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. 2026 Apr 17;35:103879. doi: 10.1016/j.fochx.2026.103879

Comparative metabolomic profiling and bioactivities of essential oils from different organs and geographic origins of Curcuma kwangsiensis

Fengke Lin a,⁎, Rui Zhao a, Zihao Liu a, Qun Zhou a, Qianyun Wang b, Edward J Kennelly c, Jun Feng b,⁎, Renchuan Hu b,⁎
PMCID: PMC13122245  PMID: 42058385

Abstract

Curcuma kwangsiensis is valued for food and medicinal applications, but its essential oils (EOs) remain poorly characterized across organs and geographic origins. This study aimed to analyze 15 EOs of five plant parts collected from three geographic origins. GC–MS characterized 88 volatiles, primarily sesquiterpenes, with significant variation across organs and locations. PCA and PLS-DA distinguished samples by organ and origin, highlighting 31 differential metabolites, including curzerene and trans-β-elemenone. MTS assays showed Lingshan EOs had potent cytotoxicity against K562 and HepG2 cells (IC50 = 10.82–18.53 μg/mL), NO inhibition assays revealed strong anti-inflammatory activity of Wuming tuberous root EO (IC50 = 18.31 μg/mL), and DPPH/ABTS assays indicated superior antioxidant capacity of tuberous root and rhizome EOs (IC50 = 0.25–2.42 mg/mL). Spearman correlation linked key compounds, including trans-β-elemenone and nootkatone, to observed bioactivities. These findings demonstrate that organ and origin influence EO chemistry and bioactivities, supporting their potential in functional foods and nutraceuticals.

Keywords: Curcuma, Chemical composition, Anticancer, Anti-inflammatory, Antioxidant, Untargeted metabolomics, Chemometrics

Highlights

  • •

    EO composition varied across plant parts and origins of Curcuma kwangsiensis.

  • •

    Thirty-one differential metabolites were identified as potential chemical markers.

  • •

    Plant parts and origins markedly influenced EO biological activities.

  • •

    Tuberous root and rhizome EOs showed superior bioactivities.

  • •

    Findings support targeted selection of C. kwangsiensis EOs for nutraceutical use.

1. Introduction

The genus Curcuma L. (Zingiberaceae) comprises more than 120 species widely distributed in tropical and subtropical regions of Asia, many of which are traditionally used for food and medicinal purposes (Sasikumar, 2005; Záveská et al., 2012). The characteristic aroma and health-promoting properties of Curcuma species are largely attributed to bioactive metabolites, particularly essential oils (EOs) (Sun et al., 2017; Yang et al., 2025). Curcuma EOs are rich in terpenoids and exhibit diverse biological activities, including antioxidant, anti-inflammatory, and antimicrobial effects, highlighting their potential applications in food and medicinal food products (Dosoky & Setzer, 2018; Hu et al., 2017). Accordingly, increasing research efforts have focused on elucidating the chemical composition and functional properties of Curcuma EOs (Xiang et al., 2018).

Curcuma kwangsiensis S. K. Lee & C. F. Liang is an endemic species native to the Guangxi Zhuang Autonomous Region of China and has been recognized as one of the region's top ten medicinal plants (Liu et al., 2023). Our ethnobotanical investigations demonstrate that the rhizomes of C. kwangsiensis are traditionally used as spices in soups and porridges. In addition, aqueous decoctions or ethanol infusions prepared from the rhizomes and tuberous roots are traditionally consumed to alleviate disorders such as food stagnation, abdominal distension, and gastric discomfort (Commission of Chinese Herbal Medicine, 1999). These traditional practices suggest that C. kwangsiensis possesses considerable potential as a functional food resource. Given that EOs represent one of its major bioactive fractions, systematic investigation of EO composition is crucial for supporting its traditional edible and health-related applications. Previous studies have primarily focused on the chemical composition and biological activities of rhizome EOs of C. kwangsiensis, reporting abundant sesquiterpenoids such as germacrone and trans-β-elemene (Xiang et al., 2018; Zhang, Yang, Huang, et al., 2017). Although the EO compositions of tuberous roots, fibrous roots, and leaves have been reported (Huang et al., 2020; Luo, 2014), the bioactivities of these EOs remain unexplored. In addition, the chemical composition and biological activities of flower EOs of C. kwangsiensis have not yet been reported.

The chemical composition and bioactivity of plant EOs can be markedly affected by factors such as plant part and geographic origin (Gururani et al., 2022; Jena et al., 2017). Although comparative analyses of EOs from different organs (leaves, rhizomes, and tuberous roots) of C. kwangsiensis have been conducted (Luo, 2014), a comprehensive chemical comparison between above-ground and underground plant parts remains lacking, and variations in EO bioactivities among different plant organs have not yet been elucidated. The influence of geographic origin on EO composition and bioactivity has been previously investigated for C. kwangsiensis rhizomes (Zhang, Yang, Huang, et al., 2017); however, corresponding studies on other organs remain limited. Based on previous reports (Wang et al., 2015; Zhang, Yang, Huang, et al., 2017) and our ethnobotanical surveys, Wuming District, Lingshan County, and Fumian District in Guangxi are among the major natural distribution areas of C. kwangsiensis and important sources for its traditional use. Comprehensive investigations of the chemical composition and bioactivities of C. kwangsiensis EOs from these regions are therefore essential for the rational selection of promising resources for nutraceutical applications.

Untargeted metabolomic analysis based on gas chromatography–mass spectrometry (GC–MS) has proven to be a powerful approach for comprehensive characterization of plant EO chemical profiles, including those of Curcuma species such as C. longa (Huang et al., 2023; Pantharos et al., 2022). When combined with chemometric methods, such as principal component analysis (PCA) and partial least squares discriminant analysis (PLS-DA), metabolomics enables the discrimination of samples and identification of key metabolites responsible for the chemical variation (Lin et al., 2023; Lin & Long, 2023). However, to our knowledge, an integrated metabolomics-chemometrics strategy has not yet been applied to systematically investigate EO variation in C. kwangsiensis.

In the present study, the EO compositions of five plant organs of C. kwangsiensis collected from three geographic locations were comprehensively characterized using GC–MS-based nontargeted metabolomic analysis. Differential compounds associated with variations in plant organ and geographic origin were identified by chemometrics. In addition, comparative cytotoxic, anti-inflammatory, and antioxidant assays were conducted to evaluate variations in the bioactivities of the EOs. Therefore, this study aimed to compare the chemical constituents and biological activities of C. kwangsiensis EOs from different organs and geographic origins using metabolomics and chemometric analysis. These findings offer new insights into the chemical diversity and functional potential of C. kwangsiensis EOs, highlighting the importance of selecting oils from specific plant organs and geographic origins for their potential use in functional food and nutraceutical applications.

2. Materials and methods

2.1. Chemicals and reagents

Ultra-pure water for EO extraction was obtained from a Milli-Q purification system (Millipore Corporation, Billerica, MA, USA). Anhydrous sodium sulfate (Na2SO4, 99.0%) was purchased from Sinopharm Chemical Reagent Co., Ltd. (Shanghai, China). For GC–MS analysis, HPLC-grade n-hexane (99.0%) was obtained from CNW Technologies (Düsseldorf, Germany), and the C7-C40 saturated alkane standard mixture was purchased from Sigma-Aldrich (St. Louis, MO, USA). For bioactivity assays, dimethyl sulfoxide (DMSO, 99.7%) was produced by Sangon Biotech (Shanghai, China). Dulbecco's modified Eagle medium (DMEM), Roswell Park Memorial Institute (RPMI)-1640 medium, and fetal bovine serum (FBS) were purchased from Boehringer Ingelheim (Ingelheim, Germany). Doxorubicin (DOX, 98.0%) was obtained from Meilunbio (Dalian, China). 3-(4,5-Dimethylthiazol-2-yl)-5-(3-carboxymethoxyphenyl)-2-(4-sulfophenyl)-2H-tetrazolium (MTS) reagent was procured from Promega (Madison, WI, USA). Griess reagent, lipopolysaccharide (LPS, ≤100%), and NG-methyl-L-arginine acetate salt (L-NMMA, ≥98.0%) were purchased from Sigma-Aldrich (St. Louis, MO, USA). 2,2-Diphenyl-1-picrylhydrazyl (DPPH, ≥98.5%), 2,2′-azino-bis(3-ethylbenzothiazoline-6-sulfonic acid) (ABTS, 99.5%), and analytical-grade ethanol (≥99.7%) and methanol (≥99.5%) were obtained from Macklin (Shanghai, China). l-ascorbic acid (LAA, 99.0%) and potassium persulfate (K2S2O8, 99.0%) were sourced from Aladdin (Shanghai, China).

2.2. Plant materials

Plant materials of C. kwangsiensis were collected from three locations in Guangxi, China: Fumian District (FM; 22°31′28.55“ N, 109°56’46.50” E, 118 m), Lingshan County (LS; 22°17′18.52“ N, 108°56’0.20” E, 27 m), and Wuming District (WM; 23°18′10.21“ N, 108°30’6.71” E, 148 m). Flowers (F) were harvested in May 2024, corresponding to their flowering period, whereas underground organs, including tuberous roots (RT), fibrous roots (FR), and rhizomes (RH), were collected in November 2024 at their mature stage. Leaves (L), which are present throughout the growing season, were also collected in November to align with the sampling of underground organs. Botanical identification was performed by Renchuan Hu at the Guangxi Key Laboratory of Traditional Chinese Medicine Quality Standards, Guangxi Institute of Chinese Medicine & Pharmaceutical Science (GICMPS). Voucher specimens were deposited at GICMPS under the accession numbers FJCKFM2405001, FJCKLS2405001, and FJCKWM2405001. After collection, the plant materials were rinsed with deionized water to remove surface dust and soil, cut into small pieces, air-dried in the shade, and ground to pass through a 40-mesh sieve. The resulting powders were stored at −20 °C until further analysis.

