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. 2026 Mar 3;35:103717. doi: 10.1016/j.fochx.2026.103717

Revealing the changes in lipids, volatile flavors, sensory effects and their correlations in cold fresh rainbow trout meat under different electron beam irradiation doses using lipidomics and flavoromics

Fanyi Gong a, Buzhou Xu a, Li Yang a,b, Chang Su a,b, Jiaxin Chen a,b, Xingzhong Zhang a,b, Yong Yu d, Shichen Zhu c, Jie Tang a,b, Dong Zhang a,b,⁎, Hongjun Li a,d
PMCID: PMC12972702  PMID: 41816766

Abstract

This study investigated the effects of electron beam irradiation at doses of 1, 3, 5, and 7 kGy on cold fresh rainbow trout meat, using non-irradiated samples as a control. Changes in sensory scores, volatile flavor compounds, and lipids were analyzed using comprehensive two-dimensional gas chromatography–mass spectrometry (GC × GC–MS) and untargeted lipidomics. The results showed that irradiation at doses ≥3 kGy significantly reduced sensory scores. GC × GC–MS analysis indicated an increase in the total content of volatile flavor compounds after irradiation. Specifically, at doses ≥3 kGy, levels of lipid oxidation products—such as hexanal, (R)-2-octanol, and (S)-2-octanol—increased significantly (P < 0.05). Lipidomics analysis identified 583 differential lipids across treatment groups, revealing decreases in phosphatidylcholine, triglyceride, and diglyceride levels, alongside an increase in phosphatidylethanolamine following irradiation. These findings demonstrate that electron beam irradiation promotes volatile compound formation by accelerating lipid oxidation. While doses below 3 kGy did not substantially alter flavor or sensory quality, doses of 3 kGy or higher induced a distinct “irradiation odor”, ultimately leading to a marked reduction in acceptability.

Keywords: Cold fresh rainbow trout meat, electron beam irradiation, Sensory scores, Volatile flavor compounds, Lipidomics analysis

Highlights

  • •

    PC, TG, DG and PE were the main lipid components in cold fresh rainbow trout meat.

  • •

    An irradiation of ≥3 kGy significantly increased unpleasant lipid oxidation products.

  • •

    LPC(22:4/20:4/18:2/20:3) and LPE(18:3) caused flavor changes in irradiated samples.

  • •

    The optimal electron beam irradiation dose should be maintained below 3 kGy.

1. Introduction

Cold fresh rainbow trout meat is highly favored by consumers worldwide due to its tender texture and high content of proteins and unsaturated fatty acids (Cengiz, Guclu, Kelebek, & Selli, 2023). However, during storage and transportation, cold fresh rainbow trout meat is susceptible to microbial contamination, oxidation, and other factors, leading to quality deterioration and flavor changes (Mozaffari et al., 2022). To extend its shelf life, various preservation technologies have been developed. Among them, electron beam irradiation, as an efficient non-thermal food sterilization method, has shown significant advantages in prolonging product shelf life and ensuring food safety (Gautam & Venugopal, 2021). However, the interaction between high-energy electrons and lipid molecules during irradiation may trigger oxidative cascades, altering flavor precursors in the muscle matrix (Zhang et al., 2023). In current industrial applications, the selection of electron beam irradiation doses is often primarily guided by sterilization efficacy, with insufficient consideration of its impact on volatile flavor compounds and the underlying lipid components. This oversight may ultimately compromise product acceptability and market competitiveness.

The dose of electron beam irradiation is a critical parameter that determines the balance between microbial safety and physicochemical quality. A review of previous studies reveals a spectrum of effects across different dose ranges. At low doses (typically 1–3 kGy), significant microbial reduction can be achieved with relatively minimal impact on quality attributes. For example, Xu et al. (2021) found that relatively low doses of electron beam irradiation (1–3 kGy) significantly extended the shelf life of weever (Lateolabrax japonicus) fillets while preserving their hardness and chewiness. Jeyakumari et al. (2023) reported that electron beam irradiation at doses ranging from 0 to 4.0 kGy notably reduced the counts of Pseudomonas and Brochothrix thermosphacta in tilapia meat, and treatments at 2–4 kGy effectively prolonged the shelf life of tilapia without significantly compromising its quality. At medium doses (around 3–5 kGy), sterilization is more thorough, but the lipid oxidation and related flavor changes often become detectable. Li, Yu, Xiong, Liao, and Zu (2020) observed that increasing irradiation doses (from 0 to 6 kGy) elevated acid value, peroxide value, and thiobarbituric acid reactive substances value in Micropterus salmoides meat, alongside increased free fatty acid content and elevated primary/secondary lipid oxidation products. At higher doses (≥5–7 kGy and above), while ensuring microbial safety, the risk of pronounced lipid oxidation, protein denaturation, and consequent deterioration in texture, color, and flavor increases substantially. For example, Yu et al. (2023) reported that 7 and 10 kGy electron beam irradiation induced significant lipid oxidation and texture changes in Atlantic salmon. Existing literature has systematically explored the effects of different doses of electron beam irradiation on microbial inactivation and changes in traditional quality indicators in aquatic products, as well as the evolution of these indicators during storage. However, there is relatively limited research on the dose-dependent molecular-level changes, particularly regarding the immediate impact of electron beam irradiation-mediated alterations in lipid composition on the volatile flavor compounds in cold fresh rainbow trout meat. Systematic conclusions in this area have yet to be established. Therefore, this study designed a dose-gradient scheme ranging from no treatment (0 kGy control group) to low dose (1 kGy), medium doses (3 kGy, 5 kGy), and up to a relatively high dose (7 kGy). This dose range was selected to identify the thresholds for lipid oxidation and flavor changes and to establish a dose–response relationship. The primary objective is to elucidate how different doses of electron beam irradiation mediate changes in lipid composition and thereby regulate the volatile flavor profile of cold fresh rainbow trout meat.

