Influence of steviol glycosides on the stability of anthocyanins during storage of blueberry juice

Chunxue Zhang; Lin Zhang; Xinmei Yun; Ahmed Adel Ashour; Hui Tan; Yuehua Wang; Xinyao Jiao; Chunxue Zhang; Lin Zhang; Xinmei Yun; Ahmed Adel Ashour; Hui Tan; Yuehua Wang; Xinyao Jiao

doi:10.48130/fia-0026-0019

Blueberry juice maintains the nutritional value of fresh berries while providing their complex flavor profile. Anthocyanins contribute to the vibrant color of blueberry juice and exhibit physiological functions. Steviol glycosides are widely utilized as a natural sweetener in fruit beverages, and investigating their regulatory role in the degradation of anthocyanins within juice systems is of considerable importance. This study aimed to examine the effects of steviol glycosides on the color, anthocyanin content, antioxidant capacity, and in vitro simulated digestion stability of anthocyanins in blueberry juice during storage. The juice was stored for 90 d at 4 and 25 °C in dark conditions. Throughout storage, the juice exhibited color fading, accompanied by a loss of its original color intensity. Notably, the sample supplemented with 20 mg/100 mL steviol glycosides showed the least color change when stored at 4 °C. The addition of steviol glycosides resulted in a reduced degradation rate constant (k) and an extended half-life (t_1/2) of anthocyanins. Thermodynamic analysis indicated that steviol glycosides enhanced the thermal stability of anthocyanins in blueberry juice. Furthermore, blueberry juice containing 12 and 16 mg/100 mL steviol glycosides demonstrated the highest hydroxyl radical scavenging capacity at 4 °C, and the strongest ABTS radical scavenging activity at 25 °C, respectively. During in vitro simulated digestion, the sample with 4 mg/100 mL steviol glycosides exhibited the highest retention rate of total anthocyanins. Molecular docking analysis revealed the formation of hydrophobic interactions and hydrogen bonds between stevioside and cyanidin-3-O-glucoside, as well as malvidin-3-O-galactoside. The findings of this study provide a solid foundation for advancing fruit juice processing technologies.

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Introduction

Blueberry (Vaccinium spp.) fruits are distinguished by their sweet-sour flavor and delicate texture, as well as their abundance of antioxidant compounds, which confer significant nutritional and economic value^[1]. However, postharvest blueberries are prone to rapid softening and decay, limiting their storage and transport potential. Consequently, processed blueberry products represent a vital strategy for extending the industrial chain^[2]. Among these, blueberry juice stands out as one of the most popular consumer products, as it preserves most of the nutritional components of fresh berries while delivering a rich flavor profile. Nevertheless, during processing and storage, environmental factors such as temperature, pH, oxygen, and light can accelerate the degradation of bioactive compounds of blueberry juice, resulting in nutrient loss and color fading. This challenge constitutes a major constraint for the development of blueberry juice beverages^[3]. Blueberries are notably rich in anthocyanins, whose physiological functions including antioxidant, anti-inflammatory, and gut microbiota regulation, have been extensively confirmed and recognized^[4]. However, the degradation of anthocyanins during storage not only compromises their potential health benefits, but also affects the product quality in the beverage industry. Therefore, enhancing the stability of anthocyanins has emerged as a key research priority in recent years.

Steviol glycosides, predominantly found in the leaves of perennial herbs, share a common sweet-tasting backbone structure, ent-kaurenoic acid. Among these compounds, stevioside, and rebaudioside-A (Reb-A) are the two most abundant glycosides^[5]. As a natural sweetener, steviol glycosides are characterized by a high sweetness intensity and a low caloric content, in addition to diverse biological functions such as antioxidant activity, blood glucose regulation, and gastrointestinal health improvement^[6,7]. In food processing, substituting sucrose with steviol glycosides can effectively overcome drawbacks associated with artificial sweeteners, such as metabolic disorders like obesity and diabetes, thereby aligning with contemporary health-conscious trends. As a result, steviol glycosides are widely used in beverages to partially or fully replace sucrose, demonstrating considerable potential for application across the beverage industry^[6,8].

In real juice systems, the addition of steviol glycosides as a low-calorie natural sweetener may substantially influence the stability of anthocyanins by modifying the physicochemical environment or facilitating intermolecular interactions. Previous studies have shown that steviol glycosides exert protective effects on anthocyanins, color, and antioxidant activity in sour cherry puree over a 6-month storage period^[9]. Comparable protective effects were reported in lingonberry jam, where steviol glycosides proved the most effective in preserving anthocyanin antioxidant capacity^[10]. Chokeberry juices with added steviol glycosides had a higher content of analyzed bioactive compounds in comparison with juice samples sweetened with sucrose, and bioactive compounds, and antioxidant capacity was not significantly reduced during the storage period^[11]. Furthermore, researchers have indicated that citrus-maqui juice supplemented with steviol glycosides maintains notable color stability, retaining its red hue and natural appearance throughout the shelf life^[12]. Similarly, adding steviol glycosides lead to the color of strawberry jam being substantially brighter^[13]. Degradation kinetic models of anthocyanins in citrus-maqui juice further suggest that molecular interactions between anthocyanins and steviol glycosides contribute to anthocyanin stabilization^[14]. However, most of these studies remain largely confined to the validation of macroscopic effects, with extremely limited investigation into the specific binding modes and interaction mechanisms at the molecular level. Although the application of spectroscopic techniques, molecular docking, and molecular dynamics simulations can elucidate the fundamental nature of this interaction, relevant studies have rarely been reported in the context of anthocyanin and steviol glycoside systems.

The objective of this study was to examine the influence of steviol glycosides on the color, stability, and degradation kinetics of total anthocyanin and antioxidant capacity of blueberry juice under different storage conditions. Additionally, this work explores the mechanism of non-covalent molecular interactions between steviol glycosides and anthocyanins to elucidate the underlying stabilizing mechanism. The findings are expected to provide a theoretical foundation for enhancing the nutritional quality of blueberry juice and support further advances in juice processing technology.