2.3. EO extraction

EOs from C. kwangsiensis samples were extracted by hydrodistillation using a Clevenger-type apparatus according to the protocol described in the Chinese Pharmacopoeia (National Pharmacopoeia Commission, 2020), with minor modifications. Briefly, approximately 50 g of each powdered sample was placed in a 2000 mL round-bottom flask containing 1000 mL of ultra-pure water, along with several glass beads to prevent bumping. The mixture was subjected to ultrasonic pretreatment for 30 min and subsequently heated using an electric heating mantle (Beijing Ever-Bright Medical Treatment Instrument Co., Ltd., Beijing, China). Based on previous reports (Luo et al., 2021; Sharma et al., 2019) and preliminary experiments, a hydrodistillation duration of 6 h was applied for each sample to ensure complete EO extraction. The collected EOs were dried over anhydrous Na2SO4 and stored in the dark at 4 °C prior to chemical composition and bioactivity analyses. EO yield was calculated as the weight of EO (mg) relative to the dry weight of the plant material (g) and expressed both as a percentage (w/w, %) and in mg/g of dry weight, with 1% corresponding to 10 mg/g.

2.4. GC-MS-based metabolomic analysis

The chemical constituents of C. kwangsiensis EOs were analyzed using a GC-2030 gas chromatograph coupled with a QP2020 NX mass spectrometer (Shimadzu, Kyoto, Japan), as described by Lin et al. (2025). Before analysis, each EO sample was diluted with n-hexane to a final concentration of 1 mg/mL. A pooled quality control (QC) sample was prepared by mixing equal aliquots (30 μL) from each EO solution. Chromatographic separation was achieved on a DB-5 MS capillary column (30 m × 250 μm i.d., 0.25 μm film thickness; J&W Scientific, Folsom, USA), with an injection volume of 2 μL. Helium was used as the carrier gas at a constant flow rate of 1.0 mL/min, and the inlet purge flow was set to 3 mL/min. The oven temperature program was as follows: an initial temperature of 50 °C held for 1 min, increased at 8 °C/min to 310 °C, and held for 11.5 min. The injector, transfer line, and ion source temperatures were maintained at 280 °C, 280 °C, and 230 °C, respectively. Mass spectra were acquired in electron impact (EI) ionization mode at 70 eV, scanning the m/z range of 50–500 at 12.5 spectra/s. Raw GC–MS data were processed using ChromaTOF software (version 4.3×, LECO Corporation, St. Joseph, MI, USA) for peak detection, deconvolution, retention time alignment, baseline correction, and peak area integration. Linear retention indices (LRIs) were calculated using a homologous series of n-alkanes (C7-C40) analyzed under identical chromatographic conditions. Compound identification was performed by matching experimental mass spectra with the NIST 23 mass spectral library, and compounds with similarity scores above 700 (out of 1000) were retained. The annotations were further verified by comparing calculated LRIs with literature values (Adams, 2017) and data reported in the NIST Chemistry WebBook (https://webbook.nist.gov/chemistry/). Compounds were accepted if the difference between the calculated and reported LRIs was less than 30 (Li et al., 2025). The relative abundance of each identified metabolite was expressed as the percentage of its peak area in the total ion chromatogram.

2.5. Cytotoxic evaluation

The cytotoxic activities of C. kwangsiensis EOs were evaluated in three human cancer cell lines, K562 (leukemia), A549 (lung carcinoma), and HepG2 (hepatocellular carcinoma) using the MTS assay, as described by Wang et al. (2010). Cells were cultured under standard conditions (37 °C, 5% CO2) in RPMI-1640 or DMEM supplemented with 10% FBS. For cell viability assessment, cells were seeded into 96-well plates at densities of 5.0 × 103 to 8.0 × 103 cells per well and allowed to adhere for 12–24 h. The culture medium was then replaced with fresh medium containing C. kwangsiensis EOs at a final concentration of 100 μg/mL. DOX (5 μg/mL) and 0.1% DMSO were included as the positive and negative controls, respectively. All treatments were performed in triplicate. After 48 h of incubation, the culture medium was removed, and 20 μL of MTS reagent together with 100 μL of fresh culture medium was added to each well. Following an additional incubation for 2–4 h, absorbance was measured at 492 nm using a MultiSkan FC microplate reader (Thermo Fisher Scientific, Waltham, MA, USA). Growth inhibition was calculated by comparing the absorbance of treated wells with that of the negative control. Samples exhibiting more than 50% inhibition at 100 μg/mL were selected for subsequent determination of half-maximal inhibitory concentration (IC50) values using GraphPad Prism 10.5.0 (GraphPad Software Inc., San Diego, CA, USA).

2.6. Anti-inflammatory assessment

The anti-inflammatory capacity of C. kwangsiensis EOs was assessed using LPS-stimulated RAW 264.7 macrophages according to the procedure described by Zhang et al. (2021). Cells were seeded into 96-well plates at a density of 8 × 104 cells per well and cultured in DMEM for 24 h under standard conditions (37 °C, 5% CO2). After incubation, the culture medium was removed, and the test EOs were added at a final concentration of 100 μg/mL in triplicate. Nitric oxide (NO) production was induced by the addition of LPS (1 μg/mL) to each well. L-NMMA (12.4 μg/mL) and 0.1% DMSO were used as the positive and negative controls, respectively. After 24 h of incubation, 50 μL of the culture supernatant was collected from each well and mixed with 100 μL of Griess reagent to quantify NO production by measuring absorbance at 570 nm. Cell viability was evaluated using the MTS assay by adding 20 μL of MTS reagent to the remaining cells in each well to exclude false-positive inhibition caused by cytotoxic effects. EOs exhibiting cytotoxicity were further subjected to serial two-fold dilutions until non-cytotoxic concentrations were obtained. For EO samples that inhibited NO production by more than 50% without inducing cytotoxicity in RAW 264.7 cells, their IC50 values were subsequently determined using GraphPad Prism 10.5.0.

2.7. Antioxidant activity

The antioxidant potential of C. kwangsiensis EOs was evaluated using both DPPH radical scavenging and ABTS+• decolorization assays (Lin et al., 2025). For the DPPH assay, a fresh 0.1 mM DPPH solution was prepared in methanol. In 96-well plates, 20 μL of EO solutions (final concentrations: 0.2–6.4 mg/mL) were mixed with 180 μL of DPPH solution in triplicate. For the ABTS assay, the ABTS+• working solution was generated by reacting 7 mM ABTS with 2.45 mM K2S2O8 and incubating the mixture in the dark for 12–16 h. Prior to use, the solution was diluted with ethanol to an absorbance of 0.70 ± 0.02 at 734 nm. EO samples (20 μL; final concentrations: 0.2–1.6 mg/mL) were mixed with 180 μL of ABTS+• solution in triplicate. LAA (10 μg/mL) served as the positive control, and DMSO (20 μL) as the negative control. All reactions were incubated at 37 °C in the dark for 30 min. Absorbance was measured at 517 nm (DPPH) and 734 nm (ABTS). Radical scavenging activity was calculated and IC50 values were determined using GraphPad Prism 10.5.0.

2.8. Statistical analysis

Chemometric analyses, including PCA and PLS-DA, were performed using SIMCA-P 14.1 (Umetrics, Umeå, Sweden). The PLS-DA models were validated using 7-fold cross-validation to evaluate their predictive ability. Permutation tests (200 permutations) were performed to assess the robustness of the models and to exclude potential overfitting. Variable importance in projection (VIP) scores for detected features were derived from the PLS-DA models. One-way ANOVA was applied to calculate p-values using IBM SPSS Statistics 27.0 (IBM Corp., Chicago, IL, USA). EO Metabolites with VIP > 1.5 and p < 0.05 were considered significantly different (Ren et al., 2025). For bioactivity evaluations, data were presented as mean ± standard deviation (SD). IC50 values for cytotoxicity and anti-inflammatory activity were presented in μg/mL, while mg/mL was used for antioxidant activity due to its relatively lower potency. Statistical differences among groups were assessed by one-way ANOVA followed by Duncan's multiple range test in SPSS. A p-value <0.05 was considered statistically significant.

3. Results and discussion

3.1. EO yield analysis

Many Curcuma species, such as C. zedoaria and C. sylvatica, are recognized as important sources of EOs (Angel et al., 2014). In the present study, hydrodistillation of C. kwangsiensis yielded EO contents ranging from 0.52% to 2.22% (5.2–22.2 mg/g of dry weight) (Fig. 1). Significant organ-dependent differences in EO yields were observed, with RH exhibiting the highest yields (1.88–2.22%, 18.8–22.2 mg/g), followed by F (0.65–1.13%, 6.5–11.3 mg/g), L (0.70–0.90%, 7.0–9.0 mg/g), FR (0.52–1.06%, 5.2–10.6 mg/g), and TR (0.48–0.68%, 4.8–6.8 mg/g) (Fig. 1). Similar organ-specific variations in EO accumulation have been reported in other Curcuma species, such as C. longa (Bansal et al., 2002), suggesting that tissue-dependent EO distribution is a common characteristic within the genus. Considerable variation in EO yields, both within C. kwangsiensis and among different Curcuma species, has been observed when comparing the present results with those reported in the literature using hydrodistillation (Kumar et al., 2016; Zhang, Yang, Chen, et al., 2017). For instance, the RH yield obtained in the present study was lower than previously reported values for C. kwangsiensis (3.05%) (Zhang, Yang, Chen, et al., 2017) and C. longa (2.60%) (Kumar et al., 2016), but higher than those reported for C. aromatica (1.48%) (Choochote et al., 2005) and C. aeruginosa (0.40%) (Jena et al., 2017). These results indicate pronounced inter- and intraspecific variability in EO yields within the genus Curcuma, which might be attributed to several factors, including genetic background, environmental conditions, and plant developmental stage (Barra, 2009). In addition to hydrodistillation, advanced techniques such as microwave-assisted, supercritical fluid, and enzyme-assisted extraction have been applied to improve EO recovery from C. kwangsiensis (Jiang et al., 2023). For example, Jiang et al. (2023) reported that enzyme-assisted extraction of C. kwangsiensis rhizomes achieved EO yields of up to 5.24%, significantly higher than that obtained by hydrodistillation (2.17%). These techniques generally require specialized equipment, higher operational costs, and more complex optimization procedures. In contrast, hydrodistillation remains one of the most widely used techniques for EO extraction due to its operational simplicity, reproducibility, and compatibility with the extensive literature on Curcuma species (Dosoky & Setzer, 2018). Nevertheless, some thermal degradation of volatile compounds may occur during prolonged hydrodistillation (Sun et al., 2025), highlighting the need for further studies on the effects of different extraction methods on the chemical composition and bioactivities of C. kwangsiensis EOs.

