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. 2026 May 25;36:104025. doi: 10.1016/j.fochx.2026.104025

Comparison of volatile compounds in Grifola frondosa under different cooking methods using HS-SPME-GC–MS and HS-GC-IMS

Zitong Liu a, Jian Zhao a, Ziheng Cheng a, Xinfang Zhang a, Meifan Pan a, Beilei Pang a, Ruiqi Zhang a, Piaopiao Wei b,c, Xuemei Cheng d, Yudong Wang a, Yongye Sun a, Lirong Xu a,b,⁎
PMCID: PMC13235474  PMID: 42255271

Abstract

Grifola frondosa is a popular mushroom, and its flavor is highly dependent on the cooking method. In this study, the volatile compounds of Grifola frondosa under different cooking methods were identified. HS-SPME-GC–MS identified 36 volatile compounds, mainly aldehydes and alcohols, e.g. nonanal, decanal, 1-octen-3-ol and 1-octanol. HS-GC-IMS indicated that volatile compounds in Grifola frondosa were optimally retained under microwaving cooking for 10 min. Among them, the content of typical mushroom flavor compounds 1-octen-3-ol and 2-butanone (D) reached the highest when microwaving cooking for 10 min. The content of isoamyl formate with plum odor was the highest in the stir-frying cooking sample for 10 min. The content of acetic acid hexyl ester and 2-pentylfuran with fruity flavor reached the highest in the sample with steaming cooking for 10 min. The flavor of boiling cooking samples was the weakest. This study can provide a theoretical foundation for enhancing the flavor quality of Grifola frondosa.

Keywords: Grifola frondosa, Volatile compounds, Cooking methods, HS-SPME-GC–MS, HS-GC-IMS

Graphical abstract

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Comparison of volatile compounds in Grifola frondosa under different cooking methods using HS-SPME-GC–MS and HS-GC-IMS TOC map.

Highlights

  • •

    The flavor retention of Grifola frondosa was better in cooking for 10 min.

  • •

    Aldehydes and esters were key contributors to the mushroom's flavor profile

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    Microwaving can retain volatile compounds well, followed by stir-frying and steaming.

1. Introduction

Grifola frondosa belongs to Basidiomycetes fungi. It is one of the edible and medicinal mushrooms, and is praised for its unique flavor and health benefits. Grifola frondosa is famous for its unique appearance and rich nutritional value. In China, Grifola frondosa is known as gray tree flower (He et al., 2017). It is not only edible, but also considered as a healthy food because it contains protein, carbohydrates, dietary fiber and minerals. Grifola frondosa has low fat content and relatively low calories. It also has antioxidant, anti-tumor, immunomodulatory and anti-diabetic effects (Wu et al., 2021). These functions are closely related to their bioactive components. Grifola frondosa has good taste and unique flavor, which is known as the first grade edible fungus (Mizuno & Zhuang, 1995). It is not only a leader in edible fungi, but also a medicinal fungus, and it is widely used as a cooking ingredient and dietary supplement in Asia (Wu et al., 2016).

Cooking is a traditional thermal processing process of food raw materials. It can improve the sensory characteristics of food. Mushrooms are usually eaten after cooking, rather than raw, which will change the sensory characteristics of edible fungi including color, flavor, texture, and taste (Soler-Rivas et al., 2009). Appropriate cooking methods and time will also make it more delicious. Cooking methods significantly impact the nutritional properties of mushrooms. Studies have found that boiling, microwaving, grilling, and frying affected the basic components of cultivated mushrooms (Irene, Mónica, Margarita & Cristina, 2017). Sun et al. (2019) discovered that four cooking methods (autoclaving, microwaving, sous-vide cooking, and stewing) can increase the polyphenols and flavor compounds in mushroom soup. Marta et al., (2021) showed that most cooking methods reduced the brightness, hardness, adhesiveness, and chewiness of wild delicious boletus edulis, with marinated mushrooms having the lowest flavor value. Previous studies have found that the nutritional components of edible fungi are affected by cooking methods (Luo et al., 2022; Yao et al., 2023). All these experiments showed that different cooking methods can change the nutritional components and flavor of mushrooms. The flavor of mushrooms significantly influence the willingness of people to eat them. Changes in the flavor of mushrooms during cooking, such as increased meat flavor, can be attributed to changes in free amino acids and volatiles during cooking (Kido & Tanaka, 2023). In addition to their high nutritional value, mushrooms also possess unique flavors, with volatile compounds being significant factors influencing their flavor (Liang et al., 2022). Xie et al. (2024) found that 1-octen-3-one was the primary component in button mushrooms. Zhu et al., (2022) pointed out that C8 volatile compounds are important flavor substances in edible fungi, among which 1-octen-3-ol has the highest proportion.

According to the current data, there were few studies on the cooking methods of Grifola frondosa compared with other mushrooms. In view of the potential edible value and medicinal value of Grifola frondosa, as well as its unique flavor, it is necessary to further study the changes of sensory traits and flavor components of Grifola frondosa under different cooking methods. Four cooking methods of stir-frying, steaming, boiling and microwaving were used to process Grifola frondosa. The cooking time is set to 5 min, 10 min, 15 min and 20 min. After cooking, the sensory evaluation of color, texture, flavor and taste in Grifola frondosa were evaluated. The volatile compound changes in Grifola frondosa were analyzed using headspace solid-phase microextraction gas chromatography–mass spectrometry (HS-SPME-GC–MS) and headspace gas chromatography-ion mobility spectrometry (HS-GC-IMS) technology. HS-SPME-GC–MS technology enables the qualitative detection of flavor compounds while combining the safety of HS-SPME with the high accuracy and separation capability of GC–MS, making it the most commonly used flavor analysis technique in edible fungi (Zhong et al., 2024). HS-GC-IMS is a novel gas separation technology that allows for visual analysis of volatile compounds. Compared to HS-SPME-GC–MS, HS-GC-IMS lacks a comprehensive database but offers advantages such as high analysis efficiency and low cost (Li et al., 2024). Therefore, the combined use of HS-SPME-GC–MS and HS-GC-IMS for detecting volatile compounds enables a more comprehensive acquisition of sample information. This study aims to identify rational cooking methods for Grifola frondosa to optimize its flavor, thereby providing references for its cooking practices.

2. Materials and methods

2.1. Materials

Grifola frondosa was harvested from Xi'an Baisite Biotechnology Co., Ltd. in 2024, and stored at 25 ± 2 °C. According to AOAC official method and slightly modified (Association of Official Analytical Chemists, 1990). The specific method is as follows: the Grifola frondosa was placed in the oven and dried at 102 °C for 4 h until a constant weight was achieved. The crude protein content was determined by Kjeldahl method accoding to AOAC official method 978.04. The crude fat content was determined by Soxhlet extraction method accoding to AOAC official method 966.15. The ash content was determined by roasting at 600 °C for 4 h accoding to AOAC official method 942.05. The carbohydrate content was calculated by the difference method, and the calculation formula was: carbohydrate content (%) = 100%-crude protein content (%)-crude fat content (%)-crude ash content (%). The main chemical components of the raw materials are: carbohydrate 70.4%, crude protein 18.6%, crude fat 4.5%, crude ash 6.5%. The final results of all components are expressed on a dry weight basis.

2.2. Cooking methods

Grifola frondosa was soaked in warm water at 50 °C for 30 min. After being completely softened, The Grifola frondosa samples were broken into small pieces and the excess water was drained. The treated Grifola frondosa samples were divided into five groups: uncooked, microwaving, stir-frying, steaming and boiling. The weight of each group of Grifola frondosa samples is 90 g. For steaming cooking (SC), water was added to a stainless steel steaming pot (samples:water = 1:10, w/v). After the water was boiled over high heat, the heat was reduced to the lowest level (approximately 120 W) and the samples were steamed for different durations (5 min, 10 min, 15 min, 20 min and 25 min). For microwaving cooking (MC), the prepared Grifola frondosa were placed in a microwave-safe bowl with ventilation holes, and then the bowl was transferred into the microwave oven. The power of 70 W was selected because it was the lowest stable power level that could achieve mild and uniform heating, and then the samples were treated with microwave for different durations (5 min, 10 min, 15 min, 20 min, 25 min). For boiling cooking (BC), water was added to a stainless steel pot (samples:water = 1:10, w/v). After the water was boiled over high heat, the Grifola frondosa was placed in the pot and boiled for different durations (5 min, 10 min, 15 min, 20 min and 25 min). In the stir-frying cooking (SFC), an induction cooker and a standard round-bottom iron wok were used for stir-frying. 10 g of soybean oil was placed in the iron wok (samples:oil = 9:1, w/w), followed by the addition of prepared Grifola frondosa, which was continuously stirred with a spatula. Grifola frondosa was stir-fried at different time periods (5 min, 10 min, 15 min, 20 min and 25 min). Throughout the process, the induction cooker was set to a constant minimum range (approximately 120 W) to maintain a stable frying temperature based on previous studies (Xu et al., 2026). After each cooking method, the samples of Grifola frondosa were evaluated by sensory evaluation. Then, the samples of Grifola frondosa cooked in different time periods of each cooking method were preserved.