Materials and methods

Materials and reagents

Blueberry fruits of the 'Northland' cultivar were sourced from a local farm in Shenyang, Liaoning Province, China. Pectinase (30,000 U/g), pepsin (1:3,000), porcine trypsin (1:250), and porcine bile salts were procured from Shanghai Macklin Biochemical Co., Ltd. (Shanghai, China). Assay kits for determining DPPH radical scavenging capacity, ABTS radical scavenging capacity, and hydroxyl radical (•OH) content were supplied by Suzhou Grace Biotechnology Co., Ltd. (Suzhou, China). All other chemicals used were of analytical grade (purity ≥ 98%). The sweeteners employed in this study were commercially obtained from a health food retailer.

Samples preparation
Blueberry fruit (1 kg) were homogenized with 1 L of deionized water using a crusher. Subsequently, 2 g of pectinase (30,000 U/g) was added to the pulp, and the mixture was stirred for 1 h to facilitate enzymatic hydrolysis, yielding a crude blueberry extract. The crude extract was then centrifuged at 3,500 r/min for 20 min, and the supernatant was collected. Aliquots (100 mL) of the juice were supplemented with sweeteners; specifically, steviol glycosides (at 2, 4, 8, 12, 16, or 20 mg), sucralose (4 mg), or sucrose (7.5 g), along with 150 mg/L of benzoic acid. After thorough mixing, the samples were centrifuged and filtered. The pH of the resulting solution was adjusted to 3.5 using 1 mol/L HCl or NaOH, thus obtaining the final blueberry juice (BBJ) solution. A total of nine distinct juice formulations were prepared, with each type replicated three times.

The concentrations of sucralose and sucrose were selected based on consumer sweetness preference. In contrast, the 4 mg dose of steviol glycosides was determined from literature data to achieve an equivalent sweetness intensity^[12,15]. The different doses of steviol glycosides were administered to investigate the dose-dependent protective effect of steviol glycosides on anthocyanins. The Brix and pH of the BBJ samples have been provided in (Supplementary Tables S1 and S2). All BBJ samples were stored under dark conditions at 4 and 25 °C for 90 d. Analyses were conducted immediately after preparation (day 0) and at 10-d intervals throughout the storage period.

Color measurements

The chromaticity parameters (L*, a*, b*) of the samples were measured using a colorimeter (NR60CP+, Shenzhen Threenh Technology Co., Ltd., China). The L* value represents lightness, while a* and b* denote the degree of redness/greenness and yellowness/blueness, respectively. The chroma (C*_ab) and total color difference (ΔE*_ab) were calculated according to Eqs. (1) and (2), as follows:

$ \mathrm{C}_{\text{ab}}^{\mathrm{*}}=\sqrt{{\left( {\mathrm{a}}^{\mathrm{*}}\right) }^{2}{+\left( {\mathrm{b}}^{\mathrm{*}}\right) }^{2}} $

(1)

$ \Delta \mathrm{E}=\sqrt{{\left( {\Delta \mathrm{L}}^{\mathrm{*}}\right) }^{2}+{\left( {\Delta \mathrm{a}}^{\mathrm{*}}\right) }^{2}{+\left( {\Delta \mathrm{b}}^{\mathrm{*}}\right) }^{2}} $

(2)

In this equation, ΔL*, Δa*, and Δb* represent the differences in L*, a*, and b* between the treatment group and the control group, respectively.

Determination of total anthocyanins content (TAC)

The total anthocyanin content (TAC) was determined using the pH differential method^[16]. Briefly, 1 mL of blueberry juice was diluted with pH 1.0 and 4.5 buffer solutions to a final volume of 20 mL, respectively. After thorough mixing, solutions were incubated in the dark for 60 min, and the absorbance of each solution was measured at 520 and 700 nm. The absorbance difference was calculated using Eq. (3), and the TAC was subsequently determined according to Eq. (4).

$ \rm A={\left({{A}}_{{520}}-{{A}}_{{700}}\right)}_{{pH1.0}}-{\left({{A}}_{{520}}-{{A}}_{{700}}\right)}_{{pH4.5}} $

(3)

$ \mathrm{TAC}=\dfrac{\left( \mathrm{A}\times \text{MW}\times \text{DF}\times 1,000\right) }{\varepsilon \times 1} $

(4)

In this equation, MW is the relative molecular weight of anthocyanins in the sample (449.2 g/mol); DF is the dilution factor of the sample; ε is the molar absorptivity of anthocyanins in the sample (26,900).

Thermodynamic parameters measurements

The thermodynamic parameters were determined following the method described by Cao et al.^[17]. Briefly, BBJ samples were incubated at 333, 343, 353, and 363 K for 5 h, with aliquots collected hourly for anthocyanin content analysis. The degradation rate constant (k), Gibbs free energy change (ΔG), enthalpy change (ΔH), and entropy change (ΔS) were calculated based on first-order kinetic equations and the Arrhenius equation, as established by Brouillard et al. and Lambert et al.^[18,19].

Degradation rate (k) was calculated using Eq. (5).

First-order kinetic equation:

$ \ln \left({\mathrm{C}}_{\mathrm{t}}-{\mathrm{C}}_{0}\right)=-\text{kt} $

(5)

In this equation, C₀ is the initial concentration of anthocyanin; C_t is the concentration of anthocyanin after heating time; t is the heating time (h) ; k is the rate constant.

Anthocyanin thermal degradation half-life (t_1/2) was calculated using Eq. (6).

$ {\mathrm{t}}_{{1}/{2}}=-\dfrac{\ln \left( 0.5\right) }{\mathrm{k}} $

(6)

Activation energy (E_a) was calculated using the Arrhenius equation (Eq. [7]).

$ \mathrm{lnk}=\ln\mathrm{K}_0-\left(\mathrm{E}_{\mathrm{a}}/\text{RT}\right) $

(7)

In this equation, E_a is the activation energy (kJ/mol) ; R is the gas constant (8.314 J/mol·K); T is the absolute temperature (K); K₀ is the frequency factor (min⁻¹) .

