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. 2025 Nov 5;32:103264. doi: 10.1016/j.fochx.2025.103264

Glycerophospholipid fatty acid remodeling optimizes lipid nutrition and volatile organic compound composition in freshwater fish with linolenic acid intervention

Zijie He a, Junli Wang b, Yun Wei a, Xiao Yan a, Yuanyou Li c, Guoxing Nie a,⁎, Dizhi Xie c
PMCID: PMC12657307  PMID: 41323677

Abstract

To investigate α-linolenic acid (ALA) intervention effects on lipid remodeling and volatile organic compounds (VOCs) in Cyprinus carpio, fish were fed soybean oil (SO, 5.37 % ALA) and flaxseed oil (SL, 32.66 % ALA) diets for 30 weeks. Key results: ALA interventions significantly increased n-3 long-chain polyunsaturated fatty acids (LC-PUFAs) without altering protein or amino acid content. ALA interventions preferentially incorporated docosahexaenoic acid at the phosphatidylethanolamines (PEs) sn-2 position while suppressing saturated fatty acids in PEs sn-1 and monounsaturated fatty acids in phosphatidylcholines (P < 0.05). ALA interventions enhanced pleasant VOCs (pent-1-en-3-ol) but reduced off-flavor VOCs (oct-1-en-3-ol, heptanal), with n-3 PUFAs correlating positively and n-6 PUFAs negatively (P < 0.05). Lipidomics revealed glycerophospholipids as a core target for ALA, with significant metabolic pathways enrichment (P < 0.05). In summary, ALA interventions optimize the nutrition and VOCs of freshwater fish by remodeling glycerophospholipid molecules, supporting the supply of high-quality aquatic foods.

Keywords: ALA, Cyprinus carpio, Lipid molecular modification, VOCs, Glycerophospholipid

Highlights

  • •

    Linolenic acid (ALA): a potential regulator for high-quality aquatic food supply.

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    ALA intervention effectively promoted DHA binding at the PE sn-2 position.

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    ARA and DHA competitively bind to the sn-2 position of PEs.

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    ALA optimizes the volatile flavor in the muscle of Yellow River carp.

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    The lipid molecule with the greatest influence by ALA was glycerophospholipids.

1. Introduction

Clinical data show that n-3 long-chain polyunsaturated fatty acids (n-3 LC-PUFAs) have particularly important physiological functions in the management of inflammation, the prevention of cardiovascular diseases, and the promotion of brain development in humans (Das, 2003; Riediger et al., 2009). They are predominantly found in aquatic animals such as fish and microalgae, yet they are not present (or small amounts) in terrestrial animals, eggs, or milk. Because of this unique nutritional value, fish is frequently regarded as a high-quality food and a significant source for the development of bioproducts and functional foods (Golden et al., 2021). In recent decades, the rapid advancement of global aquaculture technology has made aquaculture the fastest-growing area of agriculture in the world. This has led to a significant expansion in the quantity and variety of aquatic products, which have become a valuable source of provision and nutrition for the world's population (Naylor et al., 2021). However, the rapid advancement of aquaculture and intensive farming in recent years has had a detrimental impact on the quality of fish fillets, manifesting as an unpleasant aroma and taste, diminished levels of LC-PUFA, and compromised texture (Wang et al., 2024a). These factors pose obstacles to the supply of high-quality aquatic food.

Currently, there is a steady increase in consumer demand for high-quality aquatic products, necessitating a shift from a “quantity-focused” to a “quality-focused” approach to further advance aquaculture (Jiang et al., 2022). Accumulative studies indicated that nutritional intervention is a prominent approach to enhancing the quality of farmed fish (Song et al., 2024; Sun et al., 2024; Wen et al., 2023). For example, dietary additions of substances threonine can improve the sensory quality, nutritional value (essential fatty acids, EFAs; and essential amino acids, EAAs), and flavor of grass carp (Ctenopharyngodon idella) (Wen et al., 2023). Additionally, changing the dietary lipid source or fatty acid (FA) composition can also improve the quality of cultured fish. However, traditional research has primarily focused on the regulation of total fatty acid composition in flesh by dietary essential fatty acids. For instance, Xie et al. (2022) demonstrated that modulating the dietary α-linolenic acid (ALA) ratio enhanced the synthesis and endogenous content of n-3 long-chain polyunsaturated fatty acids (LC-PUFAs) in tilapia. In contrast, the impact of essential fatty acids on the fine molecular structure of lipids, particularly the stereospecific distribution (sn-positioning) of fatty acids, remains poorly understood. Furthermore, fish flesh quality encompasses multiple sensory attributes. In addition to nutritional value, texture characteristics and flavor are equally important. n-3 LC-PUFAs have been demonstrated to improve the texture characteristics of farmed fish—including firmness, chewiness, and springiness—by influencing muscle fiber development and collagen content (He et al., 2022; Li et al., 2020). In terms of flavor, although general associations between lipid metabolism and volatile organic compounds (VOCs) formation have been explored in other aquatic species, research on whether dietary fatty acids drive the remodeling of specific lipid molecules to regulate flavor profiles is still limited. Moreover, studies specifically targeting the common carp (Cyprinus carpio haematopterus) are notably lacking. Wang et al. (2024b), using electronic nose/tongue technology, discovered that dietary DHA enhanced the umami and sweet taste perception in blunt snout bream (Megalobrama amblycephala); however, this technique could not unveil the underlying molecular lipid mechanisms. Similarly, Chen et al. (2024) confirmed that a dietary ALA nutritional strategy enhanced the bioavailability of fatty acids and volatile flavor in tilapia, but their research lacked molecular-level insight into the specific lipid precursors responsible for flavor changes.