Fig. 1.

Fig. 1

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EO yields of different plant parts of C. kwangsiensis across three collection locations. L: leaves, F: flowers, TR: tuberous roots, FR: fibrous roots, RH: rhizomes, FM: Fumian District, LS: Lingshan County, WM: Wuming County. Different letters above the columns indicate statistically significant differences among samples (p < 0.05) based on Duncan's multiple range test.

In addition to organ-related differences, EO yields of the same plant part varied among geographic origins, consistent with observations in C. longa (Setzer et al., 2021). Among the three collection sites, rhizomes from FM exhibited the highest EO yield, whereas samples from WM showed the greatest floral EO production (Fig. 1). These spatial variations suggest that environmental factors play an important role in regulating EO biosynthesis and accumulation in C. kwangsiensis. Further studies are therefore warranted to identify key environmental determinants of EO yield and to inform cultivation strategies to maximize EO production. As EOs are highly valued for their aroma and flavor, their preferential accumulation in rhizomes supports the traditional use of Curcuma rhizomes as flavoring and aromatic ingredients in food (Sasikumar, 2005).

3.2. Metabolite analysis using GC-MS-based metabolomics

GC–MS-based metabolomic profiling enabled the identification of 88 EO constituents across all C. kwangsiensis samples, accounting for 77.48–92.81% of the total EO content (Table S1). The remaining unidentified fraction (7.19–22.52%), particularly in FM-RH EO, represents compounds lacking reliable annotation based on their mass spectra and retention indices. This may be attributed to the presence of structurally novel compounds or limitations in current mass spectral library coverage. The identified metabolites were categorized into eight chemical classes, including oxygenated sesquiterpenes, sesquiterpene hydrocarbons, oxygenated monoterpenes, monoterpene hydrocarbons, furans, ketones, diterpenes, and other compounds. Among these groups, oxygenated sesquiterpenes were the most abundant, with relative contents ranging from 21.90% to 63.48%, followed by sesquiterpene hydrocarbons (18.60–60.50%), oxygenated monoterpenes (0.59–21.79%), and monoterpene hydrocarbons (0.30–17.96%) (Table S1). This terpenoid-rich chemical profile is consistent with previous reports on other Curcuma species, such as C. zanthorrhiza and C. glans (Dosoky & Setzer, 2018), highlighting a genus-wide tendency to accumulate sesquiterpenes and monoterpenes as dominant EO constituents. At the individual metabolite level, five major constituents, including trans-β-elemenone (9.85–42.92%), trans-β-elemene (6.48–11.59%), curzerene (2.95–9.17%), α-humulene (0.23–10.45%), and (E)-β-caryophyllene (0.44–14.90%), were detected in all samples (Table S1). Among them, trans-β-elemenone, trans-β-elemene, curzerene, and (E)-β-caryophyllene have been widely reported as dominant or characteristic constituents in several Curcuma species, including C. longa, C. aurantiaca, and C. oligantha (Dosoky & Setzer, 2018). The recurrent occurrence of these compounds across multiple species highlights their potential chemotaxonomic significance as diagnostic markers for the genus Curcuma, particularly in distinguishing Curcuma EOs from those of other genera. The relative abundance of trans-β-elemenone in rhizome EOs across different Curcuma species varied markedly (5.2–65.0%) (Dosoky & Setzer, 2018), further indicating its potential utility for interspecific discrimination within the genus.

The relative abundances of major EOs varied substantially among different plant parts and geographical origins. For example, in LS samples, trans-β-elemenone displayed significant variability, reaching its highest level in LS-TR EO (42.92%) but decreasing sharply to 10.19% in LS-F EO. In comparison, its contents in FM-TR and WM-TR EOs were 28.70% and 17.95%, respectively. Such pronounced quantitative differences highlight the strong influence of plant organ specificity and environmental origin on the biosynthesis and accumulation of EO metabolites in C. kwangsiensis. In addition, other factors, such as harvest season or month, may potentially influence the chemical composition and bioactivities of C. kwangsiensis EOs, warranting further exploration. To our knowledge, this study provides the first comprehensive GC–MS-based EO profiling of C. kwangsiensis across different plant parts and geographic origins. Similar organ- and site-dependent chemical variations have been reported in other Curcuma species, including C. aeruginosa (Nugraheni et al., 2024; Simoh et al., 2018) and C. longa (Quemel et al., 2021), suggesting that significant chemical variability associated with intrinsic (plant organ) and extrinsic (geographical origin) factors might be a characteristic metabolic feature of the genus Curcuma.

3.3. Metabolite differences among EO samples using chemometric analysis

To further investigate the metabolic differences among EO samples from C. kwangsiensis across different plant parts and collection sites, chemometric analyses, including PCA and PLS-DA, were performed. Sample groups comprised either different plant parts from the same collection site or the same plant part from different sites (Fig. 2, Fig. 3). The PCA score plots based on PC1 and PC2 accounted for 52.8–83.8% of the total variance, indicating that the PCA models effectively captured the major variations among the samples (Figs. 2A-C, 3A-E). In PLS-DA, R2X (cum) and R2Y (cum) represent the cumulative explained variance of the predictor (X) and response (Y) variables, respectively, while Q2 indicates the predictive ability of the model. The cumulative R2X, R2Y, and Q2 values of the constructed PLS-DA models were all close to 1 (all >0.9), demonstrating the high reliability of the models (Tables S2-S3). Negative Q2 y-intercepts from permutation tests (Figs. 2A2-C2, 3A2-E2) further confirmed that the models were robust and free from overfitting (Westerhuis et al., 2008). Both PCA and PLS-DA score plots revealed distinct separation of EO samples, indicating clear metabolite differences among plant parts (Fig. 2) and among geographical origins (Fig. 3), substantiating the significant influence of plant parts and collection sites on the chemical composition of C. kwangsiensis EOs. These results also demonstrate the effectiveness of untargeted GC–MS metabolomics combined with chemometric analysis for discriminating EO samples. Such findings are consistent with previous studies reporting pronounced chemical variability in EOs from C. wenyujin rhizomes and roots (Zhu et al., 2013) and in C. aeruginosa rhizomes from different geographical origins as revealed by PCA analysis (Nugraheni et al., 2024).

Fig. 2.

Fig. 2

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Chemical differences among EO samples from different plant parts of C. kwangsiensis. (A-C) PCA score plots; (A1-C1) PLS-DA score plots; (A2-C2) permutation tests for the corresponding PLS-DA models. L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County.

Fig. 3.

Fig. 3

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Chemical differences among EO samples from different geographical locations of C. kwangsiensis. (A-E) PCA score plots; (A1-E1) PLS-DA score plots; (A2-E2) permutation tests for the corresponding PLS-DA models. L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County.

To identify key metabolites driving these chemical variations, differential constituents were characterized using VIP values and ANOVA p-values less than 0.05. Although a VIP threshold >1 is commonly used in metabolomics studies as a general cutoff for identifying important variables in PLS-DA models (Djande et al., 2021), more stringent thresholds such as VIP >1.3 (Ahmad et al., 2021) or VIP >1.5 (Ren et al., 2025; Wang et al., 2026) have been applied in previous studies to reduce false positives and highlight the most discriminative metabolites. In this study, a stricter threshold of VIP >1.5 was adopted to enhance the robustness and interpretability of the selected markers. Using these criteria, a total of 31 differential EO constituents were identified, with VIP values ranging from 1.50 to 7.72, including seven oxygenated sesquiterpenes, seven sesquiterpene hydrocarbons, eight oxygenated monoterpenes, seven monoterpene hydrocarbons, one furan, and one other compound (Table 1). Several compounds, including curzerene, trans-β-elemenone, and germacrone, were consistently identified as differential metabolites across multiple comparison groups, highlighting their high sensitivity to variations in plant part and geographical origin. In contrast, other compounds, such as β-myrcene and limonene, were identified as differential only in specific comparison groups, suggesting more localized or condition-dependent variations. Collectively, these discriminative metabolites account for the pronounced chemical differences observed among different plant parts within the exact geographical origin, as well as among the same plant part collected from different locations. Furthermore, these compounds have strong potential as chemical markers for authentication and quality control of EO products derived from specific plant parts or origins (Rebiai et al., 2022). Many of these discriminative constituents, including germacrone, trans-β-elemene, and (E)-β-caryophyllene, have been reported to exhibit bioactivities relevant to food and health, such as antioxidant, anti-inflammatory, and analgesic effects (Chen et al., 2023; Fidyt et al., 2016; Riaz et al., 2020). Accordingly, variations in the relative abundances of these constituents may contribute to differences in the functional properties of EO samples derived from different plant organs and geographic origins of C. kwangsiensis.

Table 1.

Differential EO constituents discriminating plant parts and geographical origins.