2.3. Sensory evaluation

The sensory evaluation method followed the approach of previous studies (Oh et al., 2024), and was adjusted according to the actual needs and resources of this study. In short, a sensory evaluation group was formed with 20 trained members (10 women and 10 men, aged 20 to 30 years). The group members underwent a two-week training period. Initially, the group members evaluated the samples of Grifola frondosa treated by four cooking methods, and described them from four aspects: color, texture, flavor and taste. In order to reduce the fatigue of group members and focus on the sensory indicators that best distinguish the differences between samples, we selected these four most representative attributes for evaluation after group discussion. Consensus was reached through public discussion, and corresponding terms and definitions were established. Color: intensity of yellow color. Texture: The degree of smooth and wet mushroom surface. Aroma: Odors associated with cooked mushroom. Taste: The taste of chewing mushroom. Subsequently, by continuously observing, tasting, and discussing, the group members refined the set of descriptive attributes and reached agreement. The intensity of the four descriptions was rated on a 0 to 9 scale (with 0 indicating weak and 9 indicating strong). The samples were placed on plates and marked before being given to the panelists for evaluation at room temperature (25 °C). After each tasting, the group leader provided drinking water for each member to rinse out any residual mushrooms. The sensory evaluation study was approved by the Ethics Committee of Qingdao University. Participants signed informed consent forms after fully understanding the purpose of the study and their rights.

2.4. Determination of volatile compounds

2.4.1. HS-SPME-GC–MS analysis

The volatile compounds in Grifola frondosa were extracted by headspace solid phase microextraction (HS-SPME). The SPME method is modified slightly by Xu et al. (2025). The Grifola frondosa samples were placed in a 20 mL vial, and 2 ± 0.02 g was added to each vial. Then the 10 μg pre-configured standard solution (1,2-dichlorobenzene, 51.55 μg/mL) was added to each samples. The samples was extracted using a 50 μm/30 μm DVB/CAR/PDMS fiber (Supelco Inc., Bellefonte, PA, USA). The samples bottle was heated at 60 °C for 30 min in a water bath on a heated magnetic stirrer. After extraction at 60 °C for 30 min, the fiber was immediately desorbed into the GC–MS inlet and maintained at 250 °C for 5 min. According to the scheme described by Xu, Wang, Tian, Liu, et al. (2023). Clarus690SQ8T (PerkinElmer, USA) was used to analyze volatile compounds on GC–MS equipment. The GC conditions were as follows: DB-5MS chromatographic column (30 m × 0.25 mm × 0.25 μm), the temperature program was as follows: the initial column box was kept at 40 °C for 3 min, then increased to 100 °C at a rate of 6 °C/min, and then increased to 230 °C at a rate of 10 °C/min for 3 min. Helium was used as carrier gas (1.0 mL/min).

The mass spectrometry (MS) conditions were EL ion source 230 °C, MS grade rod 150 °C, scanning mass range of 10-550u. The volatile compounds were determined by comparing the retention index (RI) and MS data with the data of the real standard. The quantification of each compound was calculated by internal standard method.

2.4.2. HS-GC-IMS analysis

In terms of headspace gas chromatography-ion mobility spectrometry (HS-GC-IMS), an Agilent 490 gas chromatograph (provided by Agilent Technologies, Palo Alto, CA, USA) and an ion mobility spectrometer (FlavourSpec®, produced by Gesellschaft für Analytische Sensorsysteme mbH, Dortmund, Germany) were used to analyze the volatile components of the samples. The detection conditions were referred to our previous study (Xu, Wang, Tian, Liu, et al., 2023; Xu, Wang, Tian, Wang, et al., 2023). Grifola frondosa samples 2 ± 0.02 g were placed in a 20 mL volumetric flask. The samples bottle was placed in an incubator at 50 °C for 20 min. The samples bottle was placed in an incubator at 50 °C for 20 min, as this temperature and duration were sufficient to reach headspace equilibrium, ensuring adequate release and stabilization of volatile compounds in the gas phase for subsequent HS-GC-IMS analysis (Xu et al., 2025). Then 500 μL of the volatile samples was injected into a 50 °C samplesr, and the mode was not split. MXT-WAX column (30 m × 0.53 mm × 1 μm) was used for detection. The flow of nitrogen was programmed as follows: starting at 2 mL/min (holding for 5 min), increasing to 5 mL/min (holding for 5 min), then to 10 mL/min (holding for 5 min), followed by 20 mL/min (holding for 5 min), 50 mL/min (holding for 5 min), and finally reaching 100 mL/min (holding for 10 min).

The qualitative analysis of the volatile compounds was carried out according to several criteria: the retention index of the volatile compounds, the mass spectra of the compounds in the NIST library, and the drift time of the standards in the GC-IMS library.

2.5. Data analysis

The experiment was designed with biological replicates to account for natural variation in the raw material. Three replicates were used in all experiments, and the data were expressed as mean ± standard deviation. One-way analysis of variance (ANOVA) and two-way analysis of variance were used for data analysis. SPSS statistical software (version 26.0) was used for data analysis. P-value<0.05 was considered statistically significant.

3. Results and discussion

3.1. Sensory evaluation of Grifola frondosa

As shown in Table 1, the sensory evaluation of Grifola frondosa was conducted based on four aspects: color, texture, aroma, and taste. The results showed that at the cooking duration of 10 min, the scores for these four aspects were significantly higher than those at other time points (P < 0.05), suggesting that this duration might represent the optimal cooking time. In terms of aroma, the scores of steamed and boiled samples after 10 min cooking were significantly lower than those of microwaved and stir-fried samples (P < 0.05). Regarding texture, the score of microwaving sample after 10 min cooking was significantly higher than those of the other cooking methods (P < 0.05). This indicated that different cooking methods had a significant impact on the sensory characteristics of Grifola frondosa, which was consistent with previous studies reporting substantial effects of cooking methods on the sensory attributes of mushrooms (Jiang et al., 2023). In summary, the results showed that the comprehensive scores of Grifola frondosa in color, texture, flavor and taste were higher at the 10-min cooking time point, suggesting that this time point may be beneficial to the maintenance of its sensory quality.

Table 1.

The results of sensory evaluation of Grifola frondosa under different cooking methods.

Volatile compounds(μg/kg)
 Cooking method
 Cooking time/min
5 10 15 20 25
Color Steam 7.02 ± 0.18Ab 8.05 ± 0.21Aa 5.98 ± 0.27Bc 5.96 ± 0.18Ac 5.96 ± 0.48Ac
Boil 6.03 ± 0.20Bb 7.05 ± 0.24Ba 5.98 ± 0.27Bb 5.96 ± 0.18Ab 5.96 ± 0.48Ab
Microwave 7.02 ± 0.18Ab 8.05 ± 0.24Aa 5.98 ± 0.27Bb 4.97 ± 0.15Bb 4.97 ± 0.40Bc
Stir-fry 7.02 ± 0.18Ab 8.05 ± 0.24Ab 6.98 ± 0.32Aa 5.96 ± 0.18Ab 4.97 ± 0.40Bb
Aroma Steam 7.02 ± 0.18Ba 7.05 ± 0.21Ca 5.98 ± 0.27Bb 5.96 ± 0.18Ab 5.96 ± 0.48Ab
Boil 8.03 ± 0.23Aa 8.05 ± 0.24Ba 6.98 ± 0.32Ab 4.97 ± 0.15Bc 4.97 ± 0.40Bc
Microwave 6.02 ± 0.15Ca 9.06 ± 0.27Aa 5.98 ± 0.27Bb 4.97 ± 0.15Bb 4.97 ± 0.40Bb
Stir-fry 8.03 ± 0.23Ab 9.05 ± 0.21Aa 6.98 ± 0.32Ac 5.96 ± 0.18Ac 3.97 ± 0.32Cd
Texture Steam 8.03 ± 0.20Aa 6.06 ± 0.27Db 7.97 ± 0.36Ac 7.95 ± 0.24Ad 6.95 ± 0.56Ad
Boil 7.02 ± 0.18Bb 7.05 ± 0.21Ca 5.98 ± 0.27Bb 5.96 ± 0.18Bc 5.96 ± 0.48Bc
Microwave 6.02 ± 0.15Cb 9.04 ± 0.18Aa 4.98 ± 0.23Cc 3.97 ± 0.12Dd 3.97 ± 0.32Cd
Stir-fry 7.02 ± 0.18Bb 8.05 ± 0.24Ba 5.98 ± 0.27Bc 4.97 ± 0.15Cd 3.97 ± 0.32Cd
Taste Steam 7.02 ± 0.18Ab 7.05 ± 0.21Ba 7.97 ± 0.36Ab 6.95 ± 0.21Ac 6.95 ± 0.56Ad
Boil 7.02 ± 0.18Aa 8.05 ± 0.24Aa 5.98 ± 0.27Cb 5.96 ± 0.18Bc 3.97 ± 0.32Cd
Microwave 7.02 ± 0.18Ab 8.05 ± 0.24Aa 5.98 ± 0.27Cc 4.97 ± 0.15Cd 4.97 ± 0.40Be
Stir-fry 7.02 ± 0.18Aa 7.05 ± 0.21Ba 6.98 ± 0.32Ba 4.97 ± 0.15Cb 3.97 ± 0.32Cc
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The values of different letters in each line are considered to be significantly different at P < 0.05. Capital letters represent the comparison of differences in different cooking methods, and lowercase letters represent the comparison of differences in different cooking time.