Enthalpy change (ΔH), Gibbs free energy (ΔG), and entropy change (ΔS) were calculated using Eqs. (8)–(10).

$ \Delta \mathrm{H}={\mathrm{E}}_{\mathrm{a}}-\text{RT} $

(8)

$ \Delta \mathrm{G}=-\text{RTln}\left(\dfrac{\text{kh}}{{\mathrm{k}}_{\mathrm{B}}\mathrm{T}}\right) $

(9)

$ \Delta \mathrm{S}=\dfrac{\Delta \mathrm{H}-\Delta \mathrm{G}}{\mathrm{T}} $

(10)

In Eq. (8), h is Planck's constant (6.6262 × 10⁻³⁴ J·s); k_B is the Boltzmann constant (1.3806 × 10⁻²³J/K).

Antioxidant capacity

The hydroxyl radical (•OH) and ABTS radical scavenging capacities of the BBJ samples were determined using commercial assay kits (Suzhou Grace Biotechnology Co., Ltd., Suzhou, China) in accordance with the manufacturer's instructions.

Hydroxyl radical scavenging capacity
A 50 μL aliquot of BBJ sample was mixed with the assay reagents. The absorbance was measured at 510 nm using a microplate reader, and the hydroxyl radical scavenging activity was calculated as described in the kit protocol.

ABTS radical scavenging capacity
A 10 μL aliquot of BBJ sample was combined with the assay reagents. The absorbance was recorded at 734 nm using a microplate reader, and the ABTS radical scavenging activity was calculated according to the manufacturer's instructions.

In vitro simulated digestion

In vitro simulated digestion was performed according to a previously described method with minor modifications^[20].

Simulated gastric digestion: a 1 mL aliquot of BBJ sample was accurately pipetted and diluted 20-fold with saline. The diluted sample was incubated in a water bath at 37 °C, and the pH was adjusted to 2.0 using 1 mol/L HCl. Then, 4 mL of simulated gastric juice was added. The mixture was placed in a refrigerated constant-temperature shaker and digested at 37 °C for 2 h. Aliquots were collected at 0, 1, and 2 h, centrifuged at 4 °C and 8,000 r/min for 15 min, and the supernatant was collected for TAC analysis.

Simulated intestinal digestion: after gastric digestion, the pH of the sample was adjusted to 7.0 using 1 mol/L NaHCO₃ solution. Then, 4 mL of trypsin–bile salt mixture was added. The sample was further digested in a refrigerated constant-temperature shaker at 37 °C for 2 h. Aliquots were taken at 0, 1, and 2 h, centrifuged under the same conditions as above, and the supernatant was used for TAC determination.

$ \mathrm{Retention\;rate\;({\text{%}})}=\rm C_{ \mathrm{digested}} \mathrm{/C}_{ \mathrm{undigested}} \times \mathrm{ 100} $

(11)

In this equation, C_digested is the TAC after gastric and intestinal digestion; C_undigested is the TAC without digestion.

Simulation of the interaction between anthocyanins and steviol glycosides

The three-dimensional (3D) structures of the small molecules were obtained from the PubChem database and energy-minimized using Discovery Studio 2019 before being saved in PDB format. The resulting files were imported into AutoDock Tools, where hydrogen atoms were added, and then water removal was performed. Steviol glycosides and anthocyanins were designated as the receptors and ligands, respectively, and both were converted into PDBQT format for molecular docking. Docking simulations were carried out using AutoDock Vina to estimate binding affinities. Finally, the docking poses were visualized and analyzed using PyMOL 3.1.

Statistical analysis
Data are expressed as mean ± standard deviation (SD) based on three replicates. Differences between the two groups were assessed using Student's t-tests. The results were considered statistically significant at p < 0.05. Data were analyzed using GraphPad Prism 8.0 (Graph Pad Software, San Diego, CA, USA) and SPSS 17.0 (IBM Corporation, Armonk, NY, USA).

Discussion

This study aimed to systematically evaluate the protective effect of steviol glycosides on anthocyanins in BBJ from a multi-scale perspective. First, the macroscopic manifestations were investigated, including color changes and degradation kinetics of anthocyanins during storage. Subsequently, the thermodynamic behavior and the in vitro simulated digestion were explored to understand the stability of anthocyanins. Finally, molecular docking was employed to probe the potential molecular-scale interactions, aiming to reveal the mechanism underlying the observed stabilization.

The susceptibility of anthocyanins to light, heat, and oxygen presents a major challenge for their application in the food industry. Under varying environmental conditions such as pH, temperature, and light, the structures of anthocyanins undergo a transition from the stable red flavylium cation, to lightly colored or nearly colorless forms, including methanol pseudobase, chalcone, and quinoidal base, under equilibrium conditions^[22]. Current strategies to enhance the stability of anthocyanins primarily fall into two categories: one focuses on modulating extrinsic factors such as environmental conditions, while the other aims at structural stabilization through intrinsic modifications to preserve color and reduce degradation rate^[17,23,24]. To date, research on environmental influences is relatively well-established, providing a solid theoretical basis for optimizing BBJ processing and storage. Effective approaches such as copigmentation, acylation, and biosynthesis have been shown to improve the stability and color retention of anthocyanins, thereby facilitating their broader application in the food and beverage processing^[25,26].