Therefore, untargeted lipidomics and comprehensive two-dimensional gas chromatography–mass spectrometry (GC × GC–MS) were employed to systematically investigate the changes in lipid composition as well as the composition and content of volatile flavor compounds in rainbow trout meat treated with different electron beam irradiation doses (0, 1, 3, 5, and 7 kGy). Additionally, the effects of different irradiation doses on sensory scores, color, and the extent of lipid oxidation (thiobarbituric acid reactive substance, TBARS) in rainbow trout meat were examined, and the inherent relationships between alterations in lipid composition and changes in volatile flavor compounds were thoroughly analyzed. This study not only elucidates the mechanism by which electron beam irradiation doses immediately impact the flavor characteristics of cold fresh rainbow trout meat through lipid metabolism but also provides a theoretical basis for optimizing irradiation processes and offers data support for establishing flavor-friendly irradiation standards.

2. Materials and methods

2.1. Materials and irradiation treatment

Live rainbow trout were purchased from Pengzhou Yongquan Cold-water Fisheries Co., Ltd. and then processed by the company staff using commercial standard procedures (removal of head, bones, and skin, the trout had been farm-raised for two years). Upon arrival at the laboratory, the trout meat was sliced into 0.7 cm–thick fillets (approximately 15 g each). The freshly sliced fillets were randomly divided into five groups. Samples for each group were placed on trays, sealed in sterile polyethylene composite bags to isolate them from the environment, and transported at 4 °C in a refrigerated container to Zhongjin Irradiation Chengdu Co., Ltd. Electron beam irradiation was performed using an IS1024 irradiator at room temperature (approximately 15 °C; maximum energy, 10 MeV; beam current: 2.1 mA; 5-step scan width; the belt speed was set at 150, 100, 60, and 45 mm/s for irradiation doses of 1, 3, 5, and 7 kGy, respectively) according to the reported method (Wang et al., 2023). The control group (DZS0) received no irradiation, while the experimental groups were irradiated at target doses of 1, 3, 5, and 7 kGy, denoted as DZS1, DZS3, DZS5, and DZS7, respectively. The entire irradiation procedure was repeated three times independently. After irradiation, the fillets were transported back to the laboratory in insulated containers with ice packs and stored appropriately until subsequent analysis.

2.2. Sensory evaluation

A sensory panel comprising ten trained assessors (five males and five females) was convened following a published method (Yao et al., 2024). Cold fresh rainbow trout meat from the different treatment groups was presented to the panelists on randomly arranged trays under standardized white lighting conditions in the Sensory Evaluation Laboratory at Xihua University. Evaluation was performed using a structured score sheet covering key attributes (detailed in Table S1). The overall sensory score for each sample was derived from the arithmetic mean of all panelists' ratings. This study was approved by the Institutional Review Board of Xihua University. Prior to participation, all panelists provided written informed consent after being fully informed of the study objectives. All collected data were anonymized and used exclusively for scientific analysis.

2.3. Color

Following the method described by Yan et al. (2025), the L*, a*, and b* values of different rainbow trout meat samples were determined at room temperature using a colorimeter (UltraScan Pro, HunterLab, USA).

2.4. TBARS

The TBARS value was determined according to Zhang et al. (2024) with slight modifications. Briefly, 10 g of minced fish meat was homogenized with 50 mL of 7.5% (w/v) trichloroacetic acid containing 0.1% EDTA. The mixture was shaken at 200 rpm for 30 min and then filtered. A 5 mL aliquot of the filtrate was mixed with 5 mL of 0.02 M thiobarbituric acid solution and heated in a boiling water bath for 40 min. After cooling, the solution was centrifuged at 4000×g for 20 min. Subsequently, 5 mL of chloroform was added, and the mixture was vortexed and allowed to separate. The upper layer was collected, and its absorbance was measured at 532 nm, and the TBARS value was calculated using a standard curve.

2.5. Analysis of volatile flavor compounds

Volatile compounds in cold fresh rainbow trout meat were extracted by static headspace solid-phase microextraction (SPME), following the method of Xu, Qiu, et al. (2025). Precisely 2.000 g (to the nearest 0.001 g) of meat was weighed into a 15 mL headspace vial, followed by the addition of 10 μL of the internal standard (2-methyl-3-heptanone). The vial was equilibrated at 60 °C for 20 min in a water bath. Subsequently, a pre-conditioned SPME fiber (75 μm CAR/PDMS, Supelco, USA) was exposed to the headspace for 30 min of extraction at the same temperature. The fiber was then desorbed in the GC injection port for 5 min. Analysis was performed using a GC × GC–MS (GCMS2020NX, Shimadzu, Japan). The GC × GC configuration consisted of a DB-Wax primary column (30 m × 0.32 mm × 0.25 μm) and a DB-17 ms secondary column (1.2 m × 0.18 mm × 0.18 μm). Helium was used as the carrier gas at a constant flow rate of 1.0 mL/min in splitless mode. The injector temperature was 250 °C. The oven temperature program was: 40 °C (hold 2 min), then increased to 240 °C at 6 °C/min. Mass spectrometry conditions were: electron impact ionization at 70 eV; interface temperature, 250 °C; ion source temperature, 230 °C; solvent delay, 3 min; mass scan range, 41–330 m/z (Zhang et al., 2026).