3.2. HS-SPME-GC–MS analysis of volatile compounds in Grifola frondosa under different cooking methods

A total of 9 aldehyde compounds, 5 alcohol compounds, 5 ester compounds, 3 amide compounds, 5 alkene compounds, 4 ketone compounds and 5 other compounds were identified by HS-SPME-GC–MS analysis (Table 2). The results indicate that aldehydes and alcohols are the main contributors to the flavor of Grifola frondosa. A variety of aldehyde compounds were detected, which exhibit relatively low odor thresholds and strong aroma activity. In contrast, although alcohols have relatively higher thresholds, they were found in significantly higher total concentrations, thereby compensating for their lower inherent potency. Overall, they dominate the volatile profile and are responsible for the characteristic flavor of Grifola frondosa. It was similar to the results of Splivallo et al. (2007), who observed aldehydes and alcohols as the main volatile flavor contributors in Lentinula edodes. Among these compounds, 2 volatile compounds were detected in the uncooked samples, while 14 volatile compounds were found after cooking.

Table 2.

GC–MS analysis results of volatile components of Grifola frondosa under different cooking methods.

Volatile compounds
 Uncooked
 Cooking method
 Cooking time/min
(μg/kg) 5 10 15 20 25
aldehydes
Benzaldehyde 78.95 ± 2.78 Steam 27.50 ± 0.97Ba 8.12 ± 0.29Bd 22.73 ± 0.80Bb 15.40 ± 0.54Bc 16.06 ± 0.57Bc
Boil 13.57 ± 0.48Ca 2.6 ± 0.09Cc 2.16 ± 0.08Cd 5.07 ± 0.18Cb 2.64 ± 0.09Dc
Microwave 34.9 ± 1.23Aa 31.15 ± 1.10Abc 29.42 ± 1.04Ac 32.81 ± 1.16Ab 19.65 ± 0.69Ad
Stir-fry 1.22 ± 0.04Dd 0.70 ± 0.02De 2.71 ± 0.10Cc 14.51 ± 0.51Ba 8.85 ± 0.31Cb
2-Nonenal, (E)- 7.95 ± 0.28 Steam 6.53 ± 0.23Aa 5.22 ± 0.18Ab 4.09 ± 0.14Bc 2.46 ± 0.09Be 3.45 ± 0.12Ad
Boil 3.65 ± 0.13Ca 2.48 ± 0.09Dc 3.33 ± 0.12Cb 1.52 ± 0.05De 1.93 ± 0.07Cd
Microwave 5.92 ± 0.21Bb 4.68 ± 0.16Bc 6.84 ± 0.24Aa 4.79 ± 0.17Ac 1.07 ± 0.04Dd
Stir-fry 1.02 ± 0.04De 2.89 ± 0.10Ca 1.35 ± 0.05Dd 1.71 ± 0.06Cc 2.15 ± 0.08Bb
5-Indanecarboxaldehyde 8.03 ± 0.28 Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry ND ND ND ND ND
Benzeneacetaldehyde 130.58 ± 4.6 Steam 25.01 ± 0.88Ba 36.39 ± 1.28Bb 18.57 ± 0.65Bc 13.28 ± 0.47Bd 11.08 ± 0.39Ce
Boil 2.97 ± 0.10Db 2.08 ± 0.07Cc 4.83 ± 0.17Da 3.04 ± 0.11Cb 2.02 ± 0.07Dc
Microwave 86.72 ± 3.06Aa 52.24 ± 1.84Ac 46.77 ± 1.65Ad 79.03 ± 2.78Ab 38.32 ± 1.35Ae
Stir-fry 11.87 ± 0.42Cd 12.49 ± 0.44Dcd 14.65 ± 0.52Cb 13.01 ± 0.46Bc 25.49 ± 0.90Ba
2,4-Nonadienal, (E,E)- ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 1.37 ± 0.05d 2.09 ± 0.07c 1.98 ± 0.07c 3.93 ± 0.14b 12.01 ± 0.42a
Decanal 9.87 ± 0.35 Steam 9.81 ± 0.35Ba 15.44 ± 0.54Ab 6.21 ± 0.22Ac 5.92 ± 0.21Ac 4.8 ± 0.17Ad
Boil 5.89 ± 0.21Db 6.08 ± 0.21Bb 6.47 ± 0.23Aa 1.57 ± 0.06Cd 3.54 ± 0.12Bc
Microwave 10.78 ± 0.38Aa 5.85 ± 0.21Bb 3.59 ± 0.13Cd 4.13 ± 0.15Bc 2.67 ± 0.09Ce
Stir-fry 1.13 ± 0.04Ca 0.43 ± 0.02Cd 0.68 ± 0.02Dc 0.89 ± 0.03Db 0.93 ± 0.03Db
Heptanal 28.46 ± 1 Steam 19.23 ± 0.68Aa 7.85 ± 0.28 Cb 4.13 ± 0.15Cc 2.98 ± 0.11Cd 2.09 ± 0.07Ce
Boil 4.94 ± 0.17Cb 5.03 ± 0.18Db 7.66 ± 0.27Ba 4.25 ± 0.15Bc 1.73 ± 0.06Dd
Microwave 4.75 ± 0.17Cc 14.40 ± 0.51Aa 12.69 ± 0.45Ab 14.88 ± 0.52Aa 2.79 ± 0.10Bd
Stir-fry 7.29 ± 0.26Bc 11.87 ± 0.42Bb 12.47 ± 0.44Aa 3.45 ± 0.12Cd 3.12 ± 0.11Ad
Hexanal 39.46 ± 1.39 Steam 9.40 ± 0.33Dc 31.24 ± 1.10Ba 18.67 ± 0.66Cb 8.87 ± 0.31Cc 7.52 ± 0.27Bd
Boil 18.26 ± 0.64Bc 25.29 ± 0.89Cb 31.00 ± 1.09Aa 1.39 ± 0.05De 5.00 ± 0.18Cd
Microwave 68.03 ± 2.4Aa 65.47 ± 2.31Aa 22.17 ± 0.78Bb 15.71 ± 0.55Bc 5.82 ± 0.21Cd
Stir-fry 12.18 ± 0.43Cd 14.72 ± 0.52Dc 17.83 ± 0.63Cb 22.68 ± 0.80Aa 23.05 ± 0.81Aa
Nonanal 44.04 ± 1.55 Steam 66.39 ± 2.34Ab 86.23 ± 3.04Aa 46.26 ± 1.63Ac 35.23 ± 1.24Ad 44.95 ± 1.58Ac
Boil 37.73 ± 1.33Cb 39.12 ± 1.38Bb 43.95 ± 1.55Aa 10.44 ± 0.37Dd 19.06 ± 0.67Bc
Microwave 53.77 ± 1.89Ba 42.37 ± 1.49Bb 37.39 ± 1.32Bc 25.63 ± 0.90Bd 14.21 ± 0.50Ce
Stir-fry 11.62 ± 0.41Dc 6.75 ± 0.24Cd 18.17 ± 0.64Ca 17.43 ± 0.61Ca 12.75 ± 0.45Cb
alcohols
1-Decanol ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 0.45 ± 0.02a 0.17 ± 0.01e 0.23 ± 0.01d 0.32 ± 0.01b 0.29 ± 0.01c
1-Nonanol ND Steam 8.96 ± 0.32Cb 15.67 ± 0.55Aa 5.33 ± 0.19Cc 3.70 ± 0.13Cd 4.85 ± 0.17Bc
Boil 12.65 ± 0.45Ba 6.17 ± 0.22Cb 4.62 ± 0.16Dc 1.52 ± 0.05Dd 1.93 ± 0.07Cd
Microwave 5.92 ± 0.21Dc 4.68 ± 0.16Dd 6.30 ± 0.22Bb 6.95 ± 0.24Aa 2.23 ± 0.08Ce
Stir-fry 14.36 ± 0.51Aa 7.48 ± 0.26Bb 7.54 ± 0.27Ab 6.01 ± 0.21Bc 7.11 ± 0.25Ab
1-Octanol 95.7 ± 3.37 Steam 122.97 ± 4.33Bb 191.00 ± 6.73Aa 63.73 ± 2.25Bc 52.91 ± 1.86Bd 44.65 ± 1.57Be
Boil 175.27 ± 6.18Aa 118.00 ± 4.16Cb 112.75 ± 3.97Ab 31.03 ± 1.09Cc 29.72 ± 1.05Cc
Microwave 63.51 ± 2.24Cd 132.49 ± 4.67Bb 110.05 ± 3.88Ac 175.97 ± 6.2Aa 64.62 ± 2.28Ad
Stir-fry 17.6 ± 0.62Da 8.46 ± 0.30Db 3.85 ± 0.14Cc 0.72 ± 0.03Dd 0.81 ± 0.03Dd
1-Octen-3-ol 1343.8 ± 47.35 Steam 303.1 ± 10.68Ba 275.17 ± 9.7Bb 148.69 ± 5.24Bc 108.75 ± 3.83Bd 114.11 ± 4.02Bd
Boil 284.63 ± 10.03Ba 183.49 ± 6.47Cb 158.91 ± 5.6Bc 49.15 ± 1.73Ce 88.27 ± 3.11Cd
Microwave 1120.55 ± 39.48Aa 565.26 ± 19.92Ac 452.98 ± 15.96Ad 563.19 ± 19.84Ac 672.37 ± 23.69Ab
Stir-fry 76.07 ± 2.68Cb 53.22 ± 1.88Dc 38.55 ± 1.36Cd 97.94 ± 3.45Ba 77.25 ± 2.72Cb
Phenylethyl Alcohol 14.17 ± 0.5 Steam 9.77 ± 0.34Ac 18.36 ± 0.65Aa 12.13 ± 0.43Bb 12.43 ± 0.44Ab 9.44 ± 0.33Ac
Boil 3.47 ± 0.12Da 3.49 ± 0.12Da 3.12 ± 0.11Db 1.91 ± 0.07Cc 1.93 ± 0.07Cc
Microwave 8.26 ± 0.29Bc 8.08 ± 0.28Bc 15.50 ± 0.55Aa 10.55 ± 0.37Bb 2.26 ± 0.08Cd
Stir-fry 4.82 ± 0.17Cc 5.14 ± 0.18Cc 9.77 ± 0.34Ca 8.74 ± 0.31Db 8.26 ± 0.29Bb
ester
Acetic acid, hexyl ester ND Steam 3.93 ± 0.14Ca 2.54 ± 0.09Cb 1.52 ± 0.05Cc 1.35 ± 0.05Cd 1.62 ± 0.06Dc
Boil 9.71 ± 0.34Bd 14.64 ± 0.52Bb 22.18 ± 0.78Aa 9.04 ± 0.32Bd 10.83 ± 0.38Sc
Microwave 22.18 ± 0.78Aa 19.51 ± 0.69Ab 7.81 ± 0.28Bd 10.10 ± 0.36Ac 7.87 ± 0.28Bd
Stir-fry 4.21 ± 0.15Ca 2.41 ± 0.08Cc 1.49 ± 0.05Cd 1.60 ± 0.06Cd 3.96 ± 0.14Cb
