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Article

The Mechanism of Electrolyzed Functional Water Combined with MA on Postharvest Physiology and Lignification of the Chinese Thorny Bamboo Shoot

by
Yuan Long
†,
Fei Li
†,
Qing Gu
,
Miao He
,
Xinya Xiao
,
Yan Tang
,
Xueying He
,
Shanglian Hu
* and
Bo Zhao
*
Engineering Research Center of Biomass Materials, Ministry of Education, College of Life Sciences and Agri-forestry, Southwest University of Science and Technology, Mianyang 621010, China
*
Authors to whom correspondence should be addressed.
†
These authors contribute equally to this work.
Horticulturae 2026, 12(3), 265; https://doi.org/10.3390/horticulturae12030265
Submission received: 20 January 2026 / Revised: 15 February 2026 / Accepted: 19 February 2026 / Published: 25 February 2026
(This article belongs to the Section Postharvest Biology, Quality, Safety, and Technology)
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Abstract

This research explored the impact of combining electrolyzed functional water (EFW) with modified atmosphere (MA) storage on postharvest ripening, aging, and lignin accumulation in Chinese thorny bamboo shoots. The effects of EFW combined with MA treatment on texture hardness, lignin content, and reactive oxygen metabolism were evaluated. The findings indicated that the EFW + MA treatment was superior in postponing weight reduction, minimizing the increase in shoot hardness and lignin build-up, avoiding epidermal browning, and successfully maintaining elevated activity levels of superoxide dismutase (SOD), catalase (CAT), and ascorbate peroxidase (APX), thus reducing oxidative stress and extending storage time. Moreover, compared with the control group, the EFW/MA treatment markedly reduced malondialdehyde (MDA) levels, thereby improving cellular integrity. In addition, KEGG enrichment analysis revealed that PAL, POD, and PPO, along with their corresponding gene expression levels, were significantly up- and down-regulated. The findings suggest that combining EFW and MA can effectively delay quality deterioration and inhibit lignification in bamboo, thereby preserving its freshness and nutritional value.

1. Introduction

Bamboo shoots are the nascent sprouts of bamboo plants, typically referring to young shoots that have not fully lignified [1]. These shoots are abundant in active constituents, including proteins, carbohydrates, minerals, vitamins, dietary fibers, and antioxidants. They are noted for their high fiber content and low levels of fat and protein [2]. The Chinese thorny bamboo shoot, known for its delicious taste and rich nutritional value, is particularly favored by consumers. They thrive on sunny slopes or along riverbanks and are predominantly found in regions such as Sichuan and Guizhou Province in China. Due to their immature developmental stage and active physiological metabolism, harvested bamboo shoots undergo rapid structural and biochemical changes during storage, including cell wall modification, lignification, and browning. These processes collectively lead to texture hardening and quality deterioration, thereby markedly shortening shelf life. The increase in cellulose and lignin content renders them highly susceptible to water loss and lignification, ultimately reducing their edible value [3]. Peroxidase (POD), phenylalanine ammonia-lyase (PAL), and polyphenol oxidase (PPO) are critical enzymes that influence the browning and lignification of bamboo shoots. Regulating the activity of these enzymes through various methods can effectively delay lignification, thereby extending the storage duration of bamboo shoots [4,5,6,7,8].
Electrolyzed Functional Water (EFW), commonly called electrolyzed water or ionized water, encompasses both acidic and alkaline ionized water. This water is generated by an electrolysis unit, which alters its pH level, oxidation–reduction potential (ORP), and available chlorine concentration (ACC) [9]. Its notable characteristics include a broad sterilization spectrum, the absence of residues, negligible side effects on human health, and low production costs. As a result, EFW has been widely used in recent years, especially for the preservation of fruits and vegetables and the disinfection of food products and equipment [10,11,12,13,14,15,16]. The production equipment for EFW is relatively simple, primarily consisting of an electrolytic cell. EFW is not only effective in maintaining the flavor and freshness of fruits and vegetables, but it also offers benefits like being non-toxic and eco-friendly. This makes it an ideal choice for broader adoption in production practices. Given that postharvest bamboo shoots are prone to microbial growth and oxidative damage, EFW is expected to reduce surface microbial load and alleviate oxidative stress, thereby delaying browning and texture deterioration.
Controlled atmosphere storage technology is currently considered one of the safest and most environmentally friendly methods to preserve fruits and vegetables [17]. It can be categorized into modified atmosphere (MA) storage and controlled atmosphere (CA) storage. Among these, MA relies on the gaps (less than 1 nm) formed by the thermal vibrations of polymer chains in membrane materials, which serve as channels for gas molecules, thus facilitating gas exchange inside and outside the bag. The fruits within the bag create an ideal controlled atmosphere through their own respiration, achieving a notable preservation effect [18]. MA in this study was achieved by packaging bamboo shoots in polyethylene bags, where respiration gradually establishes a low-O2/high-CO2 microenvironment. Modified-atmosphere packaging has been reported to retard browning and lignification in bamboo shoots [19].
Currently, the primary preservation technique for bamboo shoots involves chemical methods, such as synthetic preservatives. This approach protects the bamboo shoots from pathogenic microorganisms and extends their shelf life. Although effective to some extent, this method can leave chemical residues that pose a threat to human health. Therefore, identifying preservation methods that are both safe and effective is critically important. In postharvest fruits and vegetables, EFW is mainly used for surface sanitation due to its high oxidation–reduction potential and available chlorine, which can inactivate microorganisms and reduce decay risk. In addition, EFW treatment has been reported to retard quality deterioration by mitigating oxidative stress and delaying tissue softening/browning, partly through maintaining antioxidant capacity and slowing cell wall degradation [10,11,12,13,14,15,16]. Although its use in preserving bamboo shoots has yet to be documented, this research focuses on thorn bamboo shoots. It integrates two exceptionally safe and efficient preservation techniques: MA and EFW. The impact of using EFW with MA treatment on texture hardness, levels of cellulose and lignin, reactive oxygen metabolism, and enzyme activities related to lignification was evaluated in bamboo shoots. Analysis was conducted on essential enzymes involved in lignin production and the antioxidant system, as well as their gene expressions. The aim is to provide a safe, efficient, and effective method for storing and preserving bamboo shoots while examining in detail the mechanisms that prevent lignification using this approach.