2.6. Extraction of lipids

Lipids were extracted from rainbow trout muscle according to reported methods (Wu, He, Yang, & Li, 2024). The lipid samples were extracted using a chloroform-methanol solution (2:1, v/v). The extracts were concentrated using a vacuum concentrator, dissolved in 200 μL of isopropanol, and filtered through a 0.22 μm membrane. Finally, the samples were prepared for liquid chromatography-mass spectrometry (LC-MS) analysis using a Vanquish UHPLC system (Thermo Fisher Scientific, USA).

2.7. Lipidomics analysis

MS/MS analysis was performed using higher-energy collisional dissociation with a normalized collision energy of 30 eV and dynamic exclusion enabled, following a published method (Dasilva, Muñoz, Lois, & Medina, 2019). Chromatographic separation was achieved on an ACQUITY UPLC® BEH C18 column (1.7 μm, 2.1 × 100 mm) maintained at 50 °C. The autosampler temperature was set at 8 °C. A flow rate of 0.25 mL/min was used with a 2 μL injection volume under a gradient elution program. For mass spectrometry, data were acquired in both positive and negative ionization modes with spray voltages of 3.50 kV and 2.50 kV, respectively.

2.8. Statistical analysis

Experimental data were analyzed by using IBM SPSS 27.0 software. Analyses included one-way ANOVA and LSD and Duncan tests. The results were expressed as mean ± standard error. Data visualization was performed by using Origin 2022 software.

3. Results and discussion

3.1. Sensory score

As shown in Fig. 1a, sensory evaluation revealed that with increasing irradiation dose, the color and texture scores of cold fresh rainbow trout meat decreased, while smell and overall acceptability scores first increased (at 1 kGy) and then declined. The significant decrease in color score (P < 0.05) is likely due to the oxidative degradation of pigments. Other researchers have also observed similar phenomena in their studies, where vacuum-packaged beef exhibited significant color changes following gamma irradiation at a dose of 9 kGy. Specifically, the proportion of metmyoglobin in the meat irradiated at 9 kGy was significantly higher than that in meat irradiated at low doses (Rodrigues et al., 2020). Moreover, the study detected a statistically significant reduction in texture scores (P < 0.05). This phenomenon appears attributable to irradiation-induced protein structural changes that compromised water-holding capabilities and modified water content, ultimately leading to textural degradation (Sales et al., 2020). The trends for the smell and overall acceptability scores of cold fresh rainbow trout meat were similar. The scores exhibited an increase in the 1 kGy irradiation group, though they did not differ significantly from the control group (P > 0.05). As the irradiation dose increased, both scores progressively decreased, with a significant reduction observed under 7 kGy irradiation (P < 0.05). Other researchers have also reported that irradiation induces the production of volatile organic compounds in meat, generating odors described as burnt, bloody, sweet, or pungent. Notably, high-dose irradiation produces an unacceptable “irradiation odor” (Huang et al., 2023).

Fig. 1.

Fig. 1

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Effects of different doses of electron beam irradiation on the sensory scores (a), color parameters (b), and TBARS content (c) of cold fresh rainbow trout meat.

(Note: Data are presented as mean ± standard error (SE). Different lowercase superscript letters (a–e) within the same parameter indicate significant differences (P < 0.05). Groups treated with electron-beam irradiation doses of 0, 1, 3, 5, and 7 kGy are labeled DZS0, DZS1, DZS3, DZS5, and DZS7, respectively. This labeling convention applies throughout.)

3.2. Color

As shown in Fig. 1b, with increasing irradiation dose, the L* value of the originally light orange-red rainbow trout meat showed a significant increasing trend, while the a* and b* values exhibited a significant decreasing trend (P < 0.05). Generally, the light orange-red color of rainbow trout meat mainly originates from the astaxanthin accumulated in the flesh (a type of carotenoid pigment). Astaxanthin possesses a long-chain conjugated double-bond structure, which imparts red and orange hues (contributing to redness a* and yellowness b*) (Debnath et al., 2024). As the irradiation dose increases, high-energy electron beams may directly attack astaxanthin, disrupting its conjugated double-bond structure and leading to cleavage or isomerization of the pigment molecules (Yagiz et al., 2010). Furthermore, irradiation ionizes water molecules, generating a large number of reactive oxygen species. These radicals rapidly initiate the oxidative degradation of astaxanthin, producing small, colorless or lightly colored molecular products. The more severely astaxanthin is degraded, the more drastically its color-contributing ability declines, resulting in significantly reduced a* and b* values. The increase in L* value may be related to protein denaturation (Fallah et al., 2022). Increasing irradiation doses lead to protein oxidation and aggregation, causing deterioration of the microstructure of rainbow trout meat and altering the state and distribution of moisture (e.g., increased free water content and migration of moisture from the interior to the meat surface). These changes enhance light scattering, leading to an increase in the measured L* value. Additionally, the degradation of pigments like astaxanthin also reduces light absorption, further enhancing the “whitening” effect, making the fish meat surface appear whiter and brighter. Yu et al. (2023) also observed in their study that with increasing irradiation dose, the L* value of salmon increased while the a* and b* values decreased, which further supports the findings of this study.