Octyl chloroformate 11.41 ± 0.4 Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 3.16 ± 0.11d 3.76 ± 0.13c 3.83 ± 0.13c 8.12 ± 0.29a 4.88 ± 0.17b
Decanoic acid, 2,8-dimethyl-, methyl ester ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 0.62 ± 0.02c 0.42 ± 0.01d 0.81 ± 0.03b 0.20 ± 0.01e 0.89 ± 0.03a
d-Proline, N-methoxycarbonyl-, pentyl ester ND Steam 3.12 ± 0.11Bc 3.77 ± 0.13Ba 1.53 ± 0.05Cd 3.65 ± 0.13Ba 3.34 ± 0.12Ab
Boil 2.13 ± 0.08Cc 1.64 ± 0.06Cd 3.41 ± 0.12Ba 0.90 ± 0.03Be 3.01 ± 0.11Bb
Microwave 10.78 ± 0.38Ad 42.4 ± 1.49Ac 46.93 ± 1.65Ab 94.88 ± 3.34Aa 2.23 ± 0.08Ce
Stir-fry 1.67 ± 0.06Da 1.33 ± 0.05Cc 0.76 ± 0.03Ce 0.95 ± 0.03Bd 1.57 ± 0.06Db
2-Propyl-1-pentanol, chlorodifluoroacetate ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 1.08 ± 0.04c 1.50 ± 0.05b 0.94 ± 0.03d 1.57 ± 0.06b 1.74 ± 0.06a
amide
Acetamide, 2-fluoro- 13.84 ± 0.49 Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry ND ND ND ND ND
Azodicarbonamide ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 2.84 ± 0.10a 1.85 ± 0.07b 1.41 ± 0.05c 1.47 ± 0.05c 0.65 ± 0.02d
Hydrazinecarboxamide 29.61 ± 1.04 Steam 15.01 ± 0.53Dd 18.50 ± 0.65Cd 75.67 ± 2.67Ac 172.89 ± 6.09Ab 208.42 ± 7.34Aa
Boil 23.80 ± 0.84Bb 46.24 ± 1.63Ba 8.65 ± 0.30Dc 8.13 ± 0.29Dc 1.96 ± 0.07Cd
Microwave 34.59 ± 1.22Ad 277.22 ± 9.77Aa 63.12 ± 2.22Bc 86.82 ± 3.06Bb 1.20 ± 0.04Ce
Stir-fry 20.30 ± 0.72Cc 20.41 ± 0.72Cc 41.79 ± 1.47Cb 73.61 ± 2.59Ca 75.26 ± 2.65Ba
alkene
1-Hexene, 2,5,5-trimethyl- ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 13.49 ± 0.48d 19.02 ± 0.67c 39.73 ± 1.40a 34.67 ± 1.22b 33.33 ± 1.17b
1-Undecene, 4-methyl- ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 1.07 ± 0.04c 1.41 ± 0.05a 0.62 ± 0.02d 1.20 ± 0.04b 1.08 ± 0.04c
2,3-Dimethyl-1-hexene 83.78 ± 2.95 Steam 29.80 ± 1.05Bb 38.32 ± 1.35Ba 16.38 ± 0.58Cc 16.66 ± 0.59Cc 14.12 ± 0.50Dd
Boil 24.88 ± 0.88Cb 23.40 ± 0.82Cc 26.70 ± 0.94Ba 12.49 ± 0.44Dd 12.02 ± 0.42Cd
Microwave 117.61 ± 4.14Aa 82.19 ± 2.90Ab 76.52 ± 2.70Ac 73.27 ± 2.58Ac 30.28 ± 1.07Ad
Stir-fry 16.61 ± 0.59Dc 7.82 ± 0.28Dd 6.13 ± 0.22De 24.50 ± 0.86Ba 18.01 ± 0.63Bb
2-Decene, 8-methyl-, (Z)- ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 1.15 ± 0.04d 1.21 ± 0.04d 1.55 ± 0.05c 2.39 ± 0.08a 2.11 ± 0.07b
d-Limonene 104.62 ± 3.69 Steam 150.99 ± 5.32Ba 123.03 ± 4.34Cbc 105.46 ± 3.72Cd 118.35 ± 4.17Dc 128.98 ± 4.54Cb
Boil 183.50 ± 6.47Ab 233.89 ± 8.24Ba 188.39 ± 6.64Bb 185.98 ± 6.55Cb 143.91 ± 5.07Bc
Microwave 185.25 ± 6.53Ad 320.24 ± 11.28Ab 223.9 ± 7.89Ac 485.55 ± 17.11Aa 105.60 ± 3.72De
Stir-fry 192.02 ± 6.77Ac 115.63 ± 4.07Cd 93.57 ± 3.30De 269.83 ± 9.51Bb 287.39 ± 10.13Aa
ketone
2H-1,3-Benzoxazine-2,4(3H)-dione, 3-methyl- 13.83 ± 0.49 Steam 4.09 ± 0.14Cc 4.01 ± 0.14Bc 9.08 ± 0.32Ba 1.86 ± 0.07Bd 8.34 ± 0.29Ab
Boil 28.88 ± 1.02Aa 1.66 ± 0.06Cd 5.23 ± 0.18Cb 3.72 ± 0.13Bc 1.62 ± 0.06Dd
Microwave 17.77 ± 0.63Bc 27.44 ± 0.97Ab 16.49 ± 0.58Ac 59.7 ± 2.10Aa 4.59 ± 0.16Bd
Stir-fry 5.2 ± 0.18Ca 2.5 ± 0.09Ce 4.05 ± 0.14Db 3.08 ± 0.11Bc 2.76 ± 0.10Cd
2-Undecanone 21.57 ± 0.76 Steam 8.61 ± 0.3Ab 10.9 ± 0.38Aa 5.33 ± 0.19Ae 4.74 ± 0.17Ad 6.38 ± 0.22Ac
Boil 4.25 ± 0.15Bc 4.87 ± 0.17Bb 3.89 ± 0.14Bd 3.04 ± 0.11Ce 4.25 ± 0.15Bc
Microwave 6.16 ± 0.22Ba 3.98 ± 0.14Bc 4.61 ± 0.16Bb 4.43 ± 0.16Bb 1.29 ± 0.05Cd
Stir-fry 0.40 ± 0.01Cc 0.44 ± 0.02Cc 0.41 ± 0.01Cc 1.19 ± 0.04Da 0.76 ± 0.03Db
3-Heptanone, 2-methyl- ND Steam 2.85 ± 0.10Ce 3.48 ± 0.12Cd 5.09 ± 0.18Cb 4.80 ± 0.17Cc 5.39 ± 0.19Ca
Boil 12.41 ± 0.44Ba 11.08 ± 0.39Bb 9.90 ± 0.35Bc 9.35 ± 0.33Bc 12.41 ± 0.44Aa
Microwave 18.37 ± 0.65Ac 22.91 ± 0.81Aa 19.65 ± 0.69Ab 10.51 ± 0.37Ad 7.81 ± 0.28Be
Stir-fry 1.00 ± 0.04Dab 0.54 ± 0.02Dd 0.83 ± 0.03Dc 0.97 ± 0.03Db 1.03 ± 0.04Da
5,9-Undecadien-2-one, 6,10-dimethyl-, (E)- 12.57 ± 0.44 Steam 7.29 ± 0.26Bb 10.87 ± 0.38Aa 6.32 ± 0.22Cc 5.98 ± 0.21Bc 4.18 ± 0.15Ad
Boil 7.65 ± 0.27Ba 3.96 ± 0.14Cc 6.81 ± 0.24Bb 2.61 ± 0.09Ce 2.96 ± 0.10Bd
Microwave 11.72 ± 0.41Aa 9.29 ± 0.33Bb 7.97 ± 0.28Ac 8.32 ± 0.29Ac 4.01 ± 0.14Ad
Stir-fry 0.49 ± 0.02Cb 0.49 ± 0.02Db 0.27 ± 0.01Dd 0.55 ± 0.02Da 0.35 ± 0.01Cc
miscellaneous
Ethylbenzene ND Steam ND ND ND ND ND
Boil ND ND ND ND ND
Microwave ND ND ND ND ND
Stir-fry 28.44 ± 1.00b 3.19 ± 0.11e 5.11 ± 0.18d 8.11 ± 0.29c 29.89 ± 1.05a
Styrene 8.38 ± 0.3 Steam 8.87 ± 0.31Ca 4.33 ± 0.15Cb 4.13 ± 0.15Db 4.33 ± 0.15Cb 2.94 ± 0.10Cc
Boil 7.26 ± 0.26Dc 7.19 ± 0.25Bc 27.09 ± 0.95Ba 6.18 ± 0.22Bd 13.13 ± 0.46Bb
Microwave 34.90 ± 1.23Aa 34.27 ± 1.21Aa 30.60 ± 1.08Ab 31.93 ± 1.13Ab 1.14 ± 0.04Dc
Stir-fry 25.61 ± 0.9Bb 6.22 ± 0.22Bd 17.86 ± 0.63Cc 4.05 ± 0.14Ce 30.35 ± 1.07Aa
Toluene 16.81 ± 0.59 Steam 18.55 ± 0.65Be 25.09 ± 0.88Cd 39.83 ± 1.40Bb 30.27 ± 1.07Bc 48.40 ± 1.71Aa
Boil 15.45 ± 0.54Cd 93.95 ± 3.31Ba 66.27 ± 2.34Ab 27.70 ± 0.98Cc 12.19 ± 0.43Bd
Microwave 30.80 ± 1.09Ac 184.44 ± 6.5Aa 10.19 ± 0.36Cd 45.95 ± 1.62Ab 3.18 ± 0.11De
Stir-fry 6.15 ± 0.22Dd 7.32 ± 0.26Dc 5.71 ± 0.20Dd 10.41 ± 0.37Da 9.03 ± 0.32Cb
Oxime-, methoxy-phenyl-_ 14.84 ± 0.52 Steam 93.38 ± 3.29Ba 22.72 ± 0.80Cb 6.22 ± 0.22Bc 4.42 ± 0.16Cc 4.13 ± 0.15Cc
Boil 181.86 ± 6.41Aa 36.78 ± 1.30Ac 98.67 ± 3.48Ab 27.70 ± 0.98Bd 16.42 ± 0.58Ae
Microwave 22.91 ± 0.81Cc 26.13 ± 0.92Bb 3.83 ± 0.13Be 30.64 ± 1.08Aa 10.51 ± 0.37Bd
Stir-fry 0.27 ± 0.01Db 0.19 ± 0.01Dc 0.17 ± 0.01Cd 0.17 ± 0.01Dd 0.36 ± 0.01Da
2,4-Pentanedione, ion(1-), lithium ND Steam 5.30 ± 0.19Ca 3.33 ± 0.12Cd 3.02 ± 0.11Ce 3.96 ± 0.14Cc 4.82 ± 0.17Bb
Boil 6.41 ± 0.23Cb 14.21 ± 0.5Ba 3.46 ± 0.12Cd 3.17 ± 0.11Dd 5.18 ± 0.18Bc
Microwave 34.90 ± 1.23Aa 24.47 ± 0.86Ab 25.71 ± 0.91Ab 12.82 ± 0.45Ac 2.20 ± 0.08Cd
Stir-fry 28.44 ± 1.00Bb 3.19 ± 0.11Ce 5.11 ± 0.18Bd 8.11 ± 0.29Bc 29.89 ± 1.05Aa
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The values of different letters in each line are considered to be significantly different at P < 0.05. Capital letters represent the comparison of differences in different cooking methods, and lowercase letters represent the comparison of differences in different cooking time. ND: Not detected.