No.a Compounds Classes VIP scoresb
FM LS WM L F TR FR RH
1 Curzerene Oxygenated sesquiterpenes 3.78 2.61 3.10 2.64 2.32 4.44 1.89 2.74
9 trans-β-Elemenone Oxygenated sesquiterpenes 5.37 5.35 4.61 5.22 5.20 7.41 5.71 7.72
11 Spathulenol Oxygenated sesquiterpenes 1.64 1.83 1.59
14 β-Eudesmol Oxygenated sesquiterpenes 1.50 2.08 1.51 1.53
17 Germacrone Oxygenated sesquiterpenes 2.50 2.33 2.02 1.70 3.07 3.36 2.36 3.63
20 Nootkatone Oxygenated sesquiterpenes 4.14 3.21 3.33 1.93 2.81 4.64 2.25 5.39
21 Farnesyl acetone Oxygenated sesquiterpenes 1.69
25 trans-β-Elemene Sesquiterpene hydrocarbons 2.22 2.38 3.39 2.23 2.64 3.70 2.92 3.08
27 Caryophyllene Sesquiterpene hydrocarbons 3.50 3.23 3.39 2.79 4.73 4.87
32 (E)-β-Farnesene Sesquiterpene hydrocarbons 1.69 1.66
34 α-Humulene Sesquiterpene hydrocarbons 4.86 3.26 4.04 4.43 4.75 2.75 6.20
36 Germacrene D Sesquiterpene hydrocarbons 2.14 2.08 2.33 2.36 2.49 1.63 2.38 1.82
37 α-Farnesene Sesquiterpene hydrocarbons 1.83 1.85
38 α-Selinene Sesquiterpene hydrocarbons 1.51
42 Eucalyptol Oxygenated monoterpenes 1.51 3.74 2.96 1.82 1.84
43 Linalool Oxygenated monoterpenes 3.18 3.55 3.37 1.93 1.64
47 Camphor Oxygenated monoterpenes 1.73 1.98 1.97 4.77 3.04 3.68 3.76 4.30
50 Isoborneol Oxygenated monoterpenes 2.10 3.19 3.40 2.46 2.29 3.55 3.24 4.04
52 Borneol Oxygenated monoterpenes 2.19 1.96 1.75 2.12 2.15
55 α-Terpineol Oxygenated monoterpenes 1.71 2.50
60 Bornyl acetate Oxygenated monoterpenes 1.82 1.83 2.66 2.88
62 Myrtenyl acetate Oxygenated monoterpenes 1.71 2.50 1.68
63 α-Pinene Monoterpene hydrocarbons 2.07 3.15 4.02 3.33 1.83 1.61
64 Camphene Monoterpene hydrocarbons 2.01 1.77 1.52 2.72 1.90
65 α-Sabinene Monoterpene hydrocarbons 1.65 1.87
66 Nopinene Monoterpene hydrocarbons 2.12 2.25 1.91 5.21 2.84 2.76
67 β-Myrcene Monoterpene hydrocarbons 2.01
72 Limonene Monoterpene hydrocarbons 2.78
73 β-Phellandrene Monoterpene hydrocarbons 2.47 3.23
77 Tetrahydro-2,5-dimethylfuran Furans 3.20 2.36 2.66 1.91
88 2,4-Dimethylbenzophenone Others 2.19
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a

The order of compounds (No.) corresponds to that in Table S1.

b

VIP scores of differential compounds for discriminating metabolite differences among plant parts or across locations (L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County).

3.4. Cytotoxic activity

EOs from numerous Curcuma taxa, including C. yunnanensis and C. zedoaria, have been well documented for their cytotoxic properties (Xiang et al., 2018). In the present study, the cytotoxic effects of EO samples derived from C. kwangsiensis were evaluated against three human cancer cell lines: K562, A549, and HepG2. At a test concentration of 100 μg/mL, all EO samples exhibited significant inhibitory effects on the tested cancer cells, with inhibition rates ranging from 54.55% to 99.75% (Fig. S1). The positive control DOX at 5 μg/mL showed strong cytotoxic activity, yielding inhibition rates of 80.15–90.33% across the three cancer cell lines (Fig. S1), thereby confirming the reliability of the assay. Further analyses revealed IC50 values ranging from 10.82 to 89.74 μg/mL (Fig. 4). Among the tested samples, LS-RH and LS-TR EOs exhibited the strongest cytotoxicity against K562 and HepG2 cells, with IC50 values of 10.82 and 18.53 μg/mL, respectively. These findings further support the potential application of rhizomes and tuberous roots of C. kwangsiensis as functional or nutraceutical ingredients. In contrast, FM-FR and FM-F EOs showed the highest activity against A549 cells, with IC50 values ranging from 32.55 to 33.73 μg/mL (Fig. 4). IC50 values for EOs from other Curcuma species, including C. zedoaroides, C. wenyujin, and C. purpurascens, are generally above 43 μg/mL against K562, A549, and HepG2 cells (Hong et al. Hong et al., 2014; Thu et al., 2024; Xiao et al., 2008), suggesting that the most active EOs in this study exhibit comparatively strong cytotoxic activity. A previous study reported that the rhizome EO of C. kwangsiensis var. nanlingensis exhibited cytotoxic activity against HepG2 cells with an IC50 value of 156.43 μg/mL (Xiang et al., 2018), which is markedly higher than the values observed in this study (18.53–82.75 μg/mL) and thus indicates weaker cytotoxicity. This discrepancy may be attributed to variations in chemical composition among different plant varieties or origins of C. kwangsiensis. To our knowledge, the cytotoxic activities of EOs derived from non-rhizome plant parts of C. kwangsiensis are reported here for the first time. Overall, these findings highlight the importance of plant part selection and geographic origin in shaping the cytotoxic potential of C. kwangsiensis EOs and provide a basis for their differentiated utilization. However, the present study focused on cancer cell lines and the effects on human normal cells have not yet been evaluated. Future studies are warranted to investigate their selectivity, safety, and underlying mechanisms of action, particularly for rhizome and tuberous root EOs that exhibited prominent cytotoxic activity.

Fig. 4.

Fig. 4

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Cytotoxic effects of different EO samples from C. kwangsiensis on three cancer cells. L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County. Different lowercase letters above the columns indicate statistically significant differences (p < 0.05) among samples within the same cancer cell line, as determined by Duncan's multiple range test.

The chemical composition of the EOs provides insight into the cytotoxic activities observed in C. kwangsiensis. Sesquiterpenes, monoterpenes, and their derivatives are widely recognized for their anticancer potential (Abu-Izneid et al., 2020; Silva et al., 2021). Several major constituents identified in the present study, including trans-β-elemene, germacrone, and α-pinene, have been reported to exhibit strong cytotoxic effects against various cancer cell lines, including A549, K562, and HepG2 (Chen et al., 2015; Jiang et al., 2017; Lou et al., 2023). For instance, trans-β-elemene at 40 μg/mL showed a notable inhibitory effect on HepG2 cells, with an inhibition rate of 73.7% after 96 h of treatment (Dai et al., 2013). The relatively higher abundance of trans-β-elemene (15.22%) in WM-TR EO (Table S1) may therefore contribute to its enhanced cytotoxic activity against HepG2 cells compared with other WM samples. In addition, trans-β-elemenone, another sesquiterpenoid detected in the EOs, has been reported to undergo a Cope rearrangement to form germacrone (Setzer et al., 2008). Given that germacrone has been widely reported to possess notable anticancer activities (Lou et al., 2023), the relatively high abundance of trans-β-elemenone may also indirectly contribute to the cytotoxic activity of C. kwangsiensis EOs through this structural transformation. Taken together, the presence and relative abundance of these bioactive constituents likely contribute, at least in part, to the cytotoxic effects observed across the EO samples in this study.

3.5. Anti-inflammatory effect

Although the anti-inflammatory effects of EOs from specific plant parts, such as leaves and rhizomes of Curcuma species, have been reported (Borah et al., 2019; Xiang et al., 2018), comparative studies involving multiple plant organs remain limited. In this study, the anti-inflammatory activities of C. kwangsiensis EOs from different plant parts and geographic origins were evaluated using an LPS-stimulated RAW 264.7 macrophage model by measuring their inhibitory effects on NO production. At 100 μg/mL, all EO samples strongly inhibited NO production, with rates exceeding 93%; however, this concentration also caused significant cytotoxicity, reducing RAW 264.7 cell viability below 60%. Therefore, a non-cytotoxic concentration of 25 μg/mL, which maintained cell viability above 95%, was used for subsequent anti-inflammatory analysis. At this concentration, NO inhibition rates ranged from 10.35% to 67.69% (Fig. 5). Significant differences in NO inhibition were observed among EO samples, indicating that both plant part and geographical origin significantly influenced their anti-inflammatory activities. Notably, TR EOs from all three collection sites consistently exhibited the strongest NO inhibitory effects, with inhibition rates of 46.19–67.69% (Fig. 5), suggesting that tuberous roots may represent a particularly promising source of anti-inflammatory EOs. The most active EO was WM-TR EO, exhibiting an NO inhibitory rate of 67.69%, followed by LS-TR EO (57.71%) and WM-RH EO (56.55%). These activities are comparable to those of rhizome EOs from other Curcuma taxa, such as C. angustifolia, which inhibited NO production by 66.42% in LPS-stimulated RAW 264.7 macrophages at 25 μg/mL (Gadnayak et al., 2024). IC50 values were further determined for EO samples showing NO inhibition above 50%, namely LS-TR, WM-TR, and WM-RH, with values of 21.47, 18.31, and 22.21 μg/mL, respectively. These IC50 values were comparable to those reported for other Curcuma species, such as C. longa (IC50 = 19.16 μg/mL) (Mossmann et al., 2024). Previous studies have reported that rhizome EOs of C. kwangsiensis exhibit strong anti-inflammatory activity in a TPA-induced mouse ear edema model through the regulation of TNF-α and COX-2 expression (Zhang, Yang, Chen, et al., 2017). However, the anti-inflammatory mechanisms of EOs derived from other plant parts remain unexplored and warrant further investigation.

Fig. 5.

Fig. 5

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Anti-inflammatory effects of C. kwangsiensis EO samples at 25 μg/mL (A) and their IC50 values (B). L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County. Different lowercase letters above the columns indicate statistically significant differences among samples (p < 0.05) based on Duncan's multiple range test.