Anthocyanins are highly susceptible to nucleophilic attack by water molecules in aqueous solutions. The flavylium cation undergoes hydroxylation at the C-2 position of its C-ring, forming a hemiketal structure. This hemiketal and the corresponding chalcone glycoside exist in equilibrium via tautomerization. After deglycosylation, the chalcone degrades under specific conditions, ultimately leading to the decoloration of anthocyanins. Anthocyanins-copigment complexes form conjugated systems through non-covalent interactions, adopting a sandwich-like configuration that shields the central anthocyanin from nucleophilic attack by water molecules. This structural arrangement inhibits hydration of the flavylium cation chromophore, thereby enhancing both stability and color integrity. These complexes are primarily stabilized by van der Waals forces, hydrophobic interactions, or ionic bonds, with common copigments including phenolic acids, amino acids, polysaccharides, flavonoids, proteins, and metal ions^[27]. The observed color fading in all BBJ samples, characterized by an increase in L* value, and a decrease in a*, b*, and C*_ab values can be attributed to two primary chemical processes: the degradation of anthocyanins, and the progression of non-enzymatic browning reactions during storage. Based on the ΔE*_ab values, almost all BBJ samples exhibited visible color changes on day 30 (ΔE*_ab> 5)^[28]. Notably, the significantly smaller total color difference (ΔE*_ab) in specific steviol glycoside groups, particularly Ste20 at 4 °C and Ste4 at 25 °C, indicate steviol glycosides have better color retention, and could effectively decelerate these underlying chemical pathways leading to color deterioration. The underlying protective mechanism may be attributed to the ability of steviol glycosides to form non-covalent complexes with anthocyanins in the juice. This interaction reduces the accessibility of the C-2 position on the C-ring of anthocyanin flavylium cation to nucleophilic attack by water molecules, thereby preventing the degradation of anthocyanins; this hypothesis is further supported by molecular docking results. From day 0 to 90, the content of anthocyanins has decreased by 62.37%~71.56% at 4 °C and 77.65%~85.29% at 25 °C. The stability of anthocyanins and their degradation is influenced by a number of factors such as pH, light, temperature, co-pigmentation, sulfites, ascorbic acid, oxygen, and enzymes^[29]. The effect of color stabilization provides visual evidence for the subsequent kinetic data, which quantitatively confirms a slower degradation rate (k) and an extended half-life (t_1/2) of anthocyanins in steviol glycosides-treated groups, a 23-d extension of half-life (t_1/2) compared to the control at 4 °C. Although these colorimetric trends align with established literature, one study reported no statistically significant differences in color between stevia and sucrose-sweetened beverages^[12]. Woźniak et al. reported that steviol glycosides did not influence the degradation kinetics of anthocyanins, and found that a combination of sucrose and steviol glycosides provided optimal anthocyanin stabilization^[15].

Thermodynamic analysis revealed that steviol glycosides significantly enhanced the thermal stability of anthocyanins in BBJ, particularly at moderate temperatures. At 333 K, the Ste16 group showed markedly improved stability, with a 30% reduction in the degradation rate constant (k) and a 4 h extension of half-life (t_1/2) compared to the control. Although this protective effect diminished at higher temperatures (343–363 K), key thermodynamic parameters indicated fundamental stabilization mechanisms. Significant increases in activation energy (E_a) and enthalpy change (ΔH) for the Ste4, Ste8 and Ste16 groups suggested higher energy barriers for degradation and stronger intermolecular bonding. The observed increase in Ea and ΔH in steviol glycosides-treated groups is a hallmark of copigmentation phenomena^[30]. In a typical copigmentation complex, the anthocyanins-copigment complexes require additional energy to dissociate before anthocyanin degradation. Positive ΔG values confirmed the endergonic nature of anthocyanin degradation, while reduced entropy (ΔS) in steviol glycosides-treated groups implied enhanced molecular ordering within the system, possibly due to the stacking and reduced mobility of anthocyanin molecules within a protective matrix formed by steviol glycosides, which aligns with the proposed 'sandwich' structure in copigmentation theory^[21,31,32]. These comprehensive thermodynamic analyses affirm that steviol glycosides, particularly at optimal concentrations, stabilize the structure of anthocyanins through a combination of elevated energy barriers and entropic modulation. In line with this, Fan et al. reported that copigmentation induced molecular reordering of anthocyanins toward thermodynamic equilibrium underlies their structural stabilization^[16].

The relationship between BBJ color, anthocyanin content, and antioxidant capacity is well demonstrated^[33]. The antioxidant activity of anthocyanins stems from their molecular geometry, which facilitates hydrogen atom donation from aromatic hydroxyl groups, and is further influenced by specific hydroxylation and methoxylation patterns^[34]. Notably, the preservation of antioxidant activity in our study followed a selective pattern. Although overall antioxidant capacity declined during storage, specific steviol glycoside concentrations conferred targeted protection; Ste12 uniquely maintained hydroxyl radical scavenging capacity at 4 °C, whereas Ste16 retained superior ABTS radical scavenging at 25 °C on day 90. This differential effect suggests that steviol glycosides may stabilize specific anthocyanin subclasses with distinct antioxidant functions, rather than providing broad-spectrum protection. The improved kinetic stability and selective antioxidant preservation indicate that steviol glycosides act as multifunctional stabilizers in BBJ, offering both structural protection to anthocyanins and maintaining key antioxidant pathways. These results highlight the potential of steviol glycosides as a natural additive for extending the shelf life and functional quality of anthocyanin-rich beverages. Consistent with our findings, Nowicka & Wojdylo also reported the protective effects of steviol glycosides on polyphenol content, particularly anthocyanins, and consequently on color and antioxidant activity^[9].

Gastric digestion had no significant effect on anthocyanins, and intestinal digestion substantially decreased anthocyanins^[35]. Our study shows that steviol glycosides significantly influence the gastrointestinal stability of anthocyanins in BBJ. Under gastric conditions, the Ste4 group exhibited the highest retention rate, exceeding that of sucralose and sucrose, whereas Ste12 showed the lowest. This trend continued during intestinal digestion, with Ste4 maintaining the highest retention rate, significantly greater than that of sucralose and sucrose. These results suggest that lower concentrations of steviol glycosides (e.g., Ste4) enhance the stability of anthocyanin throughout digestion, whereas higher concentrations (e.g., Ste12) may have inhibitory effects. Steviol glycosides are superior to artificial and traditional sweeteners in maintaining the stability of anthocyanins during in vitro simulated digestion. This highlights their potential as functional additives for improving the nutritional value of anthocyanins-rich beverages. Previous studies have indicated that the effects of steviol glycosides on stability, bioactivity, and bioavailability of anthocyanins are mediated through molecular interactions^[36,37].

