 0.954, and − 0.977, respectively. Moreover, a significant positive correlation was observed between yield and the pH, hardness, chromatic value, and cap thickness, with correlation coefficients of 0.962, 0.896, 0.901, and 0.891, respectively. A highly significant negative correlation was also observed between yield and the L-value, with a correlation coefficient of − 0.953. In contrast, the correlations between PPO, α-amylase, β-amylase, MDA, ACP, and each of the other traits were found to be weak.

Discussion

The mycelium post-ripening period of edible fungi represents a significant phase of transition, marking the shift from nutritive growth to fructification. This occurs after the mycelium has colonized the bag and continues to be cultivated under suitable conditions. This study demonstrated that the optimal mycelium post-ripening time can effectively ensure the commercially acceptable yield and quality of P. geesteranus. These findings are consistent with those of Zhang et al.¹⁶ and Hu et al.¹⁷ on Hypsizygus marmoreus and P. eryngii, respectively. Thus, the mycelium post-ripening time significantly impacts the control of the appropriate fructification timing and management techniques for P. geesteranus.

The shorter mycelium post-ripening time (20–25 d) resulted in poor fruiting body traits. This is related to the insufficient absorption of nutrients by mycelia to support the nutrient conditions required for fructification¹⁸. The fruiting body characteristics of commercial importance were optimized at the mycelium post-ripening time of 35–45 days. At 50 days of mycelium post-ripening, most of the fruiting body traits had gradually deteriorated. This may be attributed to the prolonged mycelium post-ripening period of the mycelium, which exceeded the optimal duration. The excessive decomposition of the substrate by the mycelium may result in insufficient nutrients for the formation of the primordia and fruiting bodies, thereby reducing the yield¹⁹. The appropriate mycelium post-ripening time of P. geesteranus can notably enhance both the yield and quality of the fruiting body.

The high-quality fruiting rate exhibited a significant positive correlation with the mycelium post-ripening time and the yield, respectively. An appropriate mycelium post-ripening time during the growth of P. geesteranus can effectively enhance the yield and quality of mushrooms²⁰. Based on the yield and fructification of P. geesteranus observed in this study, a mycelium post-ripening time of 35–45 days is recommended for the commercial cultivation of P. geesteranus.

At present, only a limited number of studies have been conducted to investigate the impact of varying mycelium post-ripening time conditions on the nutrient composition of the fruiting bodies of P. geesteranus. Zhang²⁰ found that when the mycelium post-ripening time of Lentinus edodes was prolonged, the crude fiber did not change and the ash content decreased, potentially improving the taste of the fruiting body. The content and composition of proteins, amino acids, and total sugars are key nutritional constituents of edible mushrooms. They serve as important indicators for evaluating the nutritional and medicinal values of edible mushrooms²¹. At the mycelium post-ripening time of 40 days, the contents of protein, total sugar, and amino acids were relatively high. Pearson correlation analysis revealed a strong significant positive correlation between mycelium post-ripening time and protein and total sugar, while yield exhibited a significant positive correlation with total sugar and total amino acids. The optimal mycelium post-ripening time has a marked effect on enhancing the nutrient composition of P. geesteranus^22,23.

The cultivation of edible fungi is achieved through a mycelium post-ripening process, whereby the mycelium absorbs nutrients from the substrate, a crucial factor influencing the growth of edible fungi. The accurate determination of mycelial biomass during cultivation is a challenging process. In this context, ergosterol, a steroidal compound, plays a pivotal role in fungal cell membranes and can be employed to indirectly determine mycelial content in the substrate, as evidenced by previous literature²⁴. The ergosterol content peaked when the mycelium post-ripening time was extended to 50 days. This was accompanied by an increase in mycelial accumulation, which can effectively provide more nutrients and energy for fructification²⁵.

During substrate colonization, mushrooms primarily use sugar, such as glucose and maltose, as a carbon source. From the mycelium post-ripening stage to fructification, nutrients are obtained through crude fibers, such as lignin and cellulose. This process is a key determinant of the mushroom yield²⁶. During the mycelium post-ripening stage, mycelium uses nitrogen and cellulose in hydrolyzed substrate to provide nutrients for primordia formation²⁷. Our analysis of antioxidant enzyme activities suggests that the extension of the mycelium post-ripening stage may have contributed to this observed phenomenon. The optimal cultivation cycle not only favors the improvement of enzyme activities in the mycelium and the maintenance of active oxygen metabolism balance in the organism²⁸ but also contributes to the primordium formation²⁹.

Conclusion

A period of 35–45 days of mycelium post-ripening in the cultivation farm of P. geesteranus is sufficient to guarantee the required yield and high quality. The pH and acid protease activities can be employed as key indicators for the vegetative growth of P. geesteranus. This study provides technical support for the superior quality and yield of P. geesteranus produced in a commercial setting.