3.3. TBARS

As shown in Fig. 1c, the TBARS value of rainbow trout meat increased significantly (P < 0.05) with rising irradiation dose. This is because as the irradiation dose increases, high-energy electron beams attack lipid molecules, breaking their chemical bonds and generating lipid radicals. Furthermore, the high-energy electron beams also ionize water molecules, producing a large number of reactive oxygen species radicals. These radicals initiate chain oxidation reactions of lipids, leading to a significant increase in secondary oxidation products and, consequently, a marked rise in TBARS values (Zhang et al., 2025). Additionally, rainbow trout meat contains relatively abundant unsaturated fatty acids. The double bonds in these fatty acid molecules become increasingly unstable at higher irradiation doses, making them more susceptible to radical attack, which ultimately results in increased lipid oxidation products and significantly elevated TBARS values. Zhang et al. (2025) also found that the TBARS values of scallop adductor muscles showed a significant increasing trend as the electron beam irradiation dose increased from 0 to 8 kGy. Similarly, Zhao et al. (2023) reported that the TBARS value of shrimp increased significantly with higher electron beam irradiation doses.

3.4. Volatile flavor compounds

A total of 112 volatile flavor compounds were identified in cold fresh rainbow trout meat (Table S2), comprising 30 alcohols, 4 ketones, 5 phenols, 5 aldehydes, 15 hydrocarbons, 28 esters, 8 acids, and 17 other compounds. The total content of these compounds increased with irradiation dose, measuring 519.40, 680.87, 540.15, 682.43, and 740.70 μg/kg at 0, 1, 3, 5, and 7 kGy, respectively. As shown in Fig. 2a, alcohols, hydrocarbons, and esters were the most abundant classes. Notably, irradiation markedly altered the composition, hydrocarbon content (dominant in the control at 44.82%) decreased in all treated groups, while alcohols, ketones, and other compounds increased concurrently. Alcohols showed a particularly substantial rise. This overall increase in volatile compounds is likely attributable to the irradiation-induced oxidative degradation of lipids and proteins. Rainbow trout is rich in unsaturated fatty acids, which serve as key precursors for compounds like aldehydes and alcohols via lipid oxidation pathways promoted by irradiation (Mai et al., 2024). Furthermore, irradiation decomposes water and other components, generating reactive species (e.g., hydroxyl radicals) that initiate diverse oxidation reactions in the presence of oxygen, leading to the formation of varied volatile compounds that shape the final flavor profile (Jia, Shi, Zhang, Shi, & Chu, 2021). The distinct flavor signature of each dose is underscored by Fig. 2b, which shows that while 15 compounds were common to all groups, numerous unique compounds were detected in each (53, 50, 40, 46, and 50 compounds per group, respectively), indicating drastic, dose-dependent changes in the volatile profile.

Fig. 2.

Fig. 2

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Effects of different doses of electron-beam irradiation on the relative content of volatile flavor compounds (a), Venn diagram of volatile flavor compounds (b), and hierarchical clustering heatmap of volatile flavor compounds (c) in cold fresh rainbow trout meat.

Aldehydes, key products of lipid degradation with low odor thresholds, are crucial in defining the flavor profile of cold fresh rainbow trout meat. Among them, hexanal was detected in all groups and increased significantly with irradiation dose, especially at 7 kGy. This rise is likely due to the β-oxidation of fatty acids like linoleic acid (Schuster, Franke, Silcock, Beauchamp, & Bremer, 2018). As hexanal is a recognized marker of lipid oxidation and a primary contributor to meat off-flavors (Cordeiro, Mouro, Dos Santos, & Wagner, 2022), its accumulation at high doses may directly explain the characteristic “irradiation odor”. Additionally, irradiation elevated levels of long-chain aldehydes (C > 10), whose accumulation is known to generate intense greasy and fatty odors (Li et al., 2022), further contributing to the dose-dependent flavor deterioration. Brewer (2009) also pointed out that irradiation can increase the content of volatile flavor compounds that cause “irradiation odor”, such as characteristic aldehydes (e.g., propanal, pentanal, hexanal) and sulfur-containing compounds (e.g., dimethyl trisulfide, bismethylthiomethane). This suggests that the generation of “irradiation odor” may be due to the combined effect of unpleasant flavor compounds.

Numerous alcoholic volatile compounds were generated in the meat after irradiation. Alcohols are primarily derived from lipid oxidation, especially of polyunsaturated fatty acids (PUFAs) (Ruan et al., 2023). While their higher odor thresholds generally limit their flavor contribution, this effect diminishes with longer carbon chains (Zou et al., 2018). Specific changes were observed. The contents of (R)-2-octanol and (S)-2-octanol increased post-irradiation, with (S)-2-octanol detected exclusively at doses ≥3 kGy. These compounds impart pungent, oily, and fatty odors (Zheng et al., 2024), and their increase likely stems from the irradiation-induced decomposition of hydroperoxides derived from ω-6 PUFAs (e.g., linoleic acid). In contrast, the 1 kGy (DZS1) group exhibited a higher content of linalool. With its low odor threshold, linalool contributes floral and sweet notes (Tang et al., 2024), which may account for the improved odor scores in that group. This beneficial compound likely forms because low-dose irradiation favors the degradative release of specific flavor precursors over intense oxidation, facilitating transformations like those yielding linalool.