2. Materials and Methods

2.1. Materials and Treatments

On 31 October 2024, in Beichuan Qiang Autonomous County, Sichuan Province, Chinese thorny bamboo shoots were harvested to investigate post-harvest changes in their storage. All shoots were at the same commercial maturity stage and were visually free of mechanical injury; samples were selected to be as uniform as possible in length and diameter. The bamboo shoots were harvested and transported to the laboratory within a 3-h window, then stored at 8 °C for a 4-h pre-cooling period. Preliminary experiments had already determined the appropriate concentration of EFW, soaking duration, and the thickness of polyethylene (PE) preservation bags (See Supplementary Materials). Newly harvested bamboo shoots were divided into four groups (120 kg per treatment; 40 kg per replicate). The first group was control group (CK). The second group (EFW) was soaked in EFW at pH 2.5 for 20 min and then air-dried. The EFW was supplied by Xiongyi Agricultural Technology Service Co., Ltd., Mianyang, China. The electrolyzed water generator was independently developed by the manufacturer. The available chlorine concentration (ACC) of the acidic EFW used in this study was 40 mg/L. The third group (MA) was placed in 45 μm-thick polyethylene preservation bags, each containing approximately 10 bamboo shoot samples. The fourth group (EFW + MA) was soaked in EFW (pH 2.5) for 20 min, air-dried, then placed in 45 μm polyethylene bags with about 10 bamboo shoots per bag. Each treatment consisted of three biological replicates, and each replicate included approximately 40 kg of bamboo shoots. Measurements were conducted independently for each replicate. Each treatment was performed three times at each storage phase, with all samples kept at 4 ± 1 °C for a total of 35 days. Samples were taken on days 0, 7, 14, 21, 28, and 35 to evaluate the bamboo shoots in the preservation bags, then pulverized and stored at −80 °C for transcriptome analysis. Each measurement was performed in triplicate for each biological replicate.