In the aldehyde compounds, hexanal and heptanal can provide a grassy odor (Zorrilla & Evidente, 2022), while decanal and nonanal provide peel aroma. These aldehydes are common volatiles in mushrooms. As shown in Table 2, in the uncooked sample, the contents of hexanal and heptanal were the highest, at 39.46 μg/kg and 28.46 μg/kg respectively, significantly higher than those in samples obtained from various cooking methods (P < 0.05). In SC-10 min, the contents of hexanal and nonanal reached the highest values, at 31.24 μg/kg and 86.23 μg/kg respectively, and then significantly decreased with the extension of cooking time (P < 0.05). Additionally, in SC-10 min, the contents of decanal and benzeneacetaldehyde also increased significantly, to 15.44 μg/kg and 36.39 μg/kg respectively, and then the contents of decanal and nonanal decreased significantly (P < 0.05).

Alcohols mainly originated from amino acid metabolism and the oxidation of unsaturated fatty acids, exhibiting higher odor thresholds (Hu et al., 2024). Among the detected alcohols, 1-octen-3-ol, responsible for a strong mushroom and grassy odor, was a major contributor to mushroom aroma (Hiraide et al., 2004). In uncooked sample, the content of 1-octen-3-ol was the highest among the detected alcohols, at 1343.8 μg/kg, indicating it was the predominant alcohol in Grifola frondosa. However, after the four cooking treatments, the concentration of 1-octen-3-ol dropped significantly (P < 0.05). This finding was consistent with the research of Huang et al. (2022). For the same cooking duration, the content of 1-octen-3-ol after microwaving cooking was significantly higher than other three cooking methods (P < 0.05), suggesting microwaving cooking better preserves this compound. 1-octanol and phenethyl alcohol provide rose aroma. Their concentrations peaked in SC-10 min, at 191 μg/kg and 18.36 μg/kg respectively.