Several major constituents identified in C. kwangsiensis EOs, including trans-β-elemene (Wang et al., 2023), α-humulene, (E)-β-caryophyllene (Viveiros et al., 2022), and nootkatone (Xu et al., 2021), have been reported to possess anti-inflammatory properties and may collectively contribute to the observed activities. Although LS-TR, WM-TR, and WM-RH EOs showed stronger anti-inflammatory effects, some known bioactive compounds, such as α-humulene and (E)-β-caryophyllene, were present at lower relative contents than in other samples, such as FM-FR and LS-F EOs (Table S1). These findings suggest that the anti-inflammatory activity of C. kwangsiensis EOs was likely mediated by the combined or interactive effects of multiple constituents rather than any single compound. Trans-β-Elemenone, identified as the most abundant constituent, was especially prevalent in the highly active LS-TR and WM-TR samples (Table S1); however, its anti-inflammatory potential remains unclear and warrants further study.

3.6. Antioxidant capacity

The rhizome EOs from several Curcuma species, such as C. zedoaria and C. zanthorrhiza, are known for their antioxidant potential (Dosoky & Setzer, 2018). The present study showed that all EO samples obtained from C. kwangsiensis demonstrated scavenging activity against both DPPH and ABTS radicals (Fig. 6). In the DPPH assay, IC50 values varied from 0.54 to 5.26 mg/mL. The antioxidant activity of C. kwangsiensis EOs observed in the current study can be considered moderate to weak, as their IC50 values are comparable to those of C. longa (IC50 = 3.5–4.4 mg/mL) (Gounder & Lingamallu, 2012), but markedly weaker than those reported for other Curcuma species, such as C. aeruginosa (IC50 = 24.32 μg/mL) (Theanphong et al., 2015) and C. aromatica (IC50 = 14.45 μg/mL) (Al-Reza et al., 2010). Significant differences were detected among EOs from different plant organs and collection locations. Notably, the TR EOs (IC50 = 0.54–0.97 mg/mL) and RH EOs (IC50 = 1.71–2.42 mg/mL) collected from all three locations displayed the strongest DPPH radical scavenging activities, with the FM-TR EO exhibiting the highest activity among the collection sites. Compared with the DPPH assay, most EO samples showed greater scavenging efficiency in the ABTS assay, with IC50 values ranging from 0.24 to 1.15 mg/mL. A similar trend has been reported for rhizome EO of C. longa, which exhibited stronger activity in the ABTS assay than in the DPPH assay (Avanço et al., 2017). Consistent with the DPPH results, TR and RH EOs also demonstrated higher ABTS radical scavenging activity, with IC50 values of 0.25–0.55 mg/mL, among which the TR EO collected from FM again showed the strongest effect. Antioxidant activities at the mg/mL level have been frequently reported for EOs in previous studies, including those by Mau et al. (2003) and Gounder and Lingamallu (2012). In contrast, Zhang, Yang, Chen, et al. (2017) reported strong DPPH radical scavenging activity for rhizome EOs from four Curcuma species (IC50 = 3.47–30.19 μg/mL), with C. kwangsiensis showing the highest activity, exceeding that of ascorbic acid. These IC50 values were lower than those obtained in the present study, which may be attributed to variations in experimental conditions, including plant chemotypes, geographical origins, and harvest seasons. To our knowledge, this is the first study to report the antioxidant activities of EOs from C. kwangsiensis plant parts other than rhizomes. These results indicate that EOs derived from the tuberous roots and rhizomes of C. kwangsiensis, particularly those collected from the Fumian, represent promising natural sources for antioxidant applications. However, the effects of environmental factors such as temperature, humidity, and soil on chemical composition and bioactivities remain to be further investigated to better understand the mechanisms underlying bioactivity variation.

Fig. 6.

Fig. 6

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Antioxidant effects of C. kwangsiensis EO samples in DPPH (A) and ABTS (B) assays. L: leaves; F: flowers; TR: tuberous roots; FR: fibrous roots; RH: rhizomes; FM: Fumian District; LS: Lingshan County; WM: Wuming County. Different lowercase letters above the columns indicate statistically significant differences among samples (p < 0.05) according to Duncan's multiple range test.

Several identified sesquiterpenes, such as trans-β-elemene (Chen et al., 2023), (E)-β-caryophyllene (Gushiken et al., 2022), and nootkatone (Zhang et al., 2022), are known to possess antioxidant properties. In addition, monoterpenes such as limonene (Anandakumar et al., 2021) and linalool (An et al., 2021) have also been reported to exhibit antioxidant activity. The relatively higher contents of trans-β-elemene and nootkatone in WM-TR and WM-RH EOs (Table S1) may partly explain their stronger antioxidant performance compared with other WM samples. In the FM samples, although FM-TR and FM-RH exhibited the highest antioxidant activities, their (E)-β-caryophyllene contents were lower than those of other FM EOs (Table S1), suggesting that additional constituents may contribute to the observed effects. In addition to the characterized constituents, the remaining unidentified compounds in the EO samples may also contribute to the observed activities, either as active components or through synergistic, antagonistic, or additive interactions (Caesar & Cech, 2019). This may introduce uncertainty when attributing bioactivity solely to the identified compounds. Therefore, further isolation and structural characterization are needed to better elucidate their contributions.

3.7. Correlations between chemical composition and bioactivities

To further elucidate the relationships between EO composition and bioactivities, Spearman's rank correlation analysis was performed, taking into account variations arising from both plant parts and geographic origins. The 30 most abundant constituents, along with differential compounds, were selected to construct the correlation heatmap (Fig. 7).

Fig. 7.

Fig. 7

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Spearman's rank correlation analysis between selected volatile compounds and their related bioactivities, including cytotoxic effects (inhibition of A549, K562, and HepG2 cells), anti-inflammatory activity (NO inhibition), and antioxidant capacity (DPPH and ABTS radical scavenging). Black dashed lines separate different categories of metabolites or bioassays. Correlation coefficients are shown on the right, with positive values (above zero) indicating positive correlations and negative values (below zero) indicating negative correlations between volatiles and biological activities.

As shown in Fig. 7, both positive and negative correlations were observed between C. kwangsiensis EOs and their cytotoxic, anti-inflammatory, and antioxidant activities. Sesquiterpenes and their derivatives are widely recognized for their anticancer potential (Abu-Izneid et al., 2020). In this study, several oxygenated sesquiterpenes and sesquiterpene hydrocarbons, including trans-β-elemenone, (E)-β-caryophyllene, and α-humulene, were positively correlated with growth inhibition against K562, A549, or HepG2 cells. In contrast, compounds such as camphene and α-sabinene exhibited negative correlations (Fig. 7). (E)-β-Caryophyllene and α-humulene have previously been reported to exhibit cytotoxic activity against A549 cells, with IC50 values of 31.3 and 16.7 μg/mL, respectively (Su et al., 2015), supporting the positive correlations observed in this study. Although trans-β-elemenone showed a positive association with growth inhibition in K562 and HepG2 cells, its cytotoxic activity has not been characterized, and further investigation is therefore required. For anti-inflammatory activity, volatile constituents such as nootkatone, trans-β-elemene, and globulol showed positive correlations. Several compounds previously reported to possess anti-inflammatory properties, such as (E)-β-caryophyllene (Guo et al., 2025) and linalool (Varia et al., 2020), exhibited negative correlations in this study, possibly due to complex interactions among EO constituents, such as synergistic or antagonistic effects. Similarly, both positive and negative correlations were observed for antioxidant activity as evaluated by DPPH and ABTS assays. Nootkatone, germacrone, germacrene D, δ-elemene, and 2,4-dimethylbenzophenone showed strong positive correlations, partly explaining the stronger antioxidant activities observed in C. kwangsiensis EOs, particularly in TR and RH samples. Among them, nootkatone has been well documented for its significant antioxidant activity (Zhang et al., 2022), whereas the antioxidant potential of germacrone, germacrene D, δ-elemene, and 2,4-dimethylbenzophenone remains insufficiently explored and needs to be further investigation. Altogether, these findings provide preliminary insights into the relationships between chemical composition and bioactivities of C. kwangsiensis EOs, offering useful clues for future mechanistic studies and the discovery of potential bioactive compounds.

4. Conclusion

In this study, the chemical diversity and biological activities of EOs from five plant parts of C. kwangsiensis collected from three geographic origins were systematically investigated for the first time using a GC–MS-based metabolomics approach. Pronounced variations in EO yields were observed among plant parts, with rhizomes consistently producing the highest yields (1.88–2.22%), confirming the rhizomes as the primary EO-accumulating organ of C. kwangsiensis. A total of 88 compounds were identified across all EO samples, predominantly comprising oxygenated sesquiterpenes, sesquiterpene hydrocarbons, oxygenated monoterpenes, and monoterpene hydrocarbons, with their relative abundances varying markedly among plant parts and geographic origins. Chemometric analyses, including PCA and PLS-DA, further demonstrated that both plant part and geographic origin significantly influenced EO chemical profiles. A total of 31 compounds, mainly oxygenated monoterpenes and sesquiterpenes as well as monoterpene and sesquiterpene hydrocarbons, were identified as key differential metabolites contributing to the observed chemical variability. Biological activity evaluations revealed that C. kwangsiensis EOs possess notable cytotoxic, anti-inflammatory, and antioxidant activities, although their potencies varied substantially depending on plant part and origin. Several EOs, including LS-RH, LS-TR, and FM-FR, exhibited stronger cytotoxic effects, with IC50 values below 34 μg/mL. In addition, LS-TR, WM-TR, and WM-RH showed pronounced inhibitory effects on NO production in LPS-stimulated RAW 264.7 macrophages, with IC50 values below 23 μg/mL, indicating promising anti-inflammatory potential. Antioxidant assays further demonstrated that EOs from tuberous roots and rhizomes displayed stronger radical-scavenging activity in both the DPPH and ABTS assays than those from other plant parts, with FM-TR and FM-RH showing the highest activity (IC50 = 0.24–1.71 mg/mL). Spearman's rank correlation analysis demonstrated that major and differential volatiles were closely associated with the observed bioactivities, with several EOs, including trans-β-elemenone, nootkatone, and germacrone, showing positive contributions. Collectively, these findings demonstrate that plant part selection and geographic origin are critical factors determining the quality and functional potential of C. kwangsiensis EOs. However, several limitations of the present study warrant further investigation, including the influence of harvest season on EO composition and bioactivity, the characterization of the unannotated fractions of EO samples, the cytotoxicity of EOs toward normal cells, and the underlying mechanisms of EO activity.