2.2. Weight Loss Rate

The formula used to calculate the rate of weight loss is as follows:
T h e   i n i t i a l   w e i g h t   o f   t h e   b a m b o o   s h o o t s − T h e   w e i g h t   o f   t h e   s t o r e d   b a m b o o   s h o o t s T h e   i n i t i a l   w e i g h t   o f   t h e   b a m b o o   s h o o t s × 100 %

2.3. Decay Rate

For decay assessment, a separate batch of bamboo shoots was randomly selected from each treatment group and used exclusively for decay evaluation. A bamboo shoot was considered decayed when the visible decay area exceeded 20% of the total surface. The decay rate was calculated as the percentage of decayed shoots relative to the total number of shoots assessed in each treatment group, according to the following formula:
Q u a n t i t y   o f   R o t t e n   B a m b o o   S h o o t s T o t a l   q u a n t i t y   o f   b a m b o o   s h o o t s × 100 %

2.4. Surface Color

The color difference in bamboo shoot epidermis was measured using the SR-6 fully automatic colorimeter, Sanmai Technology Co., Ltd., Shenzhen, China. following the methodology established by Moscetti et al. [20]. A whiteboard was used to determine the L, a, and b values at the bamboo shoot’s midpoint. L* indicates lightness, with higher values showing a whiter surface. The a* value balances red and green; a positive a* value means redder, while a negative a* value means greener. The b* value balances yellow and blue; positive b* indicates yellow and negative b* indicates blue.

2.5. Respiratory Rate

The gas analyzer method [21] was employed in this study. Each instance involved weighing 500 g of bamboo shoots, which were subsequently placed in a desiccator at room temperature. A portable CO2 analyzer was used to measure CO2 concentration in the sealed desiccator. The respiratory rate calculation accounted for the bamboo shoots’ mass and duration and was repeated 3 times for accuracy. The method is as follows:
R e s p i r a t o r y   r a t e   ( m g · k g − 1 · h − 1 ) = ( C − C 0 ) × V × 10 − 3 × ρ m × t
In the formula, C represents the reading obtained from the portable CO2 analyzer (mg·L−1), while C0 denotes the initial reading recorded prior to measurement (mg·L−1). The variable V indicates the volume of the desiccator (L), and ρ refers to the gaseous density of CO2, which is 1.997 g·L−1. Additionally, m signifies the weight of the bamboo shoot sampled (kg), and t represents the measurement duration, set at 0.5 h.

2.6. Hardness

Hardness was measured weekly, following modifications to the method described by Li et al. [5], using a digital tester (LD-GY-4). A probe with a 3.5 mm diameter was employed for the procedure. Ten bamboo shoots were randomly picked from each batch, and the probe was gently inserted into the lower-middle area of the shoots. The corresponding values were recorded, with three points randomly selected on each bamboo shoot for three measurements.

2.7. Lignin and Cellulose

The procedure outlined by Wittner et al. [22] was employed to determine the lignin content. Fresh bamboo shoots, finely chopped and weighing 20 g, were dried before being treated with 30 mL of hot distilled water. After cooling, add 75 mL of 86% sulfuric acid and stir at room temperature for 4 to 5 h. Then, add 500 mL of distilled water and bring to a boil. Filter the mixture through a pre-weighed sintered glass funnel, washing with distilled water until no white precipitate forms with 10% BaCl2. The residue was dried and weighed to a constant weight. The lignin content was calculated using the following formula:
L i g n i n   c o n t e n t % = M 2 − M 3 M 1 × 100 %
Here, M1 represents the sample’s weight, M2 denotes the combined weight of the filter residue along with the funnel, and M3 refers to the funnel’s weight.
Cellulose content was measured as described in Qin et al. [23] using a 10 g sample of shredded bamboo shoots in a conical flask. Subsequently, 100 mL of 1.25% H2SO4 solution was added, and the flask was sealed with a glass stopper before being heated to boiling. Timing commenced for 30 min once boiling was achieved, with the heat reduced to prevent foam formation. The mixture was filtered, after which 100 mL of a 1.25% NaOH solution was added to the residue in the conical flask. The solution was then boiled for 30 min. Once heating was halted, vacuum filtration was promptly conducted using a pre-weighed sand filter funnel. After filtration, the residue was rinsed three times with 200 mL hot water under vacuum, dried, and heated in an oven at 105 °C for 3 h. The cellulose content was calculated with the following formula:
C e l l u l o s e   c o n t e n t % = M 2 − M 3 M 1 × 100 %
M1 represents the sample’s weight, M2 indicates the weight of the filter residue alongside the funnel, and M3 denotes the weight of just the funnel.