Hydrocarbons were the most abundant volatile substances in the control group. However, its high odor thresholds contributed minimally to the flavor profile of cold fresh rainbow trout meat. d-limonene was detected in all five treatment groups, with its content decreasing as the irradiation dose increased. This is because d-limonene is a monoterpene. Structurally, monoterpenes are prone to oxidation, which can lead to a decrease in their content (Li et al., 2018). As the irradiation dose increases, the generation of free radicals also rises. These free radicals can attack the carbon‑carbon double bonds of d-limonene, thereby triggering auto-oxidation and promoting its conversion into other compounds. Consequently, the original content of d-limonene decreases. Previous studies have suggested that d-limonene may impart citrus-like notes to cold fresh rainbow trout meat (Mahmoud & Buettner, 2017). Additionally, research indicates that d-limonene can reduce the adsorption of off-flavor compounds by meat proteins, thereby mitigating undesirable odors in meat products (Bi et al., 2025). The observed reduction in d-limonene content after irradiation might partially explain the decreased flavor scores of irradiated cold fresh rainbow trout meat. Furthermore, substantial increases in alkane content (e.g., pentadecane) were observed in irradiated samples and were primarily attributed to the decomposition of unsaturated fatty acids and amino acids under irradiation (Xu, Wang, et al., 2025).

Esters are produced by reactions between carboxylic acids and alcohols (Gao et al., 2024) and are known for their fruity aroma, which helps mask off-odors (Nie et al., 2022). The reduction in ester content observed at irradiation doses of 3 kGy and above may thus have contributed to flavor deterioration in the rainbow trout meat. Ketones and acids, formed from unsaturated fatty acid or amino acid oxidation, were detected at low levels, suggesting their effect on flavor was minimal (Yang et al., 2025).

To summarize, the volatile flavor compounds in the meat were significantly influenced by the irradiation dose. Doses lower than 3 kGy promoted moderate oxidation, yielding favorable compounds such as linalool and contributing to a positive sensory outcome. However, at doses of 3 kGy or higher, compounds associated with undesirable flavors—including hexanal, (R)-2-octanol, and (S)-2-octanol—increased substantially. These compounds, characterized by greasy and pungent odors, negatively impacted the overall flavor. The analysis of key differential flavor compounds indicated that lipid modifications induced by high-dose irradiation were the main factor behind these volatile profile changes.

3.5. Analysis of differences in volatile flavor compounds

Comparison of the GC × GC–MS data across the five groups showed that irradiation markedly altered the composition and content of volatile compounds. The partial least squares discriminant analysis (PLS-DA) was applied to analyze these data and distinguish the groups. The resulting score plot (Fig. 3a) indicates that each treatment group occupies a distinct position, evidencing considerable intergroup differences. To pinpoint key differential compounds, we selected 19 with Variable Importance in Projection (VIP) values >1.3 as critical markers. As listed in Table S3 and visualized in Fig. 3b, these included five alcohols, one ketone, two aldehydes, four hydrocarbons, four esters, and three other compounds. The fact that aldehydes, alcohols, and ketones emerged as the predominant signature compounds suggests that flavor variations originated largely from irradiation-induced lipid changes. The loading plot (Fig. 3c) further illustrates this divergence: groups DZS0 and DZS1 are located on the negative side of Component 1, while groups DZS3, DZS5, and DZS7 are on the positive side. Accordingly, compounds such as 10-octadecenal and d-limonene were more abundant in the DZS0/DZS1 clusters, while others like hexanal were higher in the DZS3/DZS5/DZS7 clusters.

Fig. 3.

Fig. 3

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Analysis of the effects of different doses of electron-beam irradiation on volatile flavor compounds in cold fresh rainbow trout meat using PLS-DA score plot (a), VIP score plot (b), and biplot (c).

3.6. Lipid composition analysis

Lipids serve as important flavor precursors in meat, and their composition and oxidative status directly influence both volatile flavor generation and sensory quality. In this study, we employed lipidomics to systematically investigate changes in lipid molecules in rainbow trout meat under different irradiation doses. A total of 2168 lipid molecules were detected and identified across the five groups (Fig. 4a). The most abundant classes were phosphatidylcholine (PC, 524 molecules), triglyceride (TG, 500), diglyceride (DG, 273), and phosphatidylethanolamine (PE, 187) (Fig. 4b). Collectively, these four classes accounted for more than 75% of all lipids, forming the key pool responsible for flavor and oxidative stability. Among them, PC had the highest proportion, ranging from 32.2% to 30.5% across the treatment groups. Both PC and PE play roles in cell membrane integrity and lipid metabolism. The high levels of PC and PE observed here, alongside lower phosphatidylinositol (PI) and phosphatidylglycerol (PG), are consistent with findings in Decapterus maruadsi (He et al., 2020). Furthermore, this study identified a diverse range of TG and DG in rainbow trout meat, a result consistent with lipid research on air-dried hairtail muscle reported by other scholars (Liao et al., 2023).

Fig. 4.

Fig. 4

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Effects of different doses of electron-beam irradiation on the overall lipid clustering heatmap (a) and pie chart of lipid classes (b) in cold fresh rainbow trout meat.

3.7. Multivariate statistical analysis of lipid components

As shown in Fig. 5a, principal component analysis (PCA) showed clear separation between the control and irradiated groups, indicating a significant treatment effect on the lipid profile. The corresponding loading plot (Fig. 5b) highlighted PC and TG as major discriminatory lipids. To further investigate these group differences, we performed orthogonal partial least squares-discriminant analysis (OPLS-DA). The OPLS-DA scores (Fig. 5c) displayed clear group separation along the first component, with wide spacing indicative of substantial differences. To guard against overfitting, the model was validated with a permutation test. The results (Fig. 5d) show that all permuted Q2 values fall below the original Q2, confirming the model's reliability and predictive ability.

Fig. 5.