To explore the mechanism by which steviol glycosides enhance the stability of anthocyanins in BBJ, we hypothesized that non-covalent interactions between them may play a crucial role. The most abundant steviol glycoside, stevioside, and the predominant anthocyanins in BBJ, C3G, and M3G, were selected for molecular docking studies. However, the docking results indicated that although the ligand occupies the binding pocket, the interactions are predominantly hydrophobic, and lack the extensive hydrogen-bonding network typically associated with high-affinity binding. This observation aligns with the low binding affinity observed, suggesting that stevioside acts as a weak binder under the simulated conditions. While the molecular docking results indicated only weak binding affinity between stevioside and individual anthocyanins (C3G, M3G), the consistent protective effects observed across color, kinetic, thermodynamic, and bioavailability assays strongly suggest the existence of an alternative or supplementary stabilization mechanism. The stabilization may not rely solely on high affinity, specific binding between single molecules; it could be governed by weak, cumulative, and non-specific interactions within a complex juice matrix. These interactions, such as hydrogen bonding, van der Waals forces, and hydrophobic effects, might collectively create a molecular environment that shields anthocyanins from water, oxygen, and nucleophilic attack. Therefore, future research should focus on alternative analytical techniques, such as isothermal titration calorimetry (ITC) to directly measure the binding stoichiometry and affinity in solution, or nuclear magnetic resonance (NMR) spectroscopy to detect weak intermolecular interactions in a model system that more closely mimics the complex juice matrix.

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Parameter	Days	Control	Ste2	Ste4	Ste8	Ste12	Ste16	Ste20	Sucralose	Sucrose
L*	0	20.63 ± 0.12^a	11.97 ± 0.14^g	18.51 ± 0.06^c	15.44 ± 0.15^e	16.78 ± 0.10^d	15.50 ± 0.08^e	19.04 ± 0.09^b	13.38 ± 0.17^f	18.38 ± 0.38^c
	30	25.12 ± 0.07^d	21.51 ± 0.02^f	24.51 ± 0.14^e	26.78 ± 0.09^a	24.64 ± 0.03^e	25.58 ± 0.02^c	25.53 ± 0.54^c	25.78 ± 0.26^c	26.24 ± 0.11^a
	60	24.73 ± 0.19^d	24.19 ± 0.29^e	24.32 ± 0.02^e	24.59 ± 0.03^d	23.75 ± 0.02^f	25.55 ± 0.06^c	26.19 ± 0.03^b	27.19 ± 0.03^a	26.00 ± 0.08^d
	90	30.74 ± 0.08^a	30.22 ± 0.01^b	28.63 ± 0.04^g	28.78 ± 0.03^e	28.36 ± 0.02ⁱ	28.49 ± 0.04^h	28.72 ± 0.02^f	29.49 ± 0.02^c	29.16 ± 0.02^d
a*	0	9.29 ± 0.14^f	13.18 ± 0.06^b	9.69 ± 0.18^f	11.56 ± 0.07^c	10.69 ± 0.22^d	11.09 ± 0.03^d	10.53 ± 0.18^de	14.52 ± 0.12^a	10.56 ± 0.69^de
	30	7.02 ± 0.06^a	7.06 ± 0.04^a	7.23 ± 0.16^a	5.50 ± 0.16^c	7.04 ± 0.02^a	6.80 ± 0.09^ab	6.71 ± 0.56^ab	6.89 ± 0.22^a	6.96 ± 0.03^a
	60	6.42 ± 0.31^ab	6.52 ± 0.35^ab	7.03 ± 0.07^a	6.89 ± 0.06^a	6.62 ± 0.28^a	6.46 ± 0.12^ab	6.18 ± 0.25^ab	6.89 ± 0.04^a	5.94 ± 0.36^b
	90	4.33 ± 0.12^e	3.90 ± 0.02^f	3.80 ± 0.07^g	4.83 ± 0.04^b	4.44 ± 0.02^d	4.75 ± 0.04^b	5.31 ± 0.01^a	4.52 ± 0.03^d	4.63 ± 0.05^c
b*	0	−3.17 ± 0.11^b	−6.39 ± 0.15^f	−4.11 ± 0.12^d	−2.79 ± 0.17^a	−3.29 ± 0.13^b	−4.31 ± 0.29^d	−2.58 ± 0.18^a	−4.48 ± 0.14^de	−3.48 ± 0.13^bc
	30	0.41 ± 0.05^de	−1.83 ± 0.05^g	0.47 ± 0.05^d	0.62 ± 0.03^c	0.43 ± 0.03^de	0.53 ± 0.05^d	0.88 ± 0.03^b	1.27 ± 0.09^a	0.16 ± 0.01^f
	60	−0.06 ± 0.01^b	−0.28 ± 0.02^d	0.22 ± 0.04^a	0.24 ± 0.02^a	−0.19 ± 0.01^c	−0.22 ± 0.03^c	−0.34 ± 0.01^e	−0.07 ± 0.01^b	−0.51 ± 0.04^f
	90	−1.12 ± 0.03^e	−1.02 ± 0.10^d	−0.62 ± 0.04^b	−0.38 ± 0.02^a	−0.70 ± 0.05^c	−0.59 ± 0.03^b	−0.59 ± 0.05^b	−0.96 ± 0.04^d	−0.61 ± 0.02^b
C*_ab	0	9.82 ± 0.17^f	14.65 ± 0.01^b	10.53 ± 0.13^de	11.90 ± 0.10^c	11.18 ± 0.20^d	11.90 ± 0.13^c	10.84 ± 0.13^d	15.19 ± 0.07^a	11.13 ± 0.62^d
	30	7.03 ± 0.06^a	7.29 ± 0.03^a	7.24 ± 0.16^a	5.54 ± 0.15^c	7.05 ± 0.02^a	6.82 ± 0.08^ab	6.77 ± 0.55^ab	7.01 ± 0.21^a	6.97 ± 0.03^a
	60	6.42 ± 0.31^ab	6.53 ± 0.35^ab	7.03 ± 0.07^a	6.89 ± 0.06^a	6.62 ± 0.28^a	6.46 ± 0.12^ab	6.19 ± 0.25^b	6.89 ± 0.04^a	5.97 ± 0.36^b
	90	4.47 ± 0.11^d	4.04 ± 0.00^e	3.85 ± 0.08^f	4.85 ± 0.04^b	4.50 ± 0.03^d	4.78 ± 0.04^b	5.34 ± 0.01^a	4.62 ± 0.03^c	4.67 ± 0.05^c
ΔE*_ab	0	0.00 ± 0.00	10.03 ± 0.12^a	2.36 ± 0.03^e	5.68 ± 0.12^c	4.10 ± 0.08^d	5.56 ± 0.10^c	2.11 ± 0.17^ef	9.04 ± 0.19^b	2.63 ± 0.66^e
	30	6.18 ± 0.01^e	2.75 ± 0.01^g	5.71± 0.11^f	8.16 ± 0.14^a	5.84 ± 0.03^f	6.66 ± 0.05^d	6.88 ± 0.51^d	7.21 ± 0.30^b	6.93 ± 0.08^c
	60	5.90 ± 0.02^d	5.36 ± 0.36^e	5.50 ± 0.01^e	5.75 ± 0.05^d	5.08 ± 0.15^f	6.40 ± 0.02^c	6.98 ± 0.08^b	7.64 ± 0.02^a	6.87 ± 0.11^b
	90	11.45 ± 0.02^a	11.21 ± 0.03^b	10.04 ± 0.03^d	9.70 ± 0.02^e	9.45 ± 0.03^f	9.44 ± 0.01^f	9.38 ± 0.02^g	10.30 ± 0.01^c	10.06 ± 0.04^d
Different superscript letters indicate significant differences among groups at the same time point (p < 0.05).