The flavor of mushrooms was also affected by alkanes, but their contribution is relatively weak (Li et al., 2019). In esters, ethyl acetate was undetectable in uncooked samples but emerged after cooking, reaching a peak concentration of 22.18 μg/kg in MC-5 min. This may be due to the oxidation reaction of aldehydes (Lu et al., 2021). In amides, Acetamide, 2-fluoro- was detected in uncooked samples at 13.84 μg/kg but was undetectable in the four cooking samples.

In summary, the results of HS-SPME-GC–MS analysis showed that the concentrations of various key volatile compounds (such as hexanal, nonanal, 1-octen-3-ol) reached or approached the peak under the corresponding cooking methods at the 10-min cooking time point, which provided a direct chemical basis for the conclusion that the sensory quality of 10-min cooking samples was better in sensory evaluation.

3.3. HS-GC-IMS analysis of volatile compounds in Grifola frondosa under different cooking methods

Based on the determination that Grifola frondosa exhibit optimal flavor performance after 10 min cooking through sensory evaluation and HS-SPME-GC–MS analysis, we selected uncooked samples and four other samples cooked for 10 min for HS-GC-IMS analysis. This method aimed to gain a deeper understanding of the changes in volatile compounds in Grifola frondosa under various cooking conditions. The data of volatile compounds under different cooking methods were first visualized in the form of 3D topographic maps (Fig. 1a). Among them, A: uncooked sample; B: microwaving sample; C: stir-frying sample; D: steaming sample; E: boiling sample. Fig. 1b showed a 2D top view and Fig. 1c a difference spectrum. In the 2D map, the background was blue, white indicated low substance concentration, and red indicates high concentration, with darker red representing higher concentration. In the different spectrum, the spectrum of uncooked sample A1 was used as a reference, and the spectra of Grifola frondosa sample processed by different cooking methods were subtracted from this reference. If two flavor compounds were the same before and after cooking, the point after subtraction is white. Red areas indicated increased substance concentration relative to the uncooked samples, and blue areas indicated decreased concentration. The results showed that the total amount and content of flavor compounds decreased after the four cooking methods. As shown in Fig. 1b, most signals fall within a retention time range of 100–600 s, and in the top view, the migration time between 1.0 and 1.8 showed significant variation. According to Fig. 1c, compared with uncooked sample, many blue spots were found in the range of 1.0–1.8 after four cooking methods, indicating that several main substances in Grifola frondosa were very sensitive to temperature and were easy to degrade or decompose, resulting in some signals disappeared or decreased after cooking. However, the signals of a few compounds increased, suggesting that these cooking methods can generate new compounds or increase the concentration of existing ones.

Fig. 1.

Fig. 1

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3D topographic map and two-dimensional map of volatile organics compounds of Grifola frondosa with different cooking methods for 10 min. A: Grifola frondosa uncooked sample; B: Grifola frondosa microwaving cooking 10 min sample; C: Grifola frondosa stir-frying cooking 10 min sample; D: Grifola frondosa steaming cooking 10 min sample; E: Grifola frondosa boiling cooking 10 min sample. (a) 3D topographic map of volatile organics compounds from Grifola frondosa with peak spectra. The ordinate represents the retention time, and the abscissa represents the drift time. (b) HS-GC-IMS two-dimensional top view of the spectra of Grifola frondosa samples. (c) HS-GC-IMS differential spectra of HS-GC-IMS samples of Grifola frondosa sample spectra.

3.4. Fingerprint analysis of volatile compounds in Grifola frondosa under different cooking methods

Fingerprinting can comprehensively show changes in the components or content of compounds in Grifola frondosa samples under different cooking methods. “M” and “D” denoted the monomer and dimer of this volatile organic compound, respectively.

The Fig. 2 demonstrated that in the uncooked samples, the main volatile compounds of Grifola frondosa were focused on aldehydes, esters, and ketones, including benzaldehyde, n-decanal, 3-octanone, 1-octen-3-ol, etc. Among these, 1-octen-3-ol was the most characteristic mushroom aroma.

Fig. 2.

Fig. 2

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Gallery plot plot of volatile organics compounds of Grifola frondosa with different cooking methods for 10 min. A: Grifola frondosa uncooked sample; B: Grifola frondosa microwaving cooking 10 min sample; C: Grifola frondosa stir-frying cooking 10 min sample; D: Grifola frondosa steaming cooking 10 min sample; E: Grifola frondosa boiling cooking 10 min sample.

Compared to the uncooked sample, the microwaving Grifola frondosa sample showed no significant difference in the content of major volatile compounds. It implied that microwaving cooking can effectively preserve the original flavor of mushroom and may result in the highest flavor acceptability. In the stir-frying sample, the content of certain volatile compounds, e.g. 3-octanone, octanal, (E)-2-octenal, heptanal, butyl formate, benzaldehyde, trans-2-butenoic acid ethyl ester, 3-methyl-2-pentanone, 1-propanethiol, and 3-(methylthio) propanal, was significantly lower than in the uncooked sample. These compounds were associated with nutty, grassy, and oily odors. Similarly, in the steaming sample, the content of these volatile compounds were also significantly reduced. However, unlike the stir-frying sample, the steaming sample contained 2-pentylfuran, which might be generated due to the Maillard reaction under high temperature conditions. In the boiling sample, except for isovaleric acid methyl ester (with a chestnut-like aroma) and 1,4-cineole (a spice), the content of other volatile compounds was significantly lower than in the uncooked sample.

3.5. Quantitative analysis of volatile compounds in Grifola frondosa by HS-GC-IMS

As shown in Table 3, a total of 27 volatile compounds were identified using HS-GC-IMS and the NIST database with retention indices. These included 4 ketones, 8 aldehydes, 5 esters, 2 alcohols, 3 ethers and 5 other compounds. Different cooking conditions differently affected the mushroom volatile compounds. Some compounds with high proton affinity formed dimers during cooking (Arce et al., 2014).

Table 3.

GC-IMS analysis of volatile components of Grifola frondosa by different cooking methods for 10 min.