2.8. Determination of PAL, POD, and PPO Activity

The method for measuring phenylalanine ammonia-lyase (PAL) activity was adapted from Kahramanoğlu et al. [24], with minor modifications. Bamboo shoots weighing 5 g were homogenized on ice with 5.0 mL of 0.1 mol/L borate-borax buffer at pH 8.8. The resulting homogenate was centrifuged at 4 °C at 10,000× g for 30 min. The enzyme extract, as the supernatant, was then kept chilled. A 3.0 mL of 50 mmol/L borate buffer (pH 8.8) was mixed with 0.5 mL of 20 mmol/L phenylalanine, and incubated at 37 °C for 60 min. Absorbance was recorded at 290 nm using distilled water as the blank. Assays were done in triplicate. A 0.01 absorbance change per hour per gram of bamboo corresponds to 1 enzyme unit.
Peroxidase (POD) activity was assessed using a modified protocol from Liu et al. [25]. A 5 g portion of bamboo shoot was weighed and placed into a pre-chilled mortar. Following this, 5.0 mL of a 0.1 mol/L sodium acetate–acetic acid extraction buffer at pH 5.5 was added, and the mixture was homogenized while kept on ice. The homogenate was then centrifuged at 4 °C and 10,000× g for 30 min, after which the supernatant was collected as the enzyme extract and stored at a low temperature. The reaction began with 3.0 mL of 0.25 mmol/L guaiacol and 0.5 mL of enzyme extract, followed immediately by 200 µL of 0.01 mol/L H2O2. Distilled water was the reference. Absorbance was measured at 470 nm and repeated three times. One unit of activity is defined by an absorbance rate change of 1 per minute per gram of bamboo shoot.
The PPO activity was evaluated following a revised procedure based on Yang et al. [26], using the same enzyme extraction method as for POD. The reaction mixture comprised 4.0 mL of 100 mmol/L acetate–sodium acetate buffer at pH 5.5, 1.0 mL of a 50 mmol/L catechol solution, and 100 µL of enzyme extract. Absorbance at 420 nm was recorded three times against distilled water immediately after combining the components. One unit of activity was defined as a change in absorbance of 1 per minute per gram of bamboo shoot.

2.9. MDA Content

The concentration of malondialdehyde (MDA) was determined using the TBA method as described by Shi et al. [27]. A sample of bamboo shoots weighing 1.0 g was mixed with 5.0 mL of a 100 g/L TCA solution. This mixture was thoroughly homogenized and then centrifuged at 4 °C and 10,000× g for 20 min. The resulting supernatant was collected and kept at a low temperature for later use. A 2.0 mL portion of the supernatant (in the control, 2.0 mL of a 100 g/L TCA solution was used instead) was mixed with 2.0 mL of 0.67% TBA. The mixture was boiled for 20 min, cooled, and centrifuged once more. Absorbance readings were taken at 450 nm, 532 nm, and 600 nm.

2.10. Measurement of CAT, SOD, and APX Activities

Enzyme extracts for CAT, SOD, and APX were prepared by homogenizing 5 g of bamboo shoot tissue in 5 mL of pre-chilled phosphate buffer (pH 7.8), followed by centrifugation at 10,000× g for 30 min at 4 °C. The supernatant was collected as the crude enzyme extract. The activity of Catalase (CAT) was determined according to the procedure outlined by Liu et al. [28]. The following reagents were combined: 0.2 mL of CAT extract, 1.5 mL of phosphate buffer (pH 7.8), and 1.0 mL of distilled water. Tubes were pre-warmed to 25 °C, after which 0.3 mL of 0.1 mol/L H2O2 was added; the absorbance was measured at 240 nm.
The determination of superoxide dismutase (SOD) activity was performed according to the method described by Qi et al. [29]. The activity of SOD (U) is quantified by the enzyme quantity required to curtail 50% of nitroblue tetrazolium’s photochemical reduction. A protected tube served as the baseline control, and absorbance was measured at 560 nm.
The activity of ascorbate peroxidase (APX) was evaluated according to the procedure outlined by Petriccione et al. [30]. In a test tube, 0.1 mL of the enzyme extract was mixed with 2.6 mL of reaction buffer. Subsequently, 0.3 mL of 2 mmol/L H2O2 was added to trigger the reaction. The solution was thoroughly mixed, and timing commenced. Absorbance at 290 nm was measured 15 s after the start, with subsequent measurements taken at 30-s intervals.