The Yellow River carp is one of the most renowned traditional fish with cultural heritage and is widely farmed in northern China (Nakajima et al., 2019). In recent years, cultured Yellow River carp have similarly faced problems such as reduced nutritional value of muscle FAs, soft taste, unpleasant odor, etc., which have brought serious negative impacts on the production and consumption of Yellow River carp (Wang et al., 2024a). Our research group previously identified that high ALA diets are beneficial for healthy growth and endogenous n-3 LC-PUFA synthesis in small-sized fish (Unpublished data). However, the impact of a high-ALA dietary intervention on the lipid profile and volatile flavor compounds of common carp remains unknown. Therefore, this study aimed to investigate the effects of dietary essential fatty acids (EFAs) on the molecular composition, abundance, and structure (sn-positional distribution) of lipids, as well as on the volatile flavor compounds in common carp. The overarching objective is to provide novel insights into the potential of EFA interventions to enhance the volatile flavor and nutritional value of farmed fish, thereby facilitating the production of high-quality aquatic foods.

2. Materials and methods

2.1. Experimental diets, fish, and feeding procedure

Two isolipid (5.5 %) and isoprotein (33.5 %) diets were formulated using soybean oil and flaxseed oil as lipid sources. The control diet (SO, contained 5.37 % ALA) contained only soybean oil as the sole source of lipids, while the experimental diet (SL, contained 32.66 %) included soybean oil and flaxseed oil (about our previous study (unpublished). The preparation and storage of the diets were identical to those previously described in detail (He et al., 2022). Table S1 presents detailed diet composition, proximate composition, and FA composition. A total of 240 Yellow River carp of similar genetic background and initial weight (∼11.64 g) were randomly allocated to six floating cages (2.0 m × 2.0 m × 1.5 m). The fish were divided into 3 portions of each dietary treatment (40 tails per portion) and were fed artificially 3 times per day (7,00, 11,30, and 17:30) during the 30-week culture experiment. The quantity of food provided was fixed at approximately 3 % of the fish's body weight. The culture conditions were as follows: dissolved oxygen levels were maintained at ≥5.0 mg/L, the ammonia concentration was kept below 0.05 mg/L, the water temperature was kept at (25.5 ± 3.5 °C), and the pH was maintained between 7.4 and 8.6.

2.2. Sample collection

The study was conducted in strict adherence to the Guidance on Treating Experimental Animals (the Ministry of Science and Technology of the People's Republic of China, 2006), and the Regulations for the Administration of Affairs Concerning Experimental Animals (Order No. 2 of the State Science and Technology Commission, 1988). Additionally, the research adhered to the guidelines established by the Science Research Experiment Ethics Committee at Henan Normal University (HNSD-SCXY-2116BS1066). All procedures were made to minimize the suffering of common carp. Following the conclusion of the culture experiment, the fish were fasted for 24 h and anesthetized using a solution of MS222 (CAS: 886-86-2, Shanghai Aoding Biochemical Technology Co., Ltd.) at a concentration of 100 mg/L to render them unconscious. Subsequently, the fish were killed by a blow to the head, and the muscle of fish was obtained. The entire sample collection process was completed on ice. Muscle samples from six fish per net box were quickly frozen in liquid nitrogen and kept at −80 °C for further examination of proximate values, FA and AA compositions, VOCs, and lipids metabonomics. Concurrently, an additional four fish from each net box were analyzed for eating quality and textural characteristics.

2.3. Analysis of proximate composition

The proximate composition of muscle was determined by the AOAC (2010) standardized method. In brief, muscle samples were sheared and dried to a constant weight at a constant temperature of 105 °C to detect moisture. Kjeldahl nitrogen determination is used to determine crude protein (CP) content, which includes digesting the samples with acid and then reacting them with an alkaline solution. Soxhlet extraction was used to evaluate the lipid content using a lipid analyzer (OPSIS, Skåne, Sweden). Subsequently, the samples were carbonized and then burned at a constant temperature of 550 °C until a constant weight was achieved to determine the ash content.

2.4. Analysis of AA composition

An accurate measurement of 100 mg of the muscle sample was taken and placed within an AA digestion tube. And 10 mL of HCl (6 mol/L) was transferred into the tube. Subsequently, the sample was sealed with N2 and hydrolyzed at 110 °C for 24 h. Thereafter, the sample was cooled to room temperature, and filtered to remove the residue, and the volume was determined to be 50 mL. Two milliliters of the volume-determined sample were pipetted into a 10-ml glass tube and blown to dryness with nitrogen at 60 degrees Celsius. The sample was then added to the sample buffer, shaken, and mixed well. It was then filtered through a 0.22-μm filter membrane into a brown vial. The AA composition of the muscle was determined by an AA analyzer.

2.5. Analysis of FA composition

Gas chromatography was used to detect the composition of muscle FAs. Total lipids were extracted by the chloroform-methanol (2:1 v/v) method, and then FAs methyl esters were prepared by reacting with a sulfuric acid-methanol solution. These were then transferred to a brown injection bottle and detected by GC. Sigma standards are used to identify the composition of FAs, and the 17:0 standard was added for quantitative calculation. The relevant calculation formulas for fatty acid nutritional value indices are detailed in Table S2. The following GC conditions were employed: the GC column was a Hewlett-Packard DB-WAX column (15 m × 250 μm × 0.25 μm); the column chamber heating conditions were as follows: the initial temperature of the column chamber was 120 °C, and the maximum temperature was 250 °C. Inlet settings: The heater temperature was adjusted to 250 °C, and the carrier gas (high-purity nitrogen) flow rate of 1.4 mL/min. Detector conditions: The heater temperature was set to 250 °C, the airflow rate was 400 mL/min, the hydrogen gas flow rate was 40 mL/min, and the tail-blown nitrogen flow rate was 25 mL/min. For detailed instructions on the specific operation procedures, refer to the previously described method (Xie et al., 2022).