CRediT authorship contribution statement

Fengke Lin: Writing – original draft, Validation, Supervision, Project administration, Methodology, Funding acquisition, Formal analysis, Data curation, Conceptualization. Rui Zhao: Data curation. Zihao Liu: Data curation. Qun Zhou: Data curation. Qianyun Wang: Investigation, Data curation. Edward J. Kennelly: Writing – review & editing, Validation. Jun Feng: Writing – review & editing, Validation, Supervision, Project administration, Investigation. Renchuan Hu: Writing – review & editing, Validation, Supervision, Project administration, Investigation, Funding acquisition.

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 work was funded by the Guangxi Key Laboratory of Traditional Chinese Medicine Quality Standards, Guangxi Institute of Chinese Medicine & Pharmaceutical Science (GZZK202406), Tianjin Natural Science Foundation Project (25JCQNJC00730), Survey and Collection of Germplasm Resources of Woody & Herbaceous Plants in Guangxi, China (GXFS-2021-34), and the Natural Science Foundation of Guangxi (2023GXNSFAA026089).

Footnotes

Appendix A

Supplementary data to this article can be found online at https://doi.org/10.1016/j.fochx.2026.103879.

Contributor Information

Fengke Lin, Email: fengkelin2023@email.tjut.edu.cn.

Jun Feng, Email: fengjun@gicmp.ac.cn.

Renchuan Hu, Email: hrcgxmi@163.com.

Appendix A. Supplementary data

Supplementary material. Table S1. Chemical composition of EOs from different plant parts of C. kwangsiensis across different geographical locations. Table S2. Quality parameters for PLS-DA plots in Figure 2. Table S3. Quality parameters for PLS-DA plots in Figure 3. Fig. S1. Cytotoxic effects of EO samples from C. kwangsiensis on three cancer cell lines at 100 μg/mL.

mmc1.docx (180.4KB, docx)

Data availability

Data will be made available on request.