2.11. Transcriptional Level and Differential Expression Gene Analysis

RNA was extracted from bamboo shoots using TRIzol, (Shenggong Bioengineering Co., Ltd., Shanghai, China) and libraries were prepared with the KCTM Digital mRNA Library Prep Kit to correct PCR and sequencing errors by adding unique molecular identifiers (UIDs) to cDNA. The library preparation process involved enriching PCR products with fragment sizes ranging from 200 to 500 bp. Using the Phyllostachys edulis genome assembly as the reference genome for transcriptome sequencing, the enriched libraries were quantified and sequenced on the DNBSEQ-T7 (MGI) platform in PE150 mode. To ensure the reliability of the transcriptomic data, relevant genes were selected from the DEGs for further analysis.

2.12. Gene Expression Analysis

Transcriptome sequencing was performed to analyze gene expression changes among treatments. Clean reads were mapped to the reference genome of Phyllostachys edulis, and gene expression levels were normalized using the FPKM (Fragments Per Kilobase of transcript per Million mapped reads) method. Differentially expressed genes (DEGs) were identified based on a fold change ≥ 2 and a false discovery rate (FDR) < 0.05. Functional annotation and pathway enrichment analyses were conducted using GO and KEGG databases. The calculation method for FPKM is as follows:
F   P   K   M = t o t a l   e x o n   f r a g m e n t s m a p p e d   f r a g m e n t s   ( m i l l i o n s ) × e x o n   l e n g t h   ( K B )

2.13. Statistical Analysis

Experimental data were analyzed using SPSS 26.0 to determine variance and perform Duncan’s test across three replicates, with a significance level of 0.05. Results are presented as mean ± standard deviation, and figures were created using Origin 2022.

3. Results and Discussion

3.1. Changes in Physical Properties of EFW- and MA-Treated Bamboo Shoots During Storage

Color is vital for evaluating the freshness of agricultural products [31]. During storage, Chinese thorny bamboo shoots undergo a color change from green to yellow to brown. As shown in Figure 1A, visual deterioration and browning were most severe in the CK group during storage. Figure 1B–D shows color changes over storage periods with different treatments. The L* value, which indicates brightness, decreases over prolonged storage, leading to browning and a loss of shine. The control group (CK) displayed the most color change. Badwaik’s research highlights notable color changes in bamboo shoots caused by enzymatic browning and microbial deterioration [32]. As storage progressed, L* and b* values decreased, while a* values increased. Chlorophyll in the shoots’ skin degrades enzymatically, forming chlorophyll quinone [33]. Shoots treated with MA and EFW + MA showed less color change than CK (p ≤ 0.05), with EFW + MA being the most effective in preventing browning. This trend may be attributed to the combined effects of surface sanitation by EFW and O2 limitation by MA, which together slow oxidative reactions and metabolism-associated quality loss during storage.
Weight reduction in fresh produce is primarily due to moisture evaporation and respiration, which reduces quality and shelf life [34]. Throughout the 35 days, every group experienced an increase in the weight loss of the bamboo shoots. The MA and EFW + MA groups experienced lower weight loss compared to the CK and EFW groups, with the EFW + MA group being the most effective. MA and EFW + MA treatments resulted in the least weight loss, significantly lower than CK (11.75% and 5.52%, respectively).
Figure 1F illustrates that the decomposition rate of bamboo shoots increases as storage time lengthens. At the conclusion of the storage period, the CK group experiences a decay rate of 100%, while the EFW + CK group’s decay rate is merely 22.22%. This indicates that the EFW + MA treatment is the most effective, delaying the onset of decay and maintaining a lower decay rate.