Parameter	Days	Control	Ste2	Ste4	Ste8	Ste12	Ste16	Ste20	Sucralose	Sucrose
L*	0	20.63 ± 0.12^a	11.97 ± 0.14^g	18.51 ± 0.06^c	15.44 ± 0.15^e	16.78 ± 0.10^d	15.50 ± 0.08^e	19.04 ± 0.09^b	13.38 ± 0.17^f	18.38 ± 0.38^c
	30	16.15 ± 0.47^b	10.85 ± 0.08^f	16.03 ± 0.33^b	12.20 ± 0.09^e	12.21 ± 0.15^e	15.35 ± 0.23^c	18.83 ± 0.12^a	13.67 ± 0.32^d	18.68 ± 0.46^a
	60	22.84 ± 0.04^a	17.34 ± 0.17^h	19.31 ± 0.03^f	22.06 ± 0.26^d	20.94 ± 0.08^b	20.20 ± 0.13^c	19.72 ± 0.29^e	18.39 ± 0.22^g	19.78 ± 0.13^e
	90	31.27 ± 0.04^a	29.37 ± 0.02^g	29.65 ± 0.03^f	28.72 ± 0.02^h	30.00 ± 0.01^d	29.84 ± 0.01^e	31.21 ± 0.03^b	30.44 ± 0.02^c	28.06 ± 0.02ⁱ
a*	0	9.29 ± 0.14^f	13.18 ± 0.06^b	9.69 ± 0.18^f	11.56 ± 0.07^c	10.69 ± 0.22^d	11.09 ± 0.03^d	10.53 ± 0.18^de	14.52 ± 0.12^a	10.56 ± 0.69
	30	10.25 ± 0.22^c	12.67 ± 0.58^a	9.54 ± 0.21^d	11.56 ± 0.33^b	9.56 ± 0.22^d	10.46 ± 0.09^c	7.53 ± 0.05^f	10.67 ± 0.34^c	8.20 ± 0.57^e
	60	5.02 ± 0.05^de	7.28 ± 0.10^a	6.92 ± 0.05^b	5.26 ± 0.10^d	5.39 ± 0.26^d	5.41 ± 0.19^d	5.28 ± 0.07^d	6.19 ± 0.36^c	6.31 ± 0.16^c
	90	2.64 ± 0.15^c	2.50 ± 0.06^d	2.65 ± 0.04^c	1.89 ± 0.08^g	1.62 ± 0.03^h	2.84 ± 0.03^b	2.06 ± 0.04^f	3.13 ± 0.05^a	2.42 ± 0.03^e
b*	0	−3.17 ± 0.11^a	−6.39 ± 0.15^c	−4.11 ± 0.12^b	−2.79 ± 0.17^a	−3.29 ± 0.13^a	−4.31 ± 0.29^b	−2.58 ± 0.18^a	−4.48 ± 0.14^b	−3.48 ± 0.13^a
	30	−2.00 ± 0.11^b	−3.29 ± 0.22^e	−2.03 ± 0.11^b	−3.02 ± 0.03^d	−2.43 ± 0.19^c	−2.39 ± 0.16^c	−2.04 ± 0.11^b	−2.16 ± 0.07^b	−1.23 ± 0.08^a
	60	3.99 ± 0.22^c	5.24 ± 0.22^a	4.94 ± 0.12^ab	5.37 ± 0.27^a	3.70 ± 0.14^c	4.77 ± 0.12^b	4.65 ± 036^b	4.85 ± 0.10^b	3.48 ± 0.10^cd
	90	−0.93 ± 0.02^bc	−1.00 ± 0.03^c	−1.62 ± 0.05^e	−1.82 ± 0.02^f	−0.80 ± 0.02^b	−1.22 ± 0.05^d	−0.33 ± 0.03^a	−1.28 ± 0.10^d	−0.86 ± 0.05^b
C*_ab	0	9.82 ± 0.24^f	14.65 ± 0.01^b	10.53 ± 0.13^e	11.90 ± 0.10^c	11.18 ± 0.20^d	11.90 ± 0.13^c	10.84 ± 0.13^de	15.19 ± 0.07^a	11.13 ± 0.62^d
	30	10.44 ± 0.06^c	13.10 ± 0.53^a	9.75 ± 0.18^d	11.95 ± 0.32^b	9.86 ± 0.16^d	10.73 ± 0.07^c	7.80 ± 0.02^e	10.88 ± 0.32^c	8.29 ± 0.55^e
	60	6.41 ± 0.17^f	8.97 ± 0.12^a	8.51 ± 0.11^b	7.52 ± 0.12^d	6.54 ± 0.25^f	7.22 ± 0.08^d	7.04 ± 0.27^de	7.86 ± 0.34^c	7.20 ± 0.18^d
	90	2.80 ± 0.05^c	2.69 ± 0.05^d	3.10 ± 0.05^b	2.62 ± 0.04^d	1.81 ± 0.02^g	3.09 ± 0.05^b	2.08 ± 0.04^f	3.38 ± 0.08^a	2.57 ± 0.03^de
ΔE*_ab	0	0.00 ± 0.00	10.03 ± 0.12^a	2.36 ± 0.03^e	5.68 ± 0.12^c	4.10 ± 0.08^d	5.56 ± 0.10^c	2.11 ± 0.17^ef	9.04 ± 0.19^b	2.63 ± 0.66^e
	30	4.74 ± 0.43^e	10.36 ± 0.25^a	4.75 ± 0.31^e	8.73 ± 0.11^b	8.46 ± 0.17^b	5.47 ± 0.25^d	2.77 ± 0.08^f	7.17 ± 0.38^c	3.01 ± 0.20^f
	60	8.63 ± 0.14^b	9.25 ± 0.27^a	8.55 ± 0.10^bc	9.56 ± 0.25^a	7.91 ± 0.15^d	8.85 ± 0.18^b	8.84 ± 0.29^b	8.90 ± 0.08^b	7.34 ± 0.04^e
	90	12.74 ± 0.01^b	11.28 ± 0.02^f	11.31 ± 0.01^f	11.05 ± 0.05^g	12.33 ± 0.02c	11.41 ± 0.03^e	13.13 ± 0.05^a	11.74 ± 0.03^d	10.38 ± 0.01^h
Different superscript letters indicate significant differences among groups at the same time point (p < 0.05).