References

  1. Abu-Izneid T., Rauf A., Shariati M.A., Khalil A.A., Imran M., Rebezov M., Uddin M.S., Mahomoodally M.F., Rengasamy K.R. Sesquiterpenes and their derivatives-natural anticancer compounds: An update. Pharmacological Research. 2020;161 doi: 10.1016/j.phrs.2020.105165. [DOI] [PubMed] [Google Scholar]
  2. Adams R.P. 4.1 ed. Allured Publishing Corporation; Illinois: 2017. Identification of essential oil components by gas chromatography/mass spectrometry. [Google Scholar]
  3. Ahmad A.A., Ismail I.S., Shaikh M.F., Abas F., Shaari K. Multi-platform metabolomics analyses revealed the complexity of serum metabolites in LPS-induced neuroinflammed rats treated with clinacanthus nutans aqueous extract. Frontiers in Pharmacology. 2021;12 doi: 10.3389/fphar.2021.629561. [DOI] [PMC free article] [PubMed] [Google Scholar]
  4. Al-Reza S.M., Rahman A., Sattar M.A., Rahman M.O., Fida H.M. Essential oil composition and antioxidant activities of curcuma aromatica salisb. Food and Chemical Toxicology. 2010;48:1757–1760. doi: 10.1016/j.fct.2010.04.008. [DOI] [PubMed] [Google Scholar]
  5. An Q., Ren J., Li X., Fan G., Qu S., Song Y., Li Y., Pan S. Recent updates on bioactive properties of linalool. Food & Function. 2021;12:10370–10389. doi: 10.1039/D1FO02120F. [DOI] [PubMed] [Google Scholar]
  6. Anandakumar P., Kamaraj S., Vanitha M.K. D-limonene: A multifunctional compound with potent therapeutic effects. Journal of Food Biochemistry. 2021;45 doi: 10.1111/jfbc.13566. [DOI] [PubMed] [Google Scholar]
  7. Angel G., Menon N., Vimala B., Nambisan B. Essential oil composition of eight starchy curcuma species. Industrial Crops and Products. 2014;60:233–238. doi: 10.1016/j.indcrop.2014.06.028. [DOI] [Google Scholar]
  8. Avanço G.B., Ferreira F.D., Bomfim N.S., Peralta R.M., Brugnari T., Mallmann C.A.…Machinski M., Jr. Curcuma longa l. essential oil composition, antioxidant effect, and effect on fusarium verticillioides and fumonisin production. Food Control. 2017;73:806–813. doi: 10.1016/j.foodcont.2016.09.032. [DOI] [Google Scholar]
  9. Bansal R., Bahl J., Garg S., Naqvi A., Kumar S. Differential chemical compositions of the essential oils of the shoot organs, rhizomes and rhizoids in the tumeric curcuma longa grown in indo-Gangetic plains. Pharmaceutical Biology. 2002;40:384–389. doi: 10.1076/phbi.40.5.384.8458. [DOI] [Google Scholar]
  10. Barra A. Factors affecting chemical variability of essential oils: A review of recent developments. Natural Product Communications. 2009;4:1147–1154. doi: 10.1177/1934578X0900400827. [DOI] [PubMed] [Google Scholar]
  11. Borah A., Paw M., Gogoi R., Loying R., Sarma N., Munda S., Pandey S.K., Lal M. Chemical composition, antioxidant, anti-inflammatory, anti-microbial and in-vitro cytotoxic efficacy of essential oil of curcuma caesia roxb. Leaves: An endangered medicinal plant of north East India. Industrial Crops and Products. 2019;129:448–454. doi: 10.1016/j.indcrop.2018.12.035. [DOI] [Google Scholar]
  12. Caesar L.K., Cech N.B. Synergy and antagonism in natural product extracts: When 1+ 1 does not equal 2. Natural Product Reports. 2019;36:869–888. doi: 10.1039/C9NP00011A. [DOI] [PMC free article] [PubMed] [Google Scholar]
  13. Chen W., Liu Y., Li M., Mao J., Zhang L., Huang R., Jin X., Ye L. Anti-tumor effect of α-pinene on human hepatoma cell lines through inducing G2/M cell cycle arrest. Journal of Pharmacological Sciences. 2015;127:332–338. doi: 10.1016/j.jphs.2015.01.008. [DOI] [PubMed] [Google Scholar]
  14. Chen X., Huang C., Li K., Liu J., Zheng Y., Feng Y., Kai G. Recent advances in biosynthesis and pharmacology of β-elemene. Phytochemistry Reviews. 2023;22:169–186. doi: 10.1007/s11101-022-09833-0. [DOI] [Google Scholar]
  15. Choochote W., Chaiyasit D., Kanjanapothi D., Rattanachanpichai E., Jitpakdi A., Tuetun B., Pitasawat B. Chemical composition and anti-mosquito potential of rhizome extract and volatile oil derived from curcuma aromatica against aedes aegypti (diptera: Culicidae) Journal of Vector Ecology. 2005;30:302. [PubMed] [Google Scholar]
  16. Commission of Chinese Herbal Medicine . In: Zhong Hua Ben Cao (Chinese herbal medicine) Song L., editor. Shanghai Scientific and Technical Publishers; Shanghai: 1999. Ting Li Zi; pp. 628–629. [Google Scholar]
  17. Dai Z., Tang W., Lu W., Gao J., Kang H., Ma X., Min W., Wang X., Wu W. Antiproliferative and apoptotic effects of β-elemene on human hepatoma HepG2 cells. Cancer Cell International. 2013;13:27. doi: 10.1186/1475-2867-13-27. [DOI] [PMC free article] [PubMed] [Google Scholar]
  18. Djande H.C.Y., Piater L.A., Steenkamp P.A., Tugizimana F., Dubery I.A. A metabolomics approach and chemometric tools for differentiation of barley cultivars and biomarker discovery. Metabolites. 2021;11:578. doi: 10.3390/metabo11090578. [DOI] [PMC free article] [PubMed] [Google Scholar]
  19. Dosoky N.S., Setzer W.N. Chemical composition and biological activities of essential oils of curcuma species. Nutrients. 2018;10:1196. doi: 10.3390/nu10091196. [DOI] [PMC free article] [PubMed] [Google Scholar]
  20. Fidyt K., Fiedorowicz A., Strządała L., Szumny A. Β-Caryophyllene and β-caryophyllene oxide—Natural compounds of anticancer and analgesic properties. Cancer Medicine. 2016;5:3007–3017. doi: 10.1002/cam4.816. [DOI] [PMC free article] [PubMed] [Google Scholar]
  21. Gadnayak A., Nayak A., Jena S., Sahoo A., Panda P.C., Ray A., Nayak S. Nitric oxide inhibitory potential of curcuma angustifolia roxb. Essential oil: An in silico and in vitro analysis. Plant Science Today. 2024;11:234–243. doi: 10.14719/pst.3410. [DOI] [Google Scholar]
  22. Gounder D.K., Lingamallu J. Comparison of chemical composition and antioxidant potential of volatile oil from fresh, dried and cured turmeric (curcuma longa) rhizomes. Industrial Crops and Products. 2012;38:124–131. doi: 10.1016/j.indcrop.2012.01.014. [DOI] [Google Scholar]
  23. Guo W., Zhang C., Yang X., Lin Z., Huang H., Shu Y., Shentu J., Xu Y. A mechanistic review on the anti-inflammatory effects of β-caryophyllene. Mini-Reviews in Organic Chemistry. 2025;22:671–684. doi: 10.2174/0118756298342319241125043305. [DOI] [Google Scholar]
  24. Gururani S., Gairola K., Kumar R., Prakash O., Dubey S.K. Altitudinal and geographical variations in phytochemical composition and biological activities of curcuma longa accession from Uttarakhand, the himalayan region. Journal of Food Processing and Preservation. 2022;46 doi: 10.1111/jfpp.16384. [DOI] [Google Scholar]
  25. Gushiken L.F.S., Beserra F.P., Hussni M.F., Gonzaga M.T., Ribeiro V.P., De Souza P.F., Campos J.C.L., Massaro T.N.C., Hussni C.A., Takahira R.K. Beta-caryophyllene as an antioxidant, anti-inflammatory and re-epithelialization activities in a rat skin wound excision model. Oxidative Medicine and Cellular Longevity. 2022;2022 doi: 10.1155/2022/9004014. [DOI] [PMC free article] [PubMed] [Google Scholar]
  26. Hong S.L., Lee G.S., Syed Abdul Rahman S.N., Ahmed Hamdi O.A., Awang K., Aznam Nugroho N., Abd Malek S.N. Essential oil content of the rhizome of Curcuma purpurascens Bl. (Temu Tis) and its antiproliferative effect on selected human carcinoma cell lines. The Scientific World Journal. 2014;2014 doi: 10.1155/2014/397430. [DOI] [PMC free article] [PubMed] [Google Scholar]
  27. Hu Y., Zhang J., Kong W., Zhao G., Yang M. Mechanisms of antifungal and anti-aflatoxigenic properties of essential oil derived from turmeric (curcuma longa l.) on aspergillus flavus. Food Chemistry. 2017;220:1–8. doi: 10.1016/j.foodchem.2016.09.179. [DOI] [PubMed] [Google Scholar]
  28. Huang P., Wang Q., Zhou G., Guan S., Li J., Wang N., Xie X. GC-MS differences of volatile components in rhizome, roots and fibrous root of curcuma kwangsiensis. Lishizhen Medicine and Mateia Medica Research. 2020;31:2366–2368. [Google Scholar]
  29. Huang Z., Yuan Y., Tan Z., Zheng J., Zhang W., Huang S., Wang Y., Chen M., Zhang L., Li H. Metabolomics in combination with network pharmacology reveals the potential anti-neuroinflammatory mechanism of essential oils from four curcuma species. Industrial Crops and Products. 2023;195 doi: 10.1016/j.indcrop.2023.116411. [DOI] [Google Scholar]
  30. Jena S., Ray A., Banerjee A., Sahoo A., Nasim N., Sahoo S., Kar B., Patnaik J., Panda P.C., Nayak S. Chemical composition and antioxidant activity of essential oil from leaves and rhizomes of curcuma angustifolia roxb. Natural Product Research. 2017;31:2188–2191. doi: 10.1080/14786419.2017.1278600. [DOI] [PubMed] [Google Scholar]
  31. Jiang D., Mo J., Jiang L., Zang Q., Xu L., Chen X. The enzymatic extraction process of volatile oil from rhizoma curcumae and its antioxidant activity study. The Food Industry. 2023;44:1–4. [Google Scholar]
  32. Jiang Z., Jacob J.A., Loganathachetti D.S., Nainangu P., Chen B. Β-Elemene: Mechanistic studies on cancer cell interaction and its chemosensitization effect. Frontiers in Pharmacology. 2017;8:105. doi: 10.3389/fphar.2017.00105. [DOI] [PMC free article] [PubMed] [Google Scholar]
  33. Kumar K.N., Venkataramana M., Allen J.A., Chandranayaka S., Murali H.S., Batra H.V. Role of curcuma longa l. essential oil in controlling the growth and zearalenone production of fusarium graminearum. LWT- Food Science and Technology. 2016;69:522–528. doi: 10.1016/j.lwt.2016.02.00. [DOI] [Google Scholar]
  34. Li X., Zhang C., Li S., Wang H., Yu P., Shao H., Wang S., Wang H., Jin F. Unveiling key aroma-active compounds of two distinct aroma-types oriental melon: An integrated sensomics approach utilizing GC×GC-O-TOF-MS, GC-MS, aroma recombination and omission experiments. Food Chemistry: X. 2025;28 doi: 10.1016/j.fochx.2025.102517. [DOI] [PMC free article] [PubMed] [Google Scholar]
  35. Lin F., Kennelly E.J., Linington R.G., Long C. Comprehensive metabolite profiling of two edible garcinia species based on UPLC-ESI-QTOF-MSE coupled with bioactivity assays. Journal of Agricultural and Food Chemistry. 2023;71:7604–7617. doi: 10.1021/acs.jafc.2c08372. [DOI] [PubMed] [Google Scholar]
  36. Lin F., Long C. GC-TOF-MS-based metabolomics correlated with bioactivity assays unveiled seasonal variations in leaf essential oils of two species in garcinia l. Industrial Crops and Products. 2023;194 doi: 10.1016/j.indcrop.2023.116356. [DOI] [Google Scholar]
  37. Lin F., Zhou Q., Lin G., Liu Z., Zhao R., Luo B. Phytochemical profiling of lepidium apetalum using GC/LC-MS metabolomics coupled with chemometric and bioactivity analyses. Food Chemistry: X. 2025;32 doi: 10.1016/j.fochx.2025.103376. [DOI] [PMC free article] [PubMed] [Google Scholar]
  38. Liu B., Yuan J., Su h., Chai L., Chen X. Progress in the composition, efficacy, analysis, and clinical application of curcuma kwangsiensis. Guangxi Sciences. 2023;30:1037–1050. [Google Scholar]
  39. Lou G., Huang Y., Wang Y., Chen S., Liu C., Li Y., Feng J. Germacrone, a novel and safe anticancer agent from genus curcuma: A review of its mechanism. Anti-Cancer Agents in Medicinal Chemistry. 2023;23:1490–1498. doi: 10.2174/187152062313230615115156. [DOI] [PubMed] [Google Scholar]
  40. Luo B., Kastrat E., Morcol T., Cheng H., Kennelly E., Long C. Gaultheria longibracteolata, an alternative source of wintergreen oil. Food Chemistry. 2021;342:128244. doi: 10.1016/j.foodchem.2020.128244. [DOI] [PubMed] [Google Scholar]