3.2. Changes in Texture and Respiratory Rate of the Chinese Thorny Bamboo Shoots During Storage

Hardness serves as a key metric for assessing the storage quality of fruits and vegetables [6]. Initially, as depicted in Figure 2A, the four bamboo shoot groups did not exhibit any notable differences in hardness. Over the storage duration, hardness in these shoots steadily increased due to lignification, a process closely linked with higher lignin and cellulose levels, consistent with Liu et al.’s findings [35]. By the end of storage, bamboo shoots treated with MA and EFW + MA were significantly less hard than the CK group, as EFW and MA treatments effectively slowed lignification, reducing the increase in hardness. Notably, the EFW + MA treatment showed the least change in hardness over the storage period. The combination of EFW with preservation bags significantly delayed the increase in hardness, prolonging the hardening and senescence of bamboo shoots.
Lignin is essential for the secondary cell wall, and its accumulation increases the hardness of post-harvest fruits and vegetables [36,37,38]. Figure 2B shows lignin levels in bamboo shoots rise over storage time. The control (CK) group showed a greater increase in lignin than the treated groups, as observed by Yang et al. [39]. The EFW + MA treatment notably reduced lignin content, with the most significant effect on delaying lignin accumulation. Since lignin biosynthesis is a major branch of secondary metabolism (phenylpropanoid pathway), the reduced lignin accumulation under EFW + MA may reflect weakened induction of secondary metabolism associated with oxidative stress during storage.
Cellulose serves as a crucial marker for assessing the aging process in bamboo shoots. During storage, increases in cellulose levels and thickening of the secondary cell wall significantly enhance hardness, leading to a rougher texture and a decline in quality [40]. In Figure 2C, cellulose levels in bamboo shoots increase with storage, aligning with previous findings [41]. The CK group shows a faster rise in cellulose than the others. From day 14 onward, cellulose in EFW + MA is lower than in CK, suggesting that EFW + MA slows postharvest cellulose synthesis. This supports Li et al.’s finding that melatonin delays the aging of bamboo shoots [5]. Polyethylene bags with low oxygen and EFW reduce enzyme activity, thereby decreasing lignin and cellulose synthesis, slowing the degradation of quality.
After being harvested, fruits and vegetables continue to respire aerobically, using up their own nutrients, which reduces their quality. Therefore, reducing respiration is crucial for quality control and for extending the post-harvest storage life of these products [42]. Figure 2D shows that the respiration of bamboo shoots under various treatments decreases with storage time, peaks, and then declines. The control (CK) group had a significantly higher respiratory rate than the others. After 35 days, the EFW, MA, and EFW + MA groups had rates of 94.88, 80.70, and 65.07 mg·kg−1·h−1, which were 16.57%, 29.04%, and 42.79% lower, respectively, than those of the CK group. The EFW + MA group exhibited a significantly lower respiratory rate than the other three groups. The EFW + MA method successfully regulated breathing by stabilizing oxygen and carbon dioxide levels. Furthermore, this approach markedly slowed the decomposition of bamboo shoots and minimized the onset of wound respiration [19].

3.3. Changes in Enzymes Related to Lignin Biosynthesis During Storage

POD is crucial in lignin biosynthesis by decomposing hydrogen peroxide (H2O2) and aiding lignin monomer polymerization in the final synthesis stage [43]. Figure 3A illustrates that POD activity across all groups gradually increased from day 0 to day 35. The trend is attributed to damage to the bamboo shoot cell membrane during storage, which increases oxidative stress and POD activity, with H2O2 serving as a key product of oxidation [44]. The rise in POD activity aligns with Liu et al. [45]. POD activity was consistently lower in the treated groups than in the control group, with EFW and MA treatments inhibiting POD activity, particularly in the EFW + MA group.
PAL is the rate-limiting enzyme in bamboo shoot lignification, converting phenylalanine to cinnamic acid, which then forms lignin precursors like coniferyl, sinapyl, and coumaryl alcohols. These aromatic alcohols undergo dehydrogenation polymerization facilitated by POD, leading to lignin formation, which plays a role in the aging process of bamboo shoots [46]. Figure 3B illustrates that during the storage period, the PAL activity in the treated group consistently remained lower than the levels found in the control group. At the final stage of storage, the PAL activity in the EFW + MA group was significantly lower than that in the other groups (p ≤ 0.05).
Throughout storage, PPO activity in treatment groups remained consistently lower than in the control group, with all bamboo shoot groups showing a similar increase in PPO activity over time. By the end of storage, PPO activity in the EFW, MA, and EFW + MA groups was significantly lower than in the CK group, at 93.40%, 86.94%, and 73.79% of the CK group’s activity, respectively. Previous studies have demonstrated that PPO activity tends to increase when the tissues of fruits or vegetables are damaged during harvesting or processing, or when storage conditions are suboptimal [47]. Similar trends have been reported under low-temperature storage conditions with modified atmosphere packaging (e.g., 4 °C storage in polyethylene bags) [19]. The EFW + MA treatment markedly reduced the activities of PAL, POD, and PPO.
This study further indicated that the activities of the enzymes PAL, POD, and PPO, as well as lignin content, showed consistent variation under different treatment conditions. The findings suggest that each treatment approach effectively mitigates lignin buildup by inhibiting the activities of PAL, POD, and PPO enzymes in the samples. As detailed by Li et al., minimizing these enzyme activities results in reduced lignin deposition or lignification in moso bamboo shoots [5]. Therefore, EFW treatment, MA treatment, and the combined EFW + MA treatment effectively inhibit lignin-associated enzymes, thereby decelerating lignification in bamboo shoots, with EFW + MA being the most effective.