Count
 Compound
 MW
 RI
 Peak Volume (a.u.)
uncooked sample microwaving sample stir-frying sample steaming cooking boiling sample
1 Propanal 58.1 507.4 1752.90 ± 80.75a 1655.47 ± 156.70a 963.98 ± 234.41c 1259.89 ± 76.69b 288.95 ± 12.53d
2 n-Octanal 128.2 1005 214.08 ± 11.15a 214.49 ± 95.49a 96.80 ± 48.97b 68.79 ± 29.51b 52.46 ± 3.19b
3 n-Nonanal 142.2 1102.4 1457.79 ± 53.17a 1483.76 ± 115.81a 942.61 ± 180.38b 1269.17 ± 480.88ab 326.89 ± 31.80c
4 Isopentyl formate 116.2 787 1441.38 ± 117.31b 1443.85 ± 30.21b 1886.09 ± 240.12a 1880.88 ± 252.99a 1083.70 ± 149.14c
5 Hexanenitrile 97.2 883.8 344.66 ± 12.90a 353.25 ± 77.30a 155.09 ± 91.66b 182.98 ± 116.51b 39.91 ± 8.37b
6 Heptanal(M) 114.2 896.9 2554.16 ± 113.35a 2208.77 ± 36.71b 1266.37 ± 108.88c 1367.91 ± 219.54c 539.60 ± 73.37d
7 Heptanal(D) 114.2 895.8 1865.92 ± 176.12a 1213.24 ± 186.05b 291.02 ± 121.86c 327.22 ± 193.18c 102.88 ± 5.07c
8 ethyl trans-2-butenoate 114.1 838.2 2184.74 ± 145.09a 1715.52 ± 20.94b 869.88 ± 56.05c 558.09 ± 37.16d 541.98 ± 36.22d
9 Ethyl butanoate 116.2 781.2 5073.59 ± 280.63a 5149.11 ± 456.4a 3570.99 ± 244.42b 3199.76 ± 222.97b 460.82 ± 77.56c
10 1-Octen-3-ol 134.2 989.8 6542.78 ± 239.44b 8573.05 ± 275.77a 4134.50 ± 1114.66c 2947.66 ± 362.52c 1143.6 ± 180.57d
11 Butyl formate(M) 102.1 731.1 239.40 ± 68.72a 168.32 ± 90.14a 268.56 ± 94.26a 185.54 ± 106.27a 92.26 ± 24.62a
12 Butyl formate(D) 102.1 728.6 248.89 ± 171.52a 144.27 ± 104.71a 142.07 ± 30.62a 98.86 ± 48.66a 51.46 ± 3.44a
13 Benzene, propyl- 120.2 957.6 211.67 ± 7.17a 123.84 ± 53.32abc 133.44 ± 87.80ab 62.30 ± 39.42bc 27.60 ± 6.48c
14 Benzaldehyde(M) 106.1 964.5 5251.13 ± 40.02a 5083.13 ± 103.92a 2272.81 ± 230.59b 2162.05 ± 342.41b 813.98 ± 95.57c
15 Benzaldehyde(D) 106.1 963.8 2457.60 ± 202.74a 1981.50 ± 287.1b 216.59 ± 39.78c 285.26 ± 121.85c 122.08 ± 30.05c
16 acetic acid hexyl ester 144.2 1011.4 732.54 ± 54.33b 813.42 ± 51.03ab 910.61 ± 61.88a 921.41 ± 133.82a 270.56 ± 50.31c
17 3-(Methylmercapto)propionaldehyde 104.2 904.2 250.79 ± 101.76a 281.67 ± 132.68a 61.52 ± 22.46b 63.24 ± 32.09b 30.65 ± 1.63b
18 2-phenylacetaldehyde 120.2 1046.5 1714.99 ± 224.73a 1553.82 ± 157.80a 672.02 ± 180.83b 489.82 ± 149.46bc 190.92 ± 15.52c
19 2-Pentyl furan 138.2 994 471.85 ± 69.10c 390.59 ± 17.33c 606.26 ± 28.79c 2046.92 ± 261.32a 909.31 ± 122.57b
20 2-pentanone, 3-methyl 100.2 752.3 1406.87 ± 83.25a 1071.36 ± 45.60b 366.51 ± 39.52c 356.88 ± 31.11c 224.71 ± 9.9d
21 2-Methoxy-2-methylpropane 88.1 549.8 2288.00 ± 127.88b 2407.92 ± 113.37b 2665.18 ± 152.35a 1693.06 ± 78.75c 647.68 ± 48.99d
22 2-Butanone(M) 72.1 589.1 769.84 ± 14.20a 604.59 ± 82.48b 772.20 ± 62.44a 710.19 ± 38.29a 637.53 ± 60.85b
23 2-Butanone(D) 72.1 580.3 2609.77 ± 265.48ab 3125.32 ± 554.28a 2044.49 ± 109.34b 1273.85 ± 165.26c 460.07 ± 65.30d
24 2-Acetyl-5-methylfuran 124.1 1032.8 45,178.79 ± 560.31a 44,789.40 ± 454.90a 24,525.17 ± 7769.60b 19,150.29 ± 185.44b 11,319.11 ± 1359.51c
25 2,3-Pentanedione 100.1 710 27.60 ± 5.59ab 32.47 ± 14.84a 23.10 ± 17.53abc 6.30 ± 2.87bc 3.45 ± 0.22c
26 1-Propanethiol 76.2 601.7 644.77 ± 10.50a 529.36 ± 183.36a 257.66 ± 65.11b 112.63 ± 24.58b 142.6 ± 11.93b
27 1-Butanol 74.1 657.4 16,142.55 ± 447.76a 16,099.24 ± 738.26a 10,245.24 ± 209.45b 9311.79 ± 223.69c 2606.23 ± 305.66d
28 1,4-Cineol(M) 154.3 1013.9 622.60 ± 7.02b 495.70 ± 21.50b 375.75 ± 125.07b 421.34 ± 48.88b 3389.52 ± 477.61a
29 1,4-Cineol(D) 154.3 1001.2 2185.79 ± 215.8b 1813.28 ± 304.65b 543.11 ± 507.98c 571.57 ± 129.72c 9011.41 ± 734.52a
30 1,2-Dimethoxyethane 90.1 625.6 122.21 ± 8.64b 101.01 ± 21.22b 190.01 ± 54.10a 142.37 ± 32.60ab 35.89 ± 1.39c
31 3-Octanone 136.2 991.5 4035.77 ± 108.58a 2386.11 ± 164.24b 688.07 ± 442.97c 709.03 ± 105.60c 451.80 ± 173.03c
32 (E)-2-Octenal(M) 126.2 1065.1 8014.26 ± 58.73a 6959.65 ± 673.88b 2090.40 ± 698.73c 1603.83 ± 15.57c 1284.00 ± 33.98c
33 (E)-2-Octenal(D) 126.2 1061.1 4367.16 ± 549.99a 3112.67 ± 645.23b 335.06 ± 105.74c 251.41 ± 26.49c 257.17 ± 56.26c
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The values of different letters in each line are considered to be significantly different at P < 0.05.

Based on the comparative analysis of HS-GC-IMS and HS-SPME-GC–MS results, the single method detected 36 and 27 volatile compounds, respectively, with a total of 57 volatile compounds identified by both methods combined. The compounds commonly detected by both techniques were benzaldehyde, benzeneacetaldehyde, heptanal, nonanal, 1-octen-3-ol, and acetic acid hexyl ester, suggesting that these substances may be present at relatively high concentrations in Grifola frondosa. Additionally, HS-GC-IMS detected three ether compounds, which were not identified by HS-SPME-GC–MS; however, HS-SPME-GC–MS detected three amides, five alkenes, and five alcohols, whereas HS-GC-IMS only detected two alcohols and failed to detect amides and alkenes. These findings indicate that HS-SPME-GC–MS demonstrates superior performance in detecting alcohols, alkenes, and nitrogen-containing compounds, which is consistent with the conclusions reported by Zhong et al. (2024). Compared with HS-SPME-GC–MS, HS-GC-IMS is capable of detecting both monomers and dimers of compounds. Therefore, the combined application of HS-GC-IMS and HS-SPME-GC–MS is highly necessary.

In HS-GC-IMS analysis, some volatile compounds were detected in the form of monomer (M) and dimer (D). The signal intensity ratio can indirectly reflect the concentration level of flavor substances and the strength of intermolecular interactions. As shown in Table 3, for heptanal and benzaldehyde, the intensity of M peak is generally higher than that of D peak, indicating that it mainly exists in the form of highly volatile free monomer in the headspace. This distribution characteristic means that the flavor substance tends to be released instantaneously and has strong olfactory explosive power. However, due to the relatively small dimer reserves, its aroma persistence is weak and easy to decay rapidly. In contrast, for 2-butanone and compounds, the intensity of D peak after cooking treatment was significantly greater than that of M peak. This phenomenon may be attributed to the fact that the cooking process promotes the release or formation of the compound, resulting in a significant increase in its concentration in the headspace, which makes it easier to form dimers through hydrogen bond association. According to the detection principle of ion mobility spectroscopy, larger dimer ions produce stronger signal response (Kanu & Hill Jr, 2008). Therefore, the strong D peak not only indicates a high concentration, but also means a dynamic flavor reservoir. This distribution pattern partially explains the more persistent and subtle characteristics of flavor in cooking samples.

Aldehydes plays a crucial role in mushrooms (Jiang et al., 2023) and have intense, overlapping flavor effects (Hou et al., 2021). According to the data listed in Table 3, compared with the uncooked sample, six aldehydes showed no significant difference in the MC-10 min sample, indicating that these six aldehydes were well retained after 10 min of microwave cooking. In SC-10 min, SFC-10 min and BC-10 min sample, except for n-Nonanal, the other aldehydes were significantly reduced (P < 0.05). Aldehydes are often produced through Strecker degradation and fatty acid oxidation (Lv et al., 2024). Microwaving cooking can better retain the six aldehydes, probably because microwave cooking rapidly heats up the interior of the sample and shortens the heat treatment time, thereby reducing the loss of aldehydes. In steaming and boiling cooking, water-soluble aldehydes may be lost with steam or water, resulting in a decrease in aldehydes. In stir-frying cooking, high temperature and oxygen promote the rapid oxidation of fatty acids, resulting in the decomposition or conversion of original aldehydes into other compounds. At the same time, the newly generated aldehydes may mask the original aldehydes, but the data show that most aldehydes are significantly reduced, indicating that the loss of aldehydes is greater than the generation.