Fig. 5

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Analysis of the effects of different doses of electron-beam irradiation on lipid components in cold fresh rainbow trout meat using PCA score plot (a), PCA loading plot (b), OPLS-DA score plot (c), and OPLS-DA permutation test plot (d).

3.8. Differential abundance of lipids (DAL) analysis

The VIP values provided by the OPLS-DA model were used to screen differential lipids from the perspective of multivariate statistical analysis, while P-values revealed differences in lipid abundance from a univariate standpoint. In this study, DALs were screened by setting the criteria of VIP > 1 and P < 0.05 simultaneously (Supplementary Table S4). The screening results were visualized, with differences between data represented by a color gradient, as shown in Fig. 6(a).

Fig. 6.

Fig. 6

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Analysis of the effects of different doses of electron-beam irradiation on differential lipid components in cold fresh rainbow trout meat using a hierarchical clustering heatmap of differential lipids (a), a categorical heatmap of differential lipids (b), and a statistical bar chart of intergroup differential metabolites (c).

Lipidomics analysis identified 583 DALs, which were classified into 36 categories based on their fatty acid chains and functional groups. They included 169 PCs, 57 PEs, 21 PIs, 54 TGs, 56 DGs, 33 cardiolipins, 12 ceramides, 33 LysoPCs (LPCs), 12 LysoPEs (LPEs), 13 monoglycerides (MGs), 14 sphingomyelins (SMs), 16 wax esters (WEs), and others. As shown in Fig. 6c, the numbers of DALs in the irradiated groups compared to the control were 662, 709, 723, and 500, respectively. In all cases, downregulated lipids outnumbered upregulated ones. Specifically, the counts of upregulated lipids were 268, 305, 282, and 176, whereas the corresponding downregulated lipids were 394, 404, 441, and 324. These widespread alterations demonstrate that irradiation profoundly alters the lipid composition and molecular profile of rainbow trout meat.

Based on the results presented in Fig. 6(a) and (b), it can be observed that after irradiation, the contents of most TG, PC, LPC, and LPE lipids in rainbow trout meat decreased, while the content of PE increased. As an important energy-storage lipid in cells, the reduction in TG content indicates that irradiation promotes TG hydrolysis or oxidation, converting it into other compounds. Studies have found that free fatty acids produced from TG decomposition can significantly influence the flavor of meat products (Zhou et al., 2021). Free fatty acids are also prone to oxidation, leading to an increase in secondary oxidation products, which explains the significant rise in TBARS values (Fig. 1c) with increasing irradiation dose. Furthermore, the total content of PC, LPC, and LPE lipids in irradiated rainbow trout meat was significantly lower than that in the control group. This is because phospholipids are rich in unsaturated fatty acids and are highly susceptible to oxidation, non-enzymatic browning, and Strecker degradation reactions (Li, Li, et al., 2020), generating a substantial pool of lipid-derived precursors for the formation of volatile flavor compounds in rainbow trout meat. For instance, oxidative cleavage of phospholipids serves as an important pathway for generating volatile flavor compounds such as aldehydes (e.g., hexanal) and alcohols (e.g., octanol) (Chen et al., 2020). This explains the significantly higher total volatile compound content in the irradiated groups compared to the control group (Table S2) and provides a precursor source for the increased levels of aldehydes and alcohols after irradiation. Fig. 6(b) also shows an increase in PE content in the irradiated groups and a decrease in LPE content. Generally, PE molecules can be degraded by phospholipase A into LPE and fatty acids (Li, Tang, et al., 2020). In this study, the lower LPE content in the irradiated groups compared to the control suggests that irradiation may have disrupted the structure of phospholipase A, thereby inhibiting this conversion reaction.

Building on the aforementioned analysis, lipids with VIP values >1.85 were selected for further examination, including LPE(18:3), DG(46:13), LPC(22:4), TG(21:3_15:3COOH), LPC(18:4), Cer(d18:3_24:1), LPC(20:5), BisMeLPA(30:8), LPC(20:3), and LPC(18:2). Among these, LPC and LPE were predominant. After irradiation, the levels of these lipids decreased significantly compared to the control. This suggests that they were released and subsequently oxidized during irradiation, contributing to the higher levels of volatile flavor compounds observed in the treated groups. LPC and LPE are hydrolysis products of PC and PE, respectively, and their formation is primarily enzyme-catalyzed (Lv et al., 2023). The observed reduction in LPC and LPE content may be attributed to irradiation disrupting the spatial structure of the relevant enzymes, leading to their inactivation and thereby suppressing this enzymatic conversion. A significant decrease in TG(21:3_15:3COOH) was also noted, which is likely because irradiation accelerated lipid oxidation, thereby reducing its abundance while simultaneously generating more volatile compounds (Liu et al., 2024).