2.6. Analysis of muscle texture and edible quality

Two muscles of identical dimensions (1.0 × 1.0 × 1.0 cm) were excised from the lateral aspect of the fish with a sharp scalpel for texture analysis. One side of the muscle samples was steamed in boiling water for five minutes to analyze the texture of the cooked muscle, while the other side was tested for the texture of the raw muscle. Texture analyzers (Brookfield, Middleboro, USA) are used to evaluate the texture of muscle samples in the TPA mode. This involved compressing the samples twice with an 8-mm cylindrical (2 mm/s), with a trigger force of 5 g, a retention time of 5 s, and a deformability of 60 %.

The muscle was weighed and found to be approximately 1 g. It was then added 9 times distilled water, homogenized, and its pH value was measured using a pH analyzer (Leici, Shanghai, China). Muscle-cooked meat percentage (COP)was obtained by the method of Xie et al. (2022). It was assessed by placing fresh muscle samples (M1) in heat-resistant plastic bags and reweighing after steaming at 100 °C for 5 min (M2). To quantify the drip loss rate (DLR), fresh muscle samples (M3) were placed in a refrigerator (4 °C, 24 h) and reweighed after absorbing the water droplets (M4).

[1] COP (%) = (M2 − M1)/M1 × 100;

[2] DLR (%) = (M4 − M3)/M3 × 100.

2.7. Identification of VOCs

The VOC composition of fish samples was detected by the FlavourSpec®Flavor HS–GC–IMS instrument (Shandong Haineng Scientific Instrument, Shandong, China) (Li et al., 2024). Briefly, 1.5 g of muscle was weighed and incubated at 60 °C for 15 min. Subsequently, 0.5 mL of sample was automatically injected into the GC-IMS instrument, and the injector temperature was 85 °C. The GC separation was carried out using an MXT-5 capillary column at a column temperature of 60 °C. The carrier gas (nitrogen purity ≥99.999 %) was maintained at 2 mL/min for the initial 2 min and 100 mL/min for the subsequent 18 min. Retention times (RTs) and retention indices (RIs) of VOCs were measured using an external standard mixture of C4–C9 n-ketones (Sinopharm Chemical Reagent Beijing Co., Ltd., Beijing, China). The reactant ion peak (RIP) originates from ionized drift gas clusters (e.g., (H₂O)ₙH+ in positive mode) and serves as the fundamental reference peak in IMS. All measured drift times were normalized by dividing by the RIP drift time to obtain the dimensionless drift time (RIP-relative). VOCs were identified by matching both RI and drift time against a commercial IMS library (library match ≥85 %). Four samples were analyzed to ensure accuracy under the same conditions.

2.8. Identification of lipid composition

The muscle sample (30 mg) was subjected to pre-cooling at −20 °C (2 min) by the addition of 300 μL of a methanol-water solution (V:V = 1:1, including a mixed internal standard of 0.1 mg/mL, methanol preparation). Tissue samples were homogenized using a fully automated high-throughput tissue grinder (Shanghai Jingxin Industrial Development Co., Ltd., Shanghai, China) with grinding beads at an oscillation frequency of 60 Hz for 2 min, and subsequently mixed with 300 μL of a chloroform solution. The mixture was then extracted by ultrasonic extraction for 10 min. Following centrifugation at 12,000 rpm (4 °C, 10 min), the chloroform in the lower layer (200 μL) was transferred to a fresh centrifuge tube (tube II). In the original centrifuge tube, 300 μL of chloroform-methanol (V:V = 2:1) was added, thoroughly mixed, and subjected to ultrasonic extraction for 10 min. The tube was placed at −20 °C for 20 min, followed by centrifugation (12,000 rpm, 4 °C, 10 min). The chloroform in the lower layer (200 μL) was transferred to tube II and combined, yielding a total of 400 μL. Half of the mixing liquid should be transferred into the LC-MS vials and volatilized. Subsequently, 300 μL of isopropanol-methanol (V:V = 1:1) was added to re-dissolve the lipid. The solution was sonicated in ice water (3 min) and transferred to a 1.5 mL centrifuge tube. Then, it was centrifuged at −20 °C for 2 h (12,000 rpm, 4 °C, 10 min), and 150 μL of the resulting upper layer was loaded into the LC-MS injection vials with lined tubes and used for the LC-MS analysis. Lipidomics analysis was performed using a UHPLC system coupled with a Q-Exactive mass spectrometer equipped with an ESI ion source (Thermo Fisher Scientific, Waltham, MA, USA). Separation was achieved on a Waters ACQUITY UPLC BEH C8 column (2.1 mm × 100 mm, 1.7 μm). The mass spectrometer was operated in both positive and negative ionization modes. Full MS scans were acquired over a range of m/z 150–1500 with a resolution of 70,000. MS/MS scans were acquired at a resolution of 17,500. The quality control (QC) sample consisted of the extracts of all samples in equal volumes. The positional distribution of fatty acyl chains in triglycerides and phospholipids was determined based on the interpretation of diagnostic fragment ions in LC-MS/MS data. Data-dependent MS/MS spectra were matched against theoretical spectra in lipid databases (LipidSearch). The identities of the specific fatty acyl chains were confirmed by analyzing their corresponding precursor ions and characteristically fragment ions. It is imperative to note that all extraction reagents were pre-cooled at −20 °C before their utilization.