  41. Luo X. Comparative analysis of volatile oil content in leaves, rhizomes and tuberous roots of cureuma wang-siensis. SG lee et CF liang by GC/MS. China Pharmacist. 2014;12:1659–1661. [Google Scholar]
  42. Mau J.-L., Lai E.Y., Wang N.-P., Chen C.-C., Chang C.-H., Chyau C.-C. Composition and antioxidant activity of the essential oil from curcuma zedoaria. Food Chemistry. 2003;82:583–591. doi: 10.1016/S0308-8146(03)00014-1. [DOI] [Google Scholar]
  43. Mossmann V., Weimer P., Nunes K.A.A., Rossi R.C., Koester L.S. Essential oil extracted from the leaves of curcuma longa l.: Application of an agro-industrial residue in the development of anti-inflammatory nanoemulsions intended for skin delivery. Chemistry Africa. 2024;7:1849–1864. doi: 10.1007/s42250-023-00872-4. [DOI] [Google Scholar]
  44. National Pharmacopoeia Commission . China Medical Science and Technology Press; Beijing: 2020. China Pharmacopoeia; p. 233. [Google Scholar]
  45. Nugraheni B., Rohman A., Susidarti R.A., Purwanto P. Influence of geographical origin on essential oil contents in curcuma aeruginosa Roxb. rhizomes in central java, Indonesia. Tropical Journal of Natural Product Research. 2024;8 doi: 10.26538/tjnpr/v8i12.36. [DOI] [Google Scholar]
  46. Pantharos P., Sukcharoen P., Phadungrakwittaya R., Akarasereenont P., Booranasubkajorn S., Lumlerdkij N. Utilization of UPLC-PDA and GC-MS/MS coupled with metabolomics analysis to identify bioactive metabolites in medicinal turmeric at different ages for the quality assurance. Phytomedicine. 2022;102 doi: 10.1016/j.phymed.2022.154157. [DOI] [PubMed] [Google Scholar]
  47. Quemel F.d.S., Dantas A.P., Sanches L., Alves Viana A.C.G., Silva E.S., Monteiro E.R., Gazim Z.C., Gonçalves J.E., Lopes A.D. Chemotypes of turmeric ('curcuma longa' L.) essential oil from four different states of Brazil. Australian Journal of Crop Science. 2021;15:1035–1042. [Google Scholar]
  48. Rebiai A., Hemmami H., Zeghoud S., Ben Seghir B., Kouadri I., Eddine L.S., Elboughdiri N., Ghareba S., Ghernaout D., Abbas N. Current application of chemometrics analysis in authentication of natural products: A review. Combinatorial Chemistry & High Throughput Screening. 2022;25:945–972. doi: 10.2174/1386207324666210309102239. [DOI] [PubMed] [Google Scholar]
  49. Ren Y., Lin F., Meng J., Liu Y., Li Y., Zhao W., Zhao R., Zhu D., Liu Y. Characterization of potential bioactive molecules in fissistigma polyanthum using UPLC-ESI-QTOF-MS-based metabolomics integrated with chemometrics approaches. Journal of Chromatography A. 2025;1746 doi: 10.1016/j.chroma.2025.465804. [DOI] [PubMed] [Google Scholar]
  50. Riaz A., Rasul A., Kanwal N., Hussain G., Shah M.A., Sarfraz I., Ishfaq R., Batool R., Rukhsar F. Germacrone: A potent secondary metabolite with therapeutic potential in metabolic diseases, cancer and viral infections. Current Drug Metabolism. 2020;21:1079–1090. doi: 10.2174/1389200221999200728144801. [DOI] [PubMed] [Google Scholar]
  51. Sasikumar B. Genetic resources of curcuma: Diversity, characterization and utilization. Plant Genetic Resources. 2005;3:230–251. doi: 10.1079/PGR200574. [DOI] [Google Scholar]
  52. Setzer W.N., Duong L., Poudel A., Mentreddy S.R. Variation in the chemical composition of five varieties of curcuma longa rhizome essential oils cultivated in North alabama. Foods. 2021;10:212. doi: 10.3390/foods10020212. [DOI] [PMC free article] [PubMed] [Google Scholar]
  53. Sharma S.K., Singh S., Tewari S.K. Study of leaf oil composition from various accessions of curcuma longa l. grown on partially reclaimed sodic soil. International Journal of Plant and Environment. 2019;5:293–296. doi: 10.18811/ijpen.v5i04.10. [DOI] [Google Scholar]
  54. Silva B.I., Nascimento E.A., Silva C.J., Silva T.G., Aguiar J.S. Anticancer activity of monoterpenes: A systematic review. Molecular Biology Reports. 2021;48:5775–5785. doi: 10.1007/s11033-021-06578-5. [DOI] [PubMed] [Google Scholar]
  55. Simoh S., Shin S.Y., Abd Rahim F., Ahmad M.A., Zainal A. Comparative analysis of metabolites and antioxidant potentials from different plant parts of curcuma aeruginosa roxb. Sains Malaysiana. 2018;47:3031–3041. [Google Scholar]
  56. Su Y.C., Hsu K.P., Wang E.I.C., Ho C.L. Composition, in vitro cytotoxic, and antimicrobial activities of the flower essential oil of diospyros discolor from Taiwan. Natural Product Communications. 2015;10:1311–1314. doi: 10.1177/1934578X15010007. [DOI] [PubMed] [Google Scholar]
  57. Sun S., Yu Y., Jo Y., Han J.H., Xue Y., Cho M.…Zhuang S. Impact of extraction techniques on phytochemical composition and bioactivity of natural product mixtures. Frontiers in Pharmacology. 2025;16 doi: 10.3389/fphar.2025.1615338. [DOI] [PMC free article] [PubMed] [Google Scholar]
  58. Sun W., Wang S., Zhao W., Wu C., Guo S., Gao H., Tao H., Lu J., Wang Y., Chen X. Chemical constituents and biological research on plants in the genus curcuma. Critical Reviews in Food Science and Nutrition. 2017;57:1451–1523. doi: 10.1080/10408398.2016.1176554. [DOI] [PubMed] [Google Scholar]
  59. Theanphong O., Mingvanish W., Kirdmanee C. Chemical constituents and biological activities of essential oil from curcuma aeruginosa roxb. Rhizome. Interprofessional Journal of Health Sciences. 2015;13:6–16. [Google Scholar]
  60. Thu N.T., Tran-Trung H., Huong H.T.D., Manh D.H., Van Trung H., Tuan N.H., Ha D.T. Chemical compositions and cytotoxicity effects of essential oils from the leaves and pseudo-stems of curcuma zedoaroides chaveer. & tanee collected in Vietnam. Journal of Biologically Active Products from Nature. 2024;14:161–170. doi: 10.1080/22311866.2024.2341730. [DOI] [Google Scholar]
  61. Varia R.D., Patel J.H., Modi F.D., Vihol P.D., Bhavsar S.K. In vitro and in vivo antibacterial and anti-inflammatory properties of linalool. International Journal of Current Microbiology and Applied Sciences. 2020;9:1481–1489. doi: 10.20546/ijcmas.2020.909.187. [DOI] [Google Scholar]
  62. Viveiros M.M.H., Silva M.G., da Costa J.G.M., de Oliveira A.G., Rubio C., Padovani C.R., Rainho C.A., Schellini S.A. Anti-inflammatory effects of α-humulene and β-caryophyllene on pterygium fibroblasts. International Journal of Ophthalmology. 2022;15:1903. doi: 10.18240/ijo.2022.12.02. [DOI] [PMC free article] [PubMed] [Google Scholar]
  63. Wang J., Hou Y., Bai Z., Fu M., Zeng J., Huang F., Diao K., Chen X. Screening the universal sequence of DNA barcodes in curcuma kwangsiensis. China Journal of Traditional Chinese Medicine and Pharmacy. 2015;5:1417–1427. [Google Scholar]
  64. Wang J., Qian C., Chen Y., Jin T., Jiang Y., Huang L., Fu X., Yang D., Jin L., Jin B. Β-Elemene alleviates hyperglycemia-induced cardiac inflammation and remodeling by inhibiting the JAK/STAT3-NF-κB pathway. Phytomedicine. 2023;119 doi: 10.1016/j.phymed.2023.154987. [DOI] [PubMed] [Google Scholar]
  65. Wang P., Henning S.M., Heber D. Limitations of MTT and MTS-based assays for measurement of antiproliferative activity of green tea polyphenols. PLoS One. 2010;5 doi: 10.1371/journal.pone.0010202. [DOI] [PMC free article] [PubMed] [Google Scholar]
  66. Wang Z., Guan L., Song W., Yang B., Tuersuntuoheti T., Zhang M., Han J., Sun Z. Integrated volatilomics and metabolomics reveal the dynamic landscape and key transformation phases of flavor compounds in indica rice during hydrothermal cooking. Food Chemistry: X. 2026;34 doi: 10.1016/j.fochx.2026.103557. [DOI] [PMC free article] [PubMed] [Google Scholar]
  67. Westerhuis J.A., Hoefsloot H.C., Smit S., Vis D.J., Smilde A.K., van Velzen E.J., van Duijnhoven J.P., van Dorsten F.A. Assessment of PLSDA cross validation. Metabolomics. 2008;4:81–89. [Google Scholar]
  68. Xiang H., Zhang L., Xi L., Yang Y., Wang X., Lei D., Zheng X., Liu X. Phytochemical profiles and bioactivities of essential oils extracted from seven curcuma herbs. Industrial Crops and Products. 2018;111:298–305. doi: 10.1016/j.indcrop.2017.10.035. [DOI] [Google Scholar]
  69. Xiao Y., Yang F.Q., Li S.P., Hu G., Lee S.M.Y., Wang Y.T. Essential oil of curcuma wenyujin induces apoptosis in human hepatoma cells. World Journal of Gastroenterology. 2008;14:4309–4318. doi: 10.3748/wjg.14.4309. [DOI] [PMC free article] [PubMed] [Google Scholar]
  70. Xu Y., Zhang M., Yang W., Xia B., Wang W., Pan X. Nootkatone protects cartilage against degeneration in mice by inhibiting NF-κB signaling pathway. International Immunopharmacology. 2021;100 doi: 10.1016/j.intimp.2021.108119. [DOI] [PubMed] [Google Scholar]
  71. Yang B., Wang W., Zhang J., Gao W., Fan L., Chitrakar B., Sang Y. Comparative study on organoleptic properties and volatile organic compounds in turmeric, turmeric essential oil, and by-products using e-nose, HS-GC-IMS, and HS-GC-MS. Food Chemistry: X. 2025;25 doi: 10.1016/j.fochx.2024.102107. [DOI] [PMC free article] [PubMed] [Google Scholar]
  72. Záveská E., Fér T., Šída O., Krak K., Marhold K., Leong-Škorničková J. Phylogeny of curcuma (zingiberaceae) based on plastid and nuclear sequences: Proposal of the new subgenus ecomata. Taxon. 2012;61:747–763. doi: 10.1002/tax.614004. [DOI] [Google Scholar]
  73. Zhang L., Chen Y., Li Z., Fan G., Li X. Production, function, and applications of the sesquiterpenes valencene and nootkatone: A comprehensive review. Journal of Agricultural and Food Chemistry. 2022;71:121–142. doi: 10.1021/acs.jafc.2c07543. [DOI] [PubMed] [Google Scholar]
  74. Zhang L., Yang Z., Chen D., Huang Z., Li Y., Lan X., Su P., Pan W., Zhou W., Zheng X. Variation on composition and bioactivity of essential oils of four common curcuma herbs. Chemistry & Biodiversity. 2017;14 doi: 10.1002/cbdv.201700280. [DOI] [PubMed] [Google Scholar]
  75. Zhang L., Yang Z., Huang Z., Zhao M., Li P., Zhou W., Zhang K., Zheng X., Lin L., Tang J. Variation in essential oil and bioactive compounds of curcuma kwangsiensis collected from natural habitats. Chemistry & Biodiversity. 2017;14 doi: 10.1002/cbdv.201700020. [DOI] [PubMed] [Google Scholar]
  76. Zhang R., Ji Y., Morcol T., Lin F., Gu R., Kennelly E.J., Long C. UPLC-QTof-MS chemical profiling and characterization of antiproliferative and anti-inflammatory compounds from seven hypericum species in China. Industrial Crops and Products. 2021;173 doi: 10.1016/j.indcrop.2021.114156. [DOI] [Google Scholar]
  77. Zhu J., Lower-Nedza A.D., Hong M., Jiec S., Wang Z., Yingmao D., Tschiggerl C., Bucar F., Brantner A.H. Chemical composition and antimicrobial activity of three essential oils from curcuma wenyujin. Natural Product Communications. 2013;8 doi: 10.1177/1934578X1300800430. [DOI] [PubMed] [Google Scholar]

Associated Data

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

Supplementary Materials

Supplementary material. Table S1. Chemical composition of EOs from different plant parts of C. kwangsiensis across different geographical locations. Table S2. Quality parameters for PLS-DA plots in Figure 2. Table S3. Quality parameters for PLS-DA plots in Figure 3. Fig. S1. Cytotoxic effects of EO samples from C. kwangsiensis on three cancer cell lines at 100 μg/mL.

mmc1.docx (180.4KB, 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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