Rainbow trout meat is rich in unsaturated fatty acids, particularly omega-3 polyunsaturated fatty acids (PUFAs). Notably, the levels of PUFAs such as arachidonic acid (20:4 n-6, AA), eicosapentaenoic acid (20:5 n-3, EPA), and docosahexaenoic acid (22:6 n-3, DHA) increased upon irradiation. Previous research indicates that a substantial portion of PUFAs in meat is deposited in glycerophospholipids (Guo et al., 2022). Consistently, this study found that the measured PUFAs were primarily concentrated in glycerophospholipids, mainly within PC and PE species (e.g., PC(14:0_20:4) + HCOO, PE(16:1_20:4)-H, PC(14:0_20:5) + HCOO, PC(16:0_20:5) + HCOO, PC(12:0_22:6) + HCOO, and PC(16:0_22:6) + HCOO). Their content increased with irradiation dose, peaking at 5 kGy, but significantly declined at 7 kGy, although it remained above the control level. This initial increase may be attributed to the irradiation-induced release of PUFAs from cell membrane phospholipids (Chiesa et al., 2022). However, at the highest dose (7 kGy), the PUFA content declined significantly. This reversal is likely because the excessive dose generates a large number of radicals, causing the oxidation rate of PUFAs to exceed their release rate. This oxidation at high doses, as evidenced by the concurrent peak in TBARS values (Fig. 1c), demonstrates that high-dose irradiation promotes the oxidation of unsaturated fatty acids. This process forms off-flavor compounds such as hexanal and (R)/(S)-2-octanol (Table S2), leading to the significant drop in smell and overall acceptability scores at 7 kGy (Fig. 1a). In contrast, a higher content of linalool, which imparts floral and sweet notes, was detected in the 1 kGy group. Lipidomics revealed that at low doses (≤5 kGy), lipid alterations were dominated by degradation rather than intense oxidation. This degradative environment may facilitate the release or transformation of specific flavor precursors, potentially explaining the generation of beneficial flavor compounds like linalool. This corresponds to the slight improvement trend in smell and overall acceptability scores observed in the 1 kGy group (Fig. 1a). The results indicate that moderate irradiation can increase the PUFA content in rainbow trout meat. This finding is consistent with previous reports on goat meat, where irradiation also significantly increased PUFA and essential fatty acid content (Jia, Shi, & Shi, 2021).

Volcano plots were used to identify differentially abundant lipids across treatment groups. As shown in Fig. 7a, irradiation significantly altered the lipid profile of rainbow trout meat. Specific lipids, such as TG (46:4) and PC (43:5), increased significantly upon irradiation. To relate these abundance changes to structural alterations, we analyzed the carbon chain length and degree of unsaturation within the four major lipid classes (PC, PE, TG, DG). In Fig. 7b, each point represents an individual lipid species, plotted by its number of carbon atoms (y-axis) and double bonds (x-axis). Overall, the total abundance of PC, DG, and TG decreased with increasing dose, whereas PE increased. A clear structural shift was observed, PC species with >40 carbons decreased, while PC and PE species with 30–40 carbons and higher unsaturation increased, peaking at 5 kGy. These shorter, more unsaturated lipids were enriched in AA, EPA, and DHA. This pattern suggests that at doses ≤5 kGy, irradiation preferentially induced the degradation of complex lipids into these smaller PUFA-rich molecules, rather than their complete oxidation. At doses >5 kGy, intense oxidation became dominant, depleting these PUFAs. This transition is supported by the concurrent peak in TBARS values (Fig. 1c), marking the accumulation of terminal oxidation products. Consequently, the role of lipid oxidation shifted from potentially generating flavor precursors to becoming the primary source of off-flavors, which correlates with the sharp decline in sensory scores (Fig. 1a).

Fig. 7.

Fig. 7

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Analysis of the effects of different electron-beam irradiation doses on lipid components in cold fresh rainbow trout meat using a volcano plot (a) and a heatmap of differential lipid features (b).

In summary, lipidomics analysis revealed that electron-beam irradiation fundamentally altered the lipid composition of rainbow trout meat by promoting the degradation and oxidation of key lipids. Low-dose irradiation primarily induced lipid degradation, releasing flavor precursors which could contribute to the formation of favorable flavor compounds. As the irradiation dose increased, oxidation became dominant, resulting in substantial oxidation of PUFAs. This generated off-flavor compounds such as hexanal and octanol and caused a sharp increase in TBARS values, ultimately leading to a marked deterioration in sensory scores.

3.9. PCA of key volatile flavor compounds and differential lipids

As shown in Fig. 8, PCA effectively consolidated multivariate data—including differential lipids, key volatile compounds, a lipid oxidation indicator (TBARS), and color parameters—to systematically elucidate the overall changes and intrinsic relationships in the quality of cold fresh rainbow trout meat under different electron-beam irradiation doses. The cumulative variance explained by PC1 and PC2 reached 92.6%, indicating that these two components captured most of the sample information. PC1 (accounting for 88.3% of the variance) represented the core dimension corresponding to the irradiation dose effect. Sample scores exhibited a clear gradient along PC1 from negative to positive values, sequentially corresponding to irradiation doses from 0 to 7 kGy. Critically, this axis clearly separated the low-dose groups (DZS0/DZS1) from the medium- and high-dose groups (DZS3/DZS5/DZS7), confirming that the treatments induced systematic, dose-dependent differences.

Fig. 8.

Fig. 8

Open in a new tab

PCA plot of key volatile flavor compounds and differential lipids.