2.9. Calculations and statistical analysis

The experimental data were statistically analyzed and visualized using SPSS 22.0 (USA, SPSS Inc.) and GraphPad Prism 9.0 (USA, GraphPad Software Inc.). Independent samples t-tests were employed to analyze the significant differences between groups (P < 0.05). The values are exhibited as the mean ± SEM (standard error of the mean). We have implemented a consistent notation scheme where * denotes P < 0.05, ** denotes P < 0.01, and *** denotes P < 0.001. Multivariate statistical analyses, including principal component analysis (PCA), partial least squares-discriminant analysis (PLS-DA), and orthogonal PLS-DA (OPLS-DA), were conducted on the Wekemo Bioincloud platform (https://www.bioincloud.tech). To validate the robustness of the PLS-DA and OPLS-DA models and prevent overfitting, 200 permutation tests were performed. Hierarchical clustering heatmaps and correlation heatmaps (Pearson's correlation coefficient) were generated using the “Advanced Clustering Heatmap” and “Advanced Correlation Heatmap” tools on the Metware Cloud platform (https://cloud.metware.cn). These cloud-based tools are built upon widely recognized open-source R packages (e.g., ComplexHeatmap), support universal file formats (.xlsx or .txt), and feature menu-driven interfaces with transparent parameter selection. Crucially, all analytical platforms are freely accessible without regional access restrictions. The differential VOCs and lipids in the muscles of Yellow River carp from different treatments were determined by employing a variable importance in projection (VIP) value of greater than 1 and P < 0.05.

3. Results and discussion

3.1. Edible quality, nutritional composition, and textural characteristics

Following a thirty-week feeding period, both groups of Yellow River carp reached the market size (>600 g). The results demonstrated that the intervention of ALA did not influence the proximate composition (Fig. 1A) and AA composition (Fig. S1) in the muscle of the Yellow River carp (P > 0.05). However, in terms of edible quality and textural characteristics (Fig. 1B, D), the ALA intervention significantly increased the COP and pH of Yellow River carp and improved muscle hardness, chewiness, and gumminess to some extent (P < 0.05). This is similar to the findings of tilapia, in which muscle texture was improved by high ALA intervention (Xie et al., 2022). This may be attributed to the role of n-3 PUFA in promoting myofiber development. Wang et al. (2020) found that DHA intervention could promote myofiber development by activating the AMPK/Sirt1 pathway, which promoted myofiber proliferation in blunt snout bream, thereby improving muscle chewiness, hardness, and gumminess. Similar results were also reported in golden pompano (Trachinotus ovatus) (Li et al., 2020) and common carp (He et al., 2022). Meanwhile, the antioxidant properties of n-3 PUFA help to reduce oxidative stress during muscle growth and further optimize muscle textural properties. This implies that the palatability and muscle textural properties of farmed fish can be improved by ALA interventions.

Fig. 1.

Fig. 1

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Nutrition, edible quality, and texture in the muscle of the Yellow River carp after ALA intervention. Proximate compositions (A), edible quality (B), fatty acid compositions (C), texture (D), and fatty acid nutrient values (E, F). CR: Carcass Ratio, COP: Cooked Meat Percentage, DLR: Drip Loss Rate. UI: Unsaturation index, n-3/n-6: omega-3/omega-6 polyunsaturated fatty acids, PUFA/SFA: Polyunsaturated fatty acids/saturated fatty acids, FLQ: Fish lipid quality/flesh lipid quality, HPI: health-promoting index, IT: index of thrombogenicity, HH: Hypocholesterolemic/hypercholesterolemic ratio. IA: Index of atherogenicity. Data in (A), (B), (D), (E), and (F) are presented as mean ± SEM (n = 4). * and ** indicate significant differences at P < 0.05 and P < 0.01, respectively. Panel (C) shows a cluster heatmap of fatty acid levels from a single biological replicate. The colour scale represents the Z-score of normalized intensity, with blue to red indicating content from low to high. The rectangular box in panel C is used to highlight fatty acids that exhibit significant differences between the control and experimental groups. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

The most important quality of fish is its unique nutritional value of fatty acids (rich in n-3 LC-PUFA, especially 20:5n-3 and 22:6n-3), which are essential for the development of the human brain and retina. However, these cannot be synthesized in humans and must be obtained from food sources. In this study (Fig. 1C), the ALA intervention led to a significant increase in ALA, 20:3n-3, 20:5n-3, 22:6n-3, n-3 PUFA, and LC-PUFA contents in the muscle of Yellow River carp, while concurrently resulting in a substantial decrease in LA, 18:3n-6, 20:3n-6, 20:4n-6, and n-6 PUFA contents (P < 0.05). Additionally, the FLQ, HPI, and HH indices describe the nutritional value of foods in terms of their healthcare functions. Generally, the larger the ratios of these indices, the higher the nutritional value. The IA and IT describe the atherogenic and thrombogenic potentials of foods. According to a summary of literature, the values of IA and IT in fish muscle are generally distributed in the ranges of 0.21–1.41 and 0.14–0.87, respectively (Chen & Liu, 2020). The smaller their ratios, the higher the nutritional value. In this study (Fig. 1E, F), the FA nutritional indices UI, PUFA/SFA, n-3/n-6 PUFA, FLQ, HPI, and HH were significantly increased in the muscle under the intervention of ALA (P < 0.05). Furthermore, IA and IT in the muscle were significantly reduced by 13.79 % and 51.02 %, respectively (P < 0.05; Fig. 1F). Similarly, n-3/n-6 PUFA, PUFA/SFA, and FLQ of tilapia muscle were significantly improved by the intervention of a high ALA (Xie et al., 2022). These suggest that the FA composition of muscle can be optimized by high ALA intervention to improve its health value.