Methods

Growth conditions and sample collection

The test strain of P. geesteranus used in the present work, named Jinxiu, was preserved in the Hangzhou Academy of Agricultural Sciences (Zhejiang, China). The experiment was conducted at Dingxin Agricultural Science and Technology Co. Ltd, Lin’an, Hangzhou, China, and the mushrooms were produced in accordance with conventional management methods. The substrate comprised of corncob 38%, cottonseed shell 30%, bran 20%, wood chips 11%, soybean meal 0.6%, lime 0.2%, and calcium bicarbonate 0.2%. The cultivation parameters and operation during the incubation period and the fruiting period refer to Rao et al.¹¹. Following the completion of the full bagging process for P. geesteranus (20 d of incubation), the mycelium post-ripening culture was continued. The mycelium post-ripening time treatments were as follows: 20 d (T1); 25 d (T2); 30 d (T3); 35 d (T4); 40 d (T5); 45 d (T6); and 50 d (T7). Three replications were arranged for each treatment and 150 bags with normal germination (no pests and diseases appeared) were selected for mushroom cultivation at each replication. The fructification and yield of P. geesteranus were recorded for each treatment. All selected fruiting body samples were mature⁷. For the substrate samples, five bags were randomly selected for each treatment the day before cold stimulation. A total of 30 g substrate was sampled respectively from the upper, middle, and lower parts of the same bag and mixed well.

Agronomic trait measurements

A total of 30 fruiting bodies were selected from each treatment to measure the cap length, width, and thickness and the stipe length and thickness using Vernier calipers. To determine the yield, yield counts were conducted for each replication and the yield per bag was then calculated. The fruiting uniformity was observed as described by Chen et al.³⁰. The high-quality fruiting rate, the weight ratio of grade A fruiting to whole fruiting, was determined as described by Rao et al.¹¹. Furthermore, 30 mushroom bags were selected randomly from each treatment to record the sub-entity number of each bag. The single sub-entity weight was then calculated from the weight of a single bag and the sub-entity number. Fruiting body cap morphology was classified as Li et al.³¹. The harvest period was taken from the start of cold stimulation to the end of harvesting for one flush.

Color analysis

The L (lightness), a (red-green), and b (yellow-blue) values in the center of the fruiting body cap were determined using a colorimeter (YS3060, Shenzhen ThreeNH Technology Co., Ltd.) in nine replicates. The chromatic value was calculated to further analyze the color and brightness of the fruiting bodies³² as follows:

Texture characterization

The textural properties, hardness and elasticity, of the fruiting bodies of P. geesteranus under different treatments were examined using a texture analyzer. The specific parameters were determined with reference to previous literature³³.

Nutrient content

The nutritional composition of the fruiting body of P. geesteranus was tested according to national standards. Crude fiber was determined following the standard GB/T 5009.10-2003. Fat was determined following GB 5009.6-2016. Ash was determined following GB/T 5009.4-2016. Protein was determined following GB/T 5009.5-2016. Water was determined following GB/T 5009.3-2016. Total sugar was determined following GB/T 15,672 − 2009. Amino acids were determined following GB 5009.4-2016.

Physicochemical properties of the substrate

The physicochemical properties of the substrates were tested according to the references listed below. Total carbon was determined by the volumetric method using potassium bicarbonate³⁴. Total nitrogen content was determined by the Kjeldahl method³⁵. The C/N value was calculated as the ratio of total carbon to total nitrogen. Water content was determined by the mass method³⁶. The pH was determined using an acidometer³⁶. Cellulose content and cellulase activity were determined using anthrone colorimetry³⁷. The activities of α-amylase and β-amylase were determined using the 3.5-dinitrosalicylic acid (DNS) colorimetric method³⁸. Acid protease activity was determined using the forintol colorimetric method³⁹. Polyphenol oxidase (PPO) activity was quantified using the catechol colorimetric method⁴⁰. Superoxide dismutase (SOD) activity was determined using the nitrogen blue tetrazolium method, as described by Beauchamp et al.⁴¹. Catalase (CAT) activity was determined by the ultraviolet absorption method⁴². Malondialdehyde (MDA) content was determined using the thiobarbituric acid colorimetric method⁴³. Acid phosphatase (ACP) concentration was determined by the colorimetric method using disodium benzene phosphate⁴⁴. Ergosterol content was determined by high-performance liquid chromatography⁴⁵.

Statistical analysis

The data were analyzed using standard deviation (SD) and one-way analysis of variance. Differences between treatments were evaluated using Duncan’s test (P = 0.05) in SPSS Statistics 25 (IBM). The results were plotted using Origin 2021 (OriginLab).

Electronic supplementary material

Below is the link to the electronic supplementary material.

Supplementary Material 1^{(555KB, pdf)}

Abbreviations

DNS: 3.5-dinitrosalicylic acid
PPO: Polyphenol oxidase
SOD: Superoxide dismutase
CAT: Catalase
MDA: Malondialdehyde
ACP: Acid phosphatase
SD: Standard deviation

Author contributions

J.S., Q.C., and W.Y. conceived and designed the experiments; X.Z. collected the samples; L.M. and Y.Z. analyzed the data and wrote the manuscript; and J.S. and L.M. revised and approved the final version of the paper. J.S. and L.M. contributed to the work equally and should be regarded as co-first authors.

Funding

Project of Collaborative Extension of Major Agricultural Technologies of Zhejiang (No. 2021XTTGSYJ01-1, 2023ZDXT08-3) and the China Agriculture Research System of MOF and MARA (No. CARS-20).

Data availability

The data of this study are included in the article or the Supplementary Materials.

Declarations

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

These authors contributed equally: Jiling Song and Lin Ma.

Contributor Information

Qing Chen, Email: chenq501@163.com.

Weidong Yuan, Email: ywd0507@126.com.

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PERMALINK

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