To elucidate the driving factors behind group separation, we screened key markers based on their VIP values (using thresholds of >1.5 for volatile compounds and > 1.85 for lipids). On the positive side of PC1, variables with high VIP values—such as TBARS, L*, and hexanal—were closely grouped together. Among these, TBARS is a direct indicator of lipid oxidation, while hexanal is a characteristic oxidation product of polyunsaturated fatty acids. The alignment of their vectors indicates that the positive side of PC1 predominantly captures the “irradiation-induced lipid oxidation” process. Consistent with this interpretation, high-dose irradiation promoted lipid oxidation, leading to the accumulation of products like hexanal and an increase in meat lightness (L* value). Hexanal was positively correlated with compounds such as 2-octanol, which exhibited spicy, oily, and fatty odors (Zheng et al., 2024). This further suggests that the deterioration of rainbow trout meat flavor caused by irradiation may be attributed to the synergistic effect of these unpleasant flavor compounds. Additionally, it can be observed from the Fig. 8 that the lipid oxidation indicator (TBARS) exhibited a negative correlation with the a* and b* values, while showing a positive correlation with the L* value. This may be attributed to the chain oxidation of lipids induced by electron beam irradiation, which leads to the generation of free radicals. These free radicals, along with the secondary products of lipid oxidation, can trigger the oxidative degradation of astaxanthin, subsequently resulting in a decrease in the a* and b* values. Furthermore, the degradation of pigments such as astaxanthin may reduce light absorption, causing the surface of the cold fresh rainbow trout meat fillets to appear whiter and brighter, thereby contributing to an increase in the L* value. In contrast, the negative side of PC1 was characterized by indicators enriched in non- or low-dose irradiated samples. These included sensory scores (overall acceptability, smell, color, texture), a* and b* values, several polyunsaturated phospholipids (e.g., LPC(18:4), LPC(20:5), LPE(18:3)), and fresh-flavor compounds like d-limonene, all of which were closely associated in the PCA space. In the plot, these lipids—particularly PUFA-rich phospholipids, which are important flavor precursors and membrane components—were positioned close to and shared similar vector directions with favorable sensory attributes. This spatial proximity visually confirms their positive correlation, underscoring that preserving these lipid components is crucial for maintaining desirable flavor and sensory quality.

Consequently, PCA revealed a clear dose-dependent pattern: as the irradiation dose increased (along PC1), lipid oxidation intensified (reflected by increased TBARS values), which led to the degradation of flavor precursor lipids (e.g., decreased polyunsaturated phospholipids) and the generation of specific undesirable flavor compounds (e.g., increased hexanal), coupled with a reduction in fresh-flavor compounds (e.g., decreased d-limonene). These changes collectively resulted in decreased sensory scores and a reduction in the a* and b* values. PC2 (contribution rate: 4.3%) accounted for within-group variation, for instance, by distinguishing among the medium-dose samples. The sensory indicator vectors were closer to the low-dose groups, reaffirming the protective effect of low-dose treatment on sensory quality. In summary, PCA not only successfully differentiated rainbow trout meat treated with different irradiation doses but, more importantly, pinpointed key driving variables based on their VIP values. By leveraging these multivariate relationships, we established a coherent causal chain linking irradiation dose to lipid oxidation, subsequent changes in lipid and volatile compound profiles, and the eventual deterioration of sensory quality. Thus, the analysis provides core multivariate statistical support for elucidating the correlations among changes in lipids, flavor, and sensory attributes.

4. Conclusion

This study clarifies the dose-dependent mechanism and key thresholds by which electron-beam irradiation affects the quality of cold fresh rainbow trout meat. Low-dose irradiation (< 3 kGy) may primarily induce lipid degradation, promoting the release of specific beneficial flavor precursors. This resulted in a slight improvement in smell and overall acceptability scores in the 1 kGy group. In contrast, high-dose irradiation (≥ 5 kGy) triggered intense lipid oxidation, as evidenced by a significantly increase in TBARS values. This process led to the oxidative degradation of key phospholipids (e.g., PC, LPC) and the accumulation of off-flavor aldehydes and alcohols (e.g., hexanal, 2-octanol), producing an “irradiation odor” and causing a significant decline in sensory scores. Multivariate analysis ultimately confirmed the association between irradiation dose and lipid oxidation, which subsequently altered volatile flavor compounds and ultimately led to deteriorated sensory quality. The findings clearly reveal a mechanism whereby irradiation promotes lipid degradation and oxidation, alters flavor precursors, generates specific volatile compounds, and ultimately influences sensory attributes. However, this study focused solely on the immediate effects of electron-beam irradiation at varying doses on the lipid composition and volatile flavor compounds of cold fresh rainbow trout meat. Future research should systematically investigate how different irradiation doses influence microorganisms, lipids, and flavor compounds in rainbow trout during storage. Additionally, applying metabolomics could help clarify the specific lipid metabolic pathways affected by different irradiation doses, thereby informing the development of “flavor-friendly” irradiation protocols. Another important direction for future work is to systematically examine the potential impact of protein oxidation on flavor, which was not addressed in the present study.

CRediT authorship contribution statement

Fanyi Gong: Writing – review & editing. Buzhou Xu: Formal analysis. Li Yang: Formal analysis. Chang Su: Formal analysis. Jiaxin Chen: Formal analysis. Xingzhong Zhang: Formal analysis. Yong Yu: Formal analysis. Shichen Zhu: Formal analysis. Jie Tang: Formal analysis. Dong Zhang: Investigation, Funding acquisition. Hongjun Li: Methodology.

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.

Acknowledgements

This research was funded by Natural Science Foundation of Sichuan Province (2024NSFSC2081), Chongqing Municipal Technology Innovation and Application Development Special Key Project (CSTB2023TIAD-KPX0029), and Science and Technology Department of Sichuan Province, China (2023YFN0015).

Footnotes

Appendix A

Supplementary data to this article can be found online at https://doi.org/10.1016/j.fochx.2026.103717.

Appendix A. Supplementary data

Supplementary material
mmc1.docx (190.1KB, docx)

Data availability

Data will be made available on request.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary material
mmc1.docx (190.1KB, docx)

Data Availability Statement

Data will be made available on request.

Articles from Food Chemistry: X are provided here courtesy of Elsevier

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