3.2. Lipid molecular modification: Distribution of FAs in key lipids

Furthermore, the nutritional value of lipids depends not only on their fatty acid composition but also on the positional distribution of fatty acids on the glycerol backbone (Kubow., 1996). Triglyceride (TG), phosphatidylcholine (PC), and phosphatidylethanolamine (PE) are important lipid components. In neutral lipids (Fig. 2A), the percentages of 16:0 and 18:1 at sn-1 positions; 14:0, 16:0, 18:1, and 20:4n-6 at sn-2 positions; and 18:1 and 18:2n-6 at sn-3 positions were significantly decreased by ALA intervention (P < 0.05). Conversely, the percentages of 16:1, 18:3n-3, and 20:5n-3 at sn-1 positions; 18:2n-6 and 18:3n-3 at sn-2 positions; and 18:3n-3, 20:5n-3, and 22:6n-3 at sn-3 positions were significantly increased (P < 0.05). In polar lipids (Fig. 2B, C), there was a significant decrease in the percentages of 18:1 at sn-1 positions; and 16:1, 18:1, and 18:2n-6 at sn-2 positions; but a significant increase in the percentage of 18:3n-3, 20:4n-6, 20:5n-3, and 22:6n-3 at sn-1 positions; and 18:3n-3, 20:5n-3, and 22:6n-3 at sn-2 positions in PCs by ALA intervention (P < 0.05). Similarly, there was a significant decrease in the percentages of 18:0 and 18:2n-6 at sn-1 positions; and 20:4n-6 at sn-2 positions; but a significant increase in the percentages of 16:0, 18:3n-3, 20:4n-6, 20:5n-3, and 22:6n-3 at sn-1 positions; and 18:3n-3, 20:5n-3, and 22:6n-3 at sn-2 positions in PEs by ALA intervention (P < 0.05).

Fig. 2.

Fig. 2

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The positional distributions of key fatty acids in triglyceride (TG), phosphatidylcholine (PC), and phosphatidylethanolamine (PE) molecules in the muscle of the Yellow River carp after ALA intervention. Data presented as mean ± SEM (n = 4). *, ** and *** represent significant differences with P < 0.05, P < 0.01 and P < 0.001, respectively. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

SFAs (16:0, 18:0) in Yellow River carp muscle were preferentially distributed at the sn-1 position of the TG molecule and the sn-1 position of the PC and PE molecules. In contrast, a high ALA nutritional intervention resulted in a reduction in the content of 16:0 in TG sn-1 and sn-2 positions. Studies on human fat absorption indicate that 16:0 at the sn-2 position is more readily digested and absorbed by the body than 16:0 at the TG sn-1 and sn-3 positions (Kubow., 1996). However, an excessive intake of 16:0 at the sn-2 position may increase the risk of obesity and atherosclerosis (Kubow., 1996). In contrast, the sn-1/3 position of 16:0 can be selectively lipidated by lipase (Wei et al., 2020). Consequently, the elevated 16:0 levels in the external position of Yellow River carp muscle lipid molecules suggest their high mobilization potential in Yellow River carp metabolism, which also indicates that humans may have a reduced risk of cardiovascular disease, hypertension, and obesity when consuming Yellow River carp fillets after a high ALA intake. 18:1 was predominantly bound in TGs and PCs. It was higher in the sn-2 position than in the sn-1 or sn-3 positions. 18:2n-6 was predominantly distributed at the sn-3 position in TGs and at the sn-2 position in PCs and PEs. From a nutritional perspective, 18:2n-6 at the sn-1/3 position is more likely to be released from TGs into the circulation, potentially increasing the risk of inflammation; 18:2n-6 at the sn-2 position is relatively stable and beneficial for retention (Naughton et al., 2016). Therefore, reducing the amount of 18:2n-6 at the sn-1/3 position may be beneficial for human health. In the present study, we found that the effect of high ALA on the change in 18:2n-6 content at different sites of TG after the intervention was positive. Compared with 18:2n-6, 18:3n-3 was mainly distributed in the sn-3 position of TG. LC-PUFA was predominantly distributed in the sn-2 position of PE and PC, with 20:4n-6 and 22:6n-3 predominantly distributed in the sn-2 position of PE and 20:5n-3 predominantly distributed in the sn-2 position of PC. Meanwhile, the 20:4n-6 and 22:6n-3 at the PE sn-2 positions of Yellow River carp muscle changed dramatically after high ALA nutritional intervention, showing a sharp decrease in 20:4n-6 and a significant increase in 22:6n-3. This indicates that 20:4n-6 and 22:6n-3 were competitively esterified at the sn-2 position of PE, which is consistent with the prevailing perspective that PUFAs were preferentially esterified at the sn-2 position of phospholipids (Ruizlopez et al., 2015; Tocher et al., 2008). This observed distribution pattern may reflect an underlying biological mechanism, warranting further investigation. In general, the high-ALA intervention remodeled the lipid structure in Yellow River carp by altering the molecular composition and positional distribution of fatty acids, leading to an enrichment of n-3 PUFAs. Considering that n-3 PUFA has been shown to have significant anti-inflammatory benefits in humans, the enrichment of site-specific n-3 PUFA in the muscle lipids seems to imply that the Yellow River carp may be a highly efficient food source for humans to obtain n-3 PUFA.