Samples	k/d⁻¹		t_1/2/d		R/%
Samples	4 °C	25 °C	4 °C	25 °C	4 °C	25 °C
Control	0.0137 ± 0.0000^a	0.0212 ± 0.0002^b	50.59 ± 0.000^g	32.65 ± 0.321^de	30.60 ± 0.391^e	17.17 ± 0.146^e
Ste2	0.0117 ± 0.0001^e	0.0201 ± 0.0000^d	59.41 ± 0.295^c	34.48 ± 0.000^c	30.89 ± 0.120^e	19.92 ± 0.213^c
Ste4	0.0109 ± 0.0001^f	0.0186 ± 0.0003^e	63.60 ± 0.584^b	37.33 ± 0.508^a	34.80 ± 0.937^c	22.35 ± 0.875^a
Ste8	0.0108 ± 0.0001^f	0.0189 ± 0.0003^e	63.99 ± 0.686^b	36.75 ± 0.592^ab	35.98 ± 0.446^b	21.29 ± 0.388^b
Ste12	0.0119 ± 0.0002^d	0.0208 ± 0.0001^bc	58.09 ± 0.741^d	33.32 ± 0.160^d	30.59 ± 0.874^e	17.43 ± 0.603^e
Ste16	0.0094 ± 0.0001^g	0.0184 ± 0.0003^ef	73.49 ± 0.906^a	37.68 ± 0.538^a	37.63 ± 0.530^a	18.76 ± 0.913^cd
Ste20	0.0125 ± 0.0001^c	0.0243 ± 0.0004^a	55.31 ± 0.512^e	28.57 ± 0.414^f	28.44 ± 1.166^f	14.71 ± 0.837^f
Sucralose	0.0120 ± 0.0001^d	0.0210 ± 0.0004^b	57.61 ± 0.556^d	33.01 ± 0.571^d	32.05 ± 0.586^d	14.78 ± 0.134^f
Sucrose	0.0132 ± 0.0001^b	0.0207 ± 0.0001^c	52.64 ± 0.231^f	33.43 ± 0.093^d	30.32 ± 0.249^e	19.37 ± 0.259^c
Different superscript letters indicate significant differences among groups under the same storage condition (p < 0.05).