Esters, common in fruits and known for their fruity aromas (Wang et al., 2021), were detected in multiple varieties in the Grifola frondosa samples. Their formation is closely related to lipid oxidation and alcohol esterification reactions (Zhao et al., 2024). According to the results in Table 3, the MC-10 min, SFC-10 min, and SC-10 min samples each retained three types of esters relatively well, whereas only one type of ester was well retained in the BC-10 min sample. In the MC-10 min sample, ethyl butyrate, which imparts a pineapple flavor, exhibited the highest peak area of 5149.11. The retention of esters in microwaving cooking may be attributed to the relatively low power, which limits steam flow, combined with good sealing. Additionally, during stir-frying cooking, the involvement of oils in lipid oxidation and alcohol esterification reactions promotes the formation of esters. In steaming cooking, the retention of esters may be because the detected esters are insoluble in water, thus avoiding a large loss of esters. Therefore, esters are also well retained in these two cooking methods. For example, the content of isoamyl formate in the SFC-10 min and SC-10 min samples was significantly higher than in other samples (P < 0.05), with values of 1886.09 and 1880.88, respectively. Regarding acetic acid hexyl ester, its content in the MC-10 min, SFC-10 min, and SC-10 min samples was significantly higher than BC-10 min sample, with the BC-10 min showing the lowest value of 270.56. This also reflects that microwaving, stir-frying and steaming cooking can better reduce the loss of the substance.

Ketones, which can form via amino acid thermolysis or polyunsaturated fatty acid oxidation (Du, Chen, et al., 2021), were only detected in three types in the Grifola frondosa samples. In MC-10 min, the concentration of 2,3-pentanedione rose to 32.47. As a product of the Maillard reaction, 2,3-pentanedione contributes caramel and nutty flavors. The content of 3-methyl-2-pentanone was 1406.87 in the uncooked samples and 1071.36 in MC-10 min, both significantly higher than in the other three cooking samples (P < 0.05). The lowest concentration of the substance in the BC-10 sample was 224.71, which further confirmed that boiling would lead to the loss of volatile substances. 3-methyl-2-pentanone in the sample is primarily a product of thermal oxidation of unsaturated fatty acids, contributing fresh fruity and grassy aromas (Zhu et al., 2024).

Among the four cooking methods, the content of 1-octen-3-ol in MC-10 min was the highest, which was 8573.05, and was significantly higher than that of stir-frying, steaming and cooking samples (P < 0.05). It was due to the presence of unsaturated double bonds in 1-octen-3-ol, which maked it chemically unstable and prone to decomposition during cooking (Peng et al., 2022). This result was similar to those of Du, Sissons, et al. (2021) and Ashmore et al. (2014). Probably because microwaving cooking promoted the release of 1-octen-3-ol, and microwaving cooking was carried out under relatively sealed conditions, reducing the volatilization loss.

2-Pentylfuran is a furan compound with a nutty or grassy flavor (Liu et al., 2014), which is commonly found in heated foods and can be generated through Maillard reaction and lipid oxidation (Ouyang et al., 2024). In SC-10 min, the concentration of 2-pentylfuran in Grifola frondosa samples was 2046.92, significantly higher than in the other cooking samples (P < 0.05). It indicated that the hot and humid environment may be conducive to lipid oxidation to produce 2-pentylfuran. 1,4-Cineol, a monoterpene with a mild camphor odor and a cool, spicy note, showed the highest concentration in the boiling sample after 10 min cooking, also significantly higher than in the other samples (P < 0.05).

Additionally, this study indicated that boiling cooking can lead to a reduction in the volatile compounds of fresh Grifola frondosa. This is similar to previous studies (Tian et al., 2016; Wu et al., 2026). In BC-10 min, apart from 1,4-cineole and isovaleric acid methyl ester, the remaining volatile compounds showed a significant decrease. This might be due to the degradation of flavor components during the heating process, which also explained why the boiling sample had the weakest flavor performance. In contrast, the volatile compounds were better preserved in the uncooked sample and the sample subjected to MC-10 min. It suggested that MC-10 min could effectively retain the original flavor profile of Grifola frondosa. For steaming cooking and stir-frying cooking, although the volatile compounds were weaker than those of microwaving cooking sample, they could also contribute special flavors to Grifola frondosa. For example, acetic acid hexyl ester and isoamyl formate were the highest in the SC-10 min and SFC-10 min, respectively, which could contribute a light fruit flavor to Grifola frondosa.

3.6. PCA analysis of different cooking methods

Typically, a PCA model is chosen for reporting when the contribution rate reaches 60% (Chen et al., 2020). In the PCA plot (Fig. 3B), the PCA results show that PC1 and PC2 account for 84% (65% and 19%) of the total variance, indicating that HS-GC-IMS can explain most volatile compound information. Overall, the uncooked and microwaving samples had similar principal components, as do the stir-frying and steaming samples. However, the boiling samples differ significantly from the others, showing that cooking methods significantly affect Grifola frondosa quality.

Fig. 3.

Fig. 3

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(a) PCA (principal component analysis) diagram of volatile organic compounds in Grifola frondosa samples with different cooking methods for 10 min. (b) The nearest neighbor fingerprint of volatile organic compounds in Grifola frondosa samples with different cooking methods for 10 min.

In the nearest neighbor fingerprint analysis, the intensity of compounds in a specified region was used to rapidly compare samples and compute the Euclidean distance between each pair of samples (Zhang et al., 2022). This method enabled the analysis of volatile chemical similarity. Fig. 3b presented the nearest neighbor Euclidean distance plot between the control and the four cooking method samples. A shorter distance signifies higher similarity, whereas a longer distance implies greater variability. As shown in the figure, the boiling sample was the farthest from the uncooked sample, followed by the stir-frying and steaming samples, indicating significant variability in volatile compounds. In contrast, the microwaving sample was the closest to the uncooked samples, suggesting less variability in volatile compounds. This finding was consistent with the PCA results. Overall, the results demonstrated that the volatile compounds of Grifola frondosa samples subjected to different cooking methods underwent significant changes.

4. Conclusion

This study systematically compared the effects of four cooking methods on the volatile flavor compounds of Grifola frondosa, integrating sensory evaluation with HS-SPME-GC–MS analysis to identify 10 min as the optimal cooking duration. HS-GC-IMS further revealed that aldehydes and esters serve as the primary contributors to its flavor profile. Microwave cooking (MC-10 min) best preserved the original flavor compounds while significantly enhancing the content of typical mushroom-flavor substances, such as 1-octen-3-ol. Additionally, stir-frying and steaming exhibited distinct flavor characteristics: stir-frying enhanced plum-like aroma (associated with isoamyl formate), whereas steaming promoted fruity notes (e.g., acetic acid hexyl ester and 2-pentylfuran). Boiled samples displayed the weakest flavor intensity. These findings offer a scientific approach to optimizing the cooking and sensory quality of Grifola frondosa. In the future, we can further explore the synergistic improvement of the overall flavor acceptance of Grifola frondosa by compound cooking or seasoning. Additionally, the study primarily assessed volatile compounds and sensory attributes, without evaluating potential changes in nutritional components or bioactive properties during cooking, which could also influence the overall quality and health value of Grifola frondosa.

CRediT authorship contribution statement

Zitong Liu: Writing – review & editing, Writing – original draft. Jian Zhao: Writing – original draft, Data curation. Ziheng Cheng: Software, Methodology. Xinfang Zhang: Software, Methodology. Meifan Pan: Formal analysis, Data curation. Beilei Pang: Formal analysis, Data curation. Ruiqi Zhang: Funding acquisition, Formal analysis. Piaopiao Wei: Validation, Methodology. Xuemei Cheng: Validation, Methodology. Yudong Wang: Methodology, Writing – review & editing. Yongye Sun: Validation, Resources, Conceptualization. Lirong Xu: Supervision, Funding acquisition, Conceptualization.

Ethical statement

The content of the sensory evaluation experiment has been informed consent of the candidates and has been carried out in accordance with The Code of Ethics of the World Medical Association (Declaration of Helsinki). Research protocols of sensory evaluation by panelists were approved by the Qingdao University Ethics Committee.

Declaration of competing interest

The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.

Acknowledgement

National Natural Science Foundation of China (Project No. 32202083), Open Research Fund of the Key Laboratory of Edible Oil Quality and Safety, State Administration for Market Regulation (SYYKF202501), Shandong Province Natural Science Foundation (ZR2024QC124), Young Talent of Lifting engineering for Science and Technology in Shandong, China (SDAST2025 QTA083), China Postdoctoral Science Foundation (Project No. 2023M731849), Qingdao Postdoctoral Innovation Project (Project No. QDBSH20220202216).

Data availability

Data will be made available on request.

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Data Availability Statement

Data will be made available on request.

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