3.3. VOCs composition was optimized by ALA intervention

In addition, the intervention of FA nutrition can also influence the development of fish flavors by regulating physiological metabolic processes (Cheng et al., 2023). The acceptability of aquatic products is closely associated with their flavor, and a fresh flavor can stimulate people's interest (Renata et al., 2022). In general, the flavor of aquatic products is comprised of both odor and taste. Odor is the initial sensory impression formed by VOCs that elicit an olfactory response, and it is a crucial element in the sensory evaluation of these products (Yu et al., 2022). The VOCs found in aquatic products mainly include acids, ketones, hydrocarbons, aldehydes, esters, alcohols, etc. (Cheng et al., 2023). Through qualitative and quantitative analysis of VOCs present in Yellow River carp muscle, a total of 48 VOCs were distinguished in the muscle (Fig. 3A and Table S3), which were classified as 16 aldehydes, 11 ketones, 16 alcohols, 2 acids, 1 furan, and 2 unknown compounds (Fig. 3B). Alcohols, ketones, and aldehydes were the main volatile organic compounds (Fig. 3C, D). Similar findings have been reported in studies on chill-stored Tilapia (Chen et al., 2024).

Fig. 3.

Fig. 3

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VOC profiles in the muscle of the Yellow River carp after ALA intervention. Panel (A) shows a representative 2D GC–IMS plot of VOC signals. The drift time (RIP-relative) represents the dimensionless value calculated by dividing all measured drift times by that of the RIP. The plot is generated using these calculated values; the axis itself is not mathematically standardized. Panel (B) shows the number of identified VOCs. The percentage (C) and relative concentration (D) of VOCs across different chemical classes. Data in (C) and (D) are presented as mean ± SEM (n = 4), derived from quantified VOC intensities across biological replicates. * and ** indicate significant differences at P < 0.05 and P < 0.01, respectively. Panel (E) shows a fingerprint gallery plot of VOCs from a single biological replicate. The colour gradient represents the normalized relative ion intensity (arbitrary units), with blue to red indicating low to high relative abundance, respectively. The rectangular box in panel E highlights VOCs that exhibited significant differences between the control and ALA groups. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Two-dimensional and three-dimensional maps of VOCs were constructed using the Gallery and Reporter plug-ins. As shown in Fig. S2A,C there is good stability between samples. Changes in VOCs are determined by comparative analysis, with SO1 as the reference and other plots deducted from the reference. In instances where the content of VOCs remained constant, the background of the deduction was set to white, with red denoting concentrations that exceeded the reference level and blue indicating concentrations that fell below it. The results indicated the presence of variations in the concentration of VOCs in Yellow River carp following ALA intervention (Fig. S2B). To ensure the robustness of the findings and enhance the precision of the results, a multivariate statistical analysis was conducted (Fig. 4A–C). The results indicated that the VOCs in the two treatments were effectively distinguished. With VIP > 1 and P < 0.05, a total of 22 VOCs were found to be significantly altered in Yellow River carp after ALA intervention, of which 12 were increased significantly and 10 were significantly decreased (Fig. 4D). These differential VOCs were categorized into four groups: 5 alcohols, 10 aldehydes, 6 ketones, and 1 furan (Table S4). Aldehydes have a lower sensory threshold. Some aldehydes, such as hexanal, heptanal, nonanal, and decanal are recognized fishy substances, reducing the generation and accumulation of these aldehydes can effectively regulate the fishy odor of fish. The threshold value for unsaturated alcohol is usually much lower than that for saturated alcohol, which significantly influences food flavors. For instance, oct-1-en-3-ol is a key substance responsible for imparting an earthy aroma to aquatic products with characteristics reminiscent of a mushroom scent (Wang et al., 2021; Zhang et al., 2019). Ketones typically emit floral fragrances, which increase gradually with carbon chain extension. The main VOCs in fermented golden pomfret contain 6-methyl-5-hepten-2-one and 2-nonanone, which give it the aroma of cheese, fruit, and lemongrass (Chen et al., 2021; Li et al., 2022). Our previous study showed that VOCs, including pent-1-en-3-ol, oct-1-en-3-ol-D, and 2-pentyl-furan, accumulate in large quantities in fish at later life stages following long-term feeding of high-LA diets (unpublished). Among these, key VOCs such as pent-1-en-3-ol and oct-1-en-3-ol are derived from the oxidation of n-3 and n-6 polyunsaturated fatty acids, respectively. In this study, the contents of oct-1-en-3-ol-D, 2-pentyl furan, Heptanal-D, Heptanal-M, (E)-hept-2-enal-M, 2-Octanone-M, (E)-hept-2-enal-D in the muscle were significantly reduced under the intervention of high ALA. These VOCs mainly have a fishy and earthy odor. The contents of pent-1-en-3-ol, (Z)-4-heptenal, (E, E)-2,4-heptadienal with fresh, plant, and fruit aromas were significantly increased, although the total amount of aldehydes and alcohols was decreased by ALA intervention (Fig. 3D). Fingerprint mapping revealed the specific alterations in VOCs following ALA intervention (Fig. 3E). This suggests that high ALA intervention can optimize the VOCs of Yellow River carp in plant-derived diets to a certain extent. Correlation analysis of the key differential VOCs with FAs revealed that those associated with ALA and n-3 PUFAs exhibited pleasant odors (sweet, floral, and fruity), while those associated with LA and n-6 PUFAs exhibited unpleasant odors (pungent) (Fig. 5 and Table S5).

Fig. 4.

Fig. 4

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Multivariate analysis of the VOCs and differential VOCs in the muscle of the Yellow River carp after ALA intervention. Principal component analysis (PCA) (A), partial least squares discriminant analysis (PLS-DA) (B), orthogonal PLS-DA (C) score plots based on flavoromics data. The number of significantly different VOCs (D). Heatmap analysis of differential VOCs (E). VOC enrichment is encoded in the heat map from low (blue) to high (red) and shows similar variations clustered together. As shown in the colored group on the right side of the heat map. Panel (E) shows a cluster heatmap of VOCs from a single biological replicate. The colour scale represents the Z-score of normalized intensity, with blue to red indicating content from low to high. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 5.