Samples	T (K)	k/h⁻¹	t_1/2/h	Eₐ (kJ/mol)	ΔH (kJ/mol)	ΔG (kJ/mol)	ΔS (J/mol·K)
Control	333	0.0628 ± 0.0003^g	11.0375 ± 0.0527^b	33.16 ± 0.162^d	30.39 ± 0.162^d	89.52 ± 0.013^b	−177.57 ± 0.502^d
	343	0.0792 ± 0.0005^g	8.7558 ± 0.0556^a		30.31 ± 0.162^d	91.64 ± 0.018^a	−178.80 ± 0.420^e
	353	0.1141 ± 0.0009^g	6.0769 ± 0.0484^a		30.23 ± 0.162^d	93.32 ± 0.023^b	−178.74 ± 0.479^d
	363	0.1676 ± 0.0008^g	4.1358 ± 0.0202^b		30.14 ± 0.162^d	94.89 ± 0.015^b	−178.36 ± 0.485^d
Ste2	333	0.0819 ± 0.0011^d	8.4608 ± 0.1084^d	19.16 ± 0.296ⁱ	16.39 ± 0.296ⁱ	88.79 ± 0.035^e	−217.42 ± 0.790ⁱ
	343	0.1148 ± 0.0015^e	6.0368 ± 0.0800^c		16.30 ± 0.296ⁱ	90.58 ± 0.038^d	−216.53 ± 0.755^h
	353	0.1311 ± 0.0019^e	5.2879 ± 0.0756^c		16.22 ± 0.296ⁱ	92.91 ± 0.042^c	−217.25 ± 0.738^h
	363	0.1474 ± 0.0005^h	4.7025 ± 0.0166^a		16.14 ± 0.296ⁱ	95.27 ± 0.011^a	−218.01 ± 0.798^h
Ste4	333	0.0767 ± 0.0009^f	9.0379 ± 0.1009^c	35.75 ± 0.160^c	32.98 ± 0.160^c	88.97 ± 0.031^c	−168.15 ± 0.392^c
	343	0.1464 ± 0.0008^c	4.7358 ± 0.0262^e		32.89 ± 0.160^c	89.88 ± 0.016^f	−166.15 ± 0.429^c
	353	0.1712 ± 0.0011^d	4.0497 ± 0.0265^d		32.81 ± 0.160^c	92.13 ± 0.019^d	−168.04 ± 0.484^c
	363	0.2369 ± 0.0022^b	2.9261 ± 0.0272^f		32.73 ± 0.160^c	93.84 ± 0.028^g	−168.36 ± 0.372^c
Ste8	333	0.0789 ± 0.0006^e	8.7892 ± 0.0655^d	38.78 ± 0.265^b	36.01 ± 0.265^b	88.89 ± 0.021^d	−158.81 ± 0.735^b
	343	0.1711 ± 0.0015^a	4.0505 ± 0.0346^f		35.93 ± 0.265^b	89.44 ± 0.024^h	−156.01 ± 0.717^b
	353	0.2011± 0.0013^b	3.4463 ± 0.0222^f		35.84 ± 0.265^b	91.66 ± 0.019^f	−158.11 ± 0.699^b
	363	0.2681 ± 0.0010^a	2.5858 ± 0.0099^g		35.76 ± 0.265^b	93.47 ± 0.012^h	−158.98 ± 0.742^b
Ste12	333	0.0767 ± 0.0025^f	9.0397 ± 0.2938^c	26.81 ± 0.580^g	24.04 ± 0.580^g	88.97 ± 0.091^c	−194.97 ± 1.472^g
	343	0.1669 ± 0.0016^b	4.1525 ± 0.0397^f		23.96 ± 0.580^g	89.51 ± 0.027^g	−191.10 ± 1.634^f
	353	0.1728 ± 0.0038^d	4.0118 ± 0.0881^d		23.88 ± 0.580^g	92.10 ± 0.065^d	−193.27 ± 1.462^f
	363	0.1820 ± 0.0016^e	3.8080 ± 0.0327^c		23.79 ± 0.580^g	94.64 ± 0.026^d	−195.16 ± 1.526^f
Ste16	333	0.0455 ± 0.0006^h	15.2247 ± 0.2035^a	47.31± 0.209^a	44.54 ± 0.209^a	90.41 ± 0.037^a	−137.76 ± 0.547^a
	343	0.0798 ± 0.0013^g	8.6912 ± 0.1413^a		44.46 ± 0.209^a	91.61 ± 0.046^a	−137.49 ± 0.477^a
	353	0.1139 ± 0.0011^g	6.0842 ± 0.0585^a		44.37 ± 0.209^a	93.32 ± 0.028^b	−138.67 ± 0.563^a
	363	0.1942 ± 0.0019^d	3.5695 ± 0.0348^d		44.29 ± 0.209^a	94.44 ± 0.029^e	−138.16 ± 0.519^a
Ste20	333	0.1006 ± 0.0009^a	6.8883 ± 0.0644^f	27.64 ± 0.165^f	24.87 ± 0.165^f	88.22 ± 0.026^h	−190.23 ± 0.423^f
	343	0.1398 ± 0.0010^d	4.9583 ± 0.0338^d		24.79 ± 0.165^f	90.01 ± 0.019^e	−190.16 ± 0.463^f
	353	0.1771 ± 0.0005^c	3.9139 ± 0.0101^e		24.71 ± 0.165^f	92.03 ± 0.008^e	−190.72 ± 0.463^e
	363	0.2325 ± 0.0011^c	2.9817 ± 0.0138^e		24.62 ± 0.165^f	93.90 ± 0.014^f	−190.85 ± 0.420^e
Sucralose	333	0.0844 ± 0.0016^c	8.2114 ± 0.1580^d	23.48 ± 0.654^h	20.71 ± 0.654^h	88.70 ± 0.053^f	−204.19 ± 1.803^h
	343	0.1025 ± 0.0016^f	6.7635 ± 0.1065^b		20.63 ± 0.654^h	90.90 ± 0.045^c	−204.88 ± 1.775^g
	353	0.1216 ± 0.0012^f	5.7006 ± 0.0555^b		20.54 ± 0.654^h	93.13 ± 0.029^a	−205.63 ± 1.771^g
	363	0.1744 ± 0.0001^f	3.9752 ± 0.0026^b		20.46 ± 0.654^h	94.77 ± 0.002^c	−204.70 ± 1.805^g
Sucrose	333	0.0906 ± 0.0009^b	7.6539 ± 0.0719^e	31.57 ± 0.056^e	28.80 ± 0.056^e	88.51 ± 0.026^g	−179.30 ± 0.112^e
	343	0.1724 ± 0.0010^a	4.0214 ± 0.0235^fg		28.72 ± 0.056^e	89.42 ± 0.017^h	−176.96 ± 0.129^d
	353	0.2130 ± 0.0015^a	3.2548 ± 0.0224^g		28.64 ± 0.056^e	91.49 ± 0.020^g	−178.05 ± 0.121^d
	363	0.2382 ± 0.0018^b	2.9105 ± 0.0223^f		28.55 ± 0.056^e	93.83 ± 0.023^g	−179.82 ± 0.114^d
Different superscript letters indicate significant differences among groups under the same temperature (p < 0.05).

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Influence of steviol glycosides on the stability of anthocyanins during storage of blueberry juice

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Influence of steviol glycosides on the stability of anthocyanins during storage of blueberry juice