Fig. 5

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Correlation analysis between key differential VOCs and fatty acids. Colour scale indicates the strength of correlation, ranging from positive (red) to negative (blue). Data presented as mean ± SEM (n = 4). *, ** and *** represent significant differences with P < 0.05, P < 0.01 and P < 0.001, respectively. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

3.4. Lipid composition was changed by ALA intervention

Research has shown that the composition of VOCs is related to lipids and proteins (Chen et al., 2023; Chu et al., 2023). In this study, ALA intervention did not significantly change the protein content and amino acid compositions of Yellow River carp muscle but significantly altered its FA composition. As key precursors for VOCs, UFAs are more often attached to the side chains of GL, GP, and SM molecules rather than in the form of free fatty acids (Jia et al., 2021). In comparison with glycerolipids (GLs), glycerophospholipids (GPs) are abundant in PUFA, which play an instrumental role in the composition of VOCs in meat (Zhang et al., 2020). Xiong et al. (2022) ascertained that PC (18:2/18:0) was the paramount factor in the formation of aldehydes, alcohols, and ketones in yak meat. Feng et al. (2022) found that the increase in aldehydes was primarily attributable to the oxidation of unsaturated fatty acids (C18:1, C16:1, C18:2, and C20:4) in ePE and ePC. Wu et al. (2019) found that PE(16:0/18:2), PE(18:0/18:2), PE(18:0/20:4), PE(16:0/22:6), and PE(18:0p/22:6) were the most important factors in the formation of hexanal, heptanal, nonanal, 2- pentylfuran, and 2-octanone as important precursors. Lipidomics can be used to detect and characterize lipid types and content (Li et al., 2021). To further investigate the changes in lipid composition during ALA intervention, the muscle lipids of Yellow River carp after ALA intervention were examined by LC-MS assay. Fig. 6 shows the amount and percentage of lipids in both groups. A total of 1188 different types of lipids (28 subclasses in 5 classes) were identified in muscles using LC-MS, including 748 glycerophospholipids (GPs), 350 glycerolipids (GLs), 75 sphingolipids (SPs), 6 saccharolipids (SLs), and 9 FAs (Fig. 6A). The subclasses with the highest number were, in order, TG (25.51 %), PC (23.32 %), PE (14.48 %), and phosphatidylserine (PS, 6.31 %) (Fig. 6B). Fish lipid classes principally comprise TG, PC, PE, PS, and phosphatidylinositol (PI) (Ruizlopez et al., 2015), and the total station ratio of TG, PC, and PE in Yellow River carp muscle surpasses 70 % (Fig. 6C). To observe the lipid changes in Yellow River carp after ALA intervention with greater accuracy, we performed multivariate statistical analysis. The results of PCA showed (Fig. 7A) that the lipids of the two groups of Yellow River carp were effectively differentiated, with the first principal component explaining 41.9 % of the variance, and the second principal component explaining 31.1 % of the variance. Consistent results were obtained from PLS—DA and OPLS-DA analyses (Fig. 7B, C). Based on the criteria of VIP > 1 and P < 0.05, the total number of differentiated lipids in Yellow River carp after ALA intervention was 143, including 88 lipids up-regulated and 55 lipids down-regulated (Fig. 7D and Table S6). The clustering heatmap visualizes information about the different substances. The lipids increased in muscle after ALA intervention consisted of 27 PCs, 22 PEs, 14 phosphatidylserines (PSs), 9 TGs, 3 lyso-phosphatidylcholines (LPCs), 2 lyso-phosphatidylethanolamine (LPEs), 1 PI, 1 phosphatidylglycerol (PG), 1 dimethylphosphatidylethanolamine (dMePE), 1 sphingomyelin (SM), and 1 FA (Fig. 7E, G). In contrast, downregulated lipids included 14 PCs, 11 PEs, 8 PSs, 7 LPCs, 3 LPEs, 3 phosphatidic acids, 2 PIs, 2 dMePEs, 2 (O-acyl)-1-hydroxy fatty acids, 1 SM, and 1 FA (Fig. 7F). These differential lipids were predominantly present in GPs (89 %). Meanwhile, the KEGG enrichment results indicated that the glycerophospholipid metabolism pathway was mainly activated after ALA intervention (Fig. 8). This indicates that GPs are the core target of ALA intervention. Integrating the abundance and structural characteristics of the major lipid molecules, dietary ALA intake was found to induce remodeling of the muscle phospholipid membrane (rather than affecting protein metabolism), characterized by the specific enrichment of n-3 polyunsaturated fatty acids in membrane phospholipids such as phosphatidylethanolamine. This compositional alteration increased the availability and susceptibility of membrane lipids as substrates for oxidation, thereby steering the flux of lipid oxidation and shaping the final profile of volatile flavor compounds. This process effectively enhanced the nutritional value of Yellow River carp and optimized its VOC composition. To our knowledge, the linkage between dietary ALA intake and changes in lipid molecules and structure, and its subsequent association with the formation of the volatile flavor profile, has not been explicitly reported in other species—where related research often focuses primarily on lipid metabolism. This study is the first to elucidate the linkage between muscle lipid molecular composition and volatile flavor substances in Yellow River carp under ALA intervention, providing novel insights for the fields of aquatic nutrition and food science.

Fig. 6.

Fig. 6

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Lipid compound composition in the muscle of the Yellow River carp after ALA intervention. Number (A), percentage (B), and relative content (C) of lipid compound categories. GL: Glycerolipids, GP: Glycerophospholipids, SL: Saccharolipids, SP: Sphingolipids; OAHFA: (O-acyl)-1-hydroxy fatty acid, TG: Triglyceride, DG: Diglyceride, MG: Monoglyceride, PC: Phosphatidylcholine, PE: Phosphatidylethanolamine, PS: Phosphatidylserine, LPC: Lyso-phosphatidylcholine, PG: Phosphatidylglycerol, PA: Phosphatidic acid, dMePE: Dimethylphosphatidylethanolamine, LPE: Lyso-phosphatidylethanolamine, PEt: Phosphatidylethanol, CL: Cardiolipin, LdMePE: Lysodimethylphosphatidylethanolamine, LPEt: Lyso-phosphatidyl ethanol, PMe: Phosphatidylmethanol, PI, PIP, PIP2: Phosphatidylinositol, CerG1: Monogylcosylceramide, MGDG: Monogalactosyldiacylglycerol, SM: Sphingomyelin, Cer: Ceramides, LSM: Lysosphingomyelin, phSM: Sphingomyelin (phytosphingosine), So: Sphingosine. Data presented as mean ± SEM (n = 4). * and ** represent significant differences with P < 0.05 and P < 0.01. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 7.

Fig. 7

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Multivariate analysis of the lipids and differential lipids in the muscle of the Yellow River carp after ALA intervention. Principal component analysis (PCA) (A), partial least squares discriminant analysis (PLS-DA) (B), orthogonal PLS-DA (C) score plots based on flavoromics data. The number of significantly different lipid components (D). Panels (D, E, F) display hierarchical clustering heatmaps of lipid. The Z-score, calculated from the normalized intensity across all samples, is shown in colour, scaling from blue (low) to red (high). Red triangles mark lipids, including PE(22:6) and PC(20:5), that were identified as significant contributors to the observed metabolic phenotype. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

Fig. 8.

Fig. 8

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Enrichment analysis of the KEGG pathway of different lipids in the muscle of Yellow River carp after ALA intervention. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

4. Conclusions

In summary, the study demonstrated that the ALA intervention strategy resulted in a significant increase in the n-3 LC-PUFA content of the Yellow River carp muscle. Lipidomic analysis of Yellow River carp revealed a distinct positional distribution of fatty acids in glycerolipids: SFAs were predominantly esterified at the sn-1 position of TGs, whereas ALA was primarily located at the sn-3 position of TGs. In phospholipids, ARA and DHA showed a strong preference for the sn-2 position of PEs. Dietary ALA intervention significantly promoted the incorporation of DHA at the sn-2 position of PEs. Conversely, it reduced the esterification of SFAs at the sn-1 position of both TGs and PEs, as well as the incorporation of MUFAs in TGs and (PCs. These ALA-induced alterations in the composition and positional distribution of fatty acids enhanced the nutritional quality of the fish lipids. Volatile metabolomics analyses revealed that the ALA intervention led to a substantial reduction in the levels of earthy substances (e.g., oct-1-en-3-ol-D) and a concomitant increase in the levels of aromatic substances (e.g., pent-1-en-3-ol). Among the diverse array of lipids influenced by dietary ALA, glycerophospholipids exhibited the most pronounced alterations, accompanied by the activation of their metabolic pathways in response to ALA. To the best of our knowledge, this is the first report on the exploration of the relationship between lipid molecular composition and volatile flavor substances in Yellow River carp muscle under ALA intervention, providing new insights into the field of fish nutrition and food science. Nevertheless, the specific regulatory mechanisms at the transcriptional and protein levels require further elucidation. Future studies integrating transcriptomics and proteomics will be crucial to directly validate the roles of key enzymes in the observed lipid metabolic shifts and VOC generation.

CRediT authorship contribution statement

Zijie He: Writing – original draft, Investigation, Formal analysis, Data curation. Junli Wang: Writing – review & editing, Funding acquisition. Yun Wei: Investigation, Data curation. Xiao Yan: Writing – review & editing, Funding acquisition. Yuanyou Li: Writing – review & editing, Supervision. Guoxing Nie: Writing – review & editing, Methodology, Funding acquisition, Conceptualization. Dizhi Xie: Writing – review & editing, Methodology, Conceptualization.

Ethical statement

The protocols for animal care and handling adopted in the present study were approved by the Institutional Animal Care and Use Committee of Henan Normal University. All efforts were made to minimize the suffering of common carp.

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.

Acknowledgments

This work was supported by the National Natural Science Foundation of China (U22A20532, 32072991, 32373149, 32373142); Special Fund for Henan Agriculture Research System (HARS-22-16-S). All authors have approved the final version of this manuscript and declared that no competing interests exist.

Footnotes

Appendix A

Supplementary data to this article can be found online at https://doi.org/10.1016/j.fochx.2025.103264.

Appendix A. Supplementary data

Supplementary material 1
mmc1.docx (201KB, docx)
Supplementary material 2
mmc2.docx (1.7MB, docx)
Supplementary material 3
mmc3.docx (22.3KB, docx)
Supplementary material 4
mmc4.docx (20.6KB, docx)
Supplementary material 5
mmc5.docx (29.1KB, docx)
Supplementary material 6
mmc6.docx (24.7KB, docx)
Supplementary material 7
mmc7.docx (23.9KB, docx)
Supplementary material 8
mmc8.docx (21.6KB, 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 1
mmc1.docx (201KB, docx)
Supplementary material 2
mmc2.docx (1.7MB, docx)
Supplementary material 3
mmc3.docx (22.3KB, docx)
Supplementary material 4
mmc4.docx (20.6KB, docx)
Supplementary material 5
mmc5.docx (29.1KB, docx)
Supplementary material 6
mmc6.docx (24.7KB, docx)
Supplementary material 7
mmc7.docx (23.9KB, docx)
Supplementary material 8
mmc8.docx (21.6KB, 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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