3.4. Changes in Antioxidant System-Related Enzymes During Storage

Aging in plant tissues is typically associated with elevated levels of MDA, increased production of reactive oxygen species (ROS), and reduced activity of enzymes involved in free radical scavenging [48]. Enzymes with antioxidant properties found in postharvest fruits and vegetables have the potential to diminish reactive oxygen species (ROS) and delay aging throughout storage, according to Cheng et al. [49]. The enzymes catalase (CAT), superoxide dismutase (SOD), and ascorbate peroxidase (APX) play significant roles in the antioxidant defense system. They work to prevent the accumulation of hydrogen peroxide (H2O2), minimize lipid peroxidation in membranes, lower MDA concentrations, and inhibit lignin and cellulose synthesis. Xu et al. reported that enhanced ROS-scavenging capacity was associated with delayed senescence in bamboo shoots during cold storage [50]. Under our storage condition (4 ± 1 °C), EFW + MA maintained higher antioxidant enzyme activities and reduced MDA accumulation (Figure 4), consistent with this observation.
MDA is a byproduct of membrane lipid peroxidation, commonly occurring in fruits and vegetables affected by diseases, chilling injuries, or stress, leading to plant cell damage or death [51]. As shown in Figure 4A, bamboo shoots exhibited a rising trend in MDA content over extended storage durations across all groups. At the conclusion of the storage period, MDA levels declined by 13.42%, 18.76%, and 33.10% in the EFW, MA, and EFW + MA groups, respectively, relative to the control (CK) group. Notably, the EFW + MA treatment proved the most effective at curbing MDA accumulation in bamboo shoots, thereby reducing membrane lipid peroxidation and preserving membrane integrity. Menaka et al. [52] examined the postharvest quality of guava when treated with melatonin. They found this approach effectively reduced MDA in the fruit, protected the cell membrane, and significantly prevented peel browning. EFW and MA treatments also contributed to slowing MDA accumulation, thereby prolonging the shelf life of bamboo shoots.
CAT is an essential antioxidant enzyme in plants that catalyzes the decomposition of H2O2 into H2O and O2. This process is vital for removing reactive oxygen species and reducing oxidative harm [53]. Figure 4B demonstrates that, during low-temperature storage, CAT activity in bamboo shoots across four groups initially rose, then fell, and eventually showed a gradual rise. CAT activity peaked at 7 days of storage in the control group (CK), the enzyme-free water group (EFW), the malic acid group (MA), and the EFW + MA group. Notably, CAT activity in the treated bamboo shoots exceeded that of the control group, highlighting its protective role during storage. From a physiological perspective, MA reduces O2 availability and suppresses respiratory metabolism, thereby limiting excessive ROS production during storage. Meanwhile, EFW treatment may alleviate initial oxidative and microbial stress at the tissue surface. The combined effect likely stabilizes cellular redox homeostasis and membrane integrity, which explains the relatively higher antioxidant enzyme activities and lower MDA accumulation observed under EFW + MA treatment.
SOD can convert superoxide anions (O2−) into H2O2 and O2, thereby neutralizing reactive oxygen species in plants. Increased SOD activity can reduce the damage caused by reactive oxygen species in fruits and vegetables [49]. According to Figure 4C, during low-temperature storage, SOD activity in four groups of bamboo shoots initially rose and then eventually fell. On day 14 of storage, SOD activity reached its peak in the control group (CK), EFW group, MA group, and EFW + MA group, with levels of 88.70 U·g−1 FW h−1, 112.14 U·g−1 FW h−1, 121.54 U·g−1 FW h−1, and 139.56 U·g−1 FW h−1, respectively. SOD activity in the treatment groups was significantly higher than in the control group, effectively reducing reactive oxygen species damage and extending the storage life of bamboo shoots.
APX utilizes ascorbic acid to convert H2O2 into H2O, playing a crucial role in the plant’s antioxidant defense mechanism. The enhanced activity of the APX enzyme indicates an improved ability of plants to neutralize reactive oxygen species, inhibit lipid peroxidation in fruits, and mitigate stress-induced damage [54]. Thus, higher APX activity benefits fruit and vegetable quality and lengthens their storage life [55]. Figure 4D shows that the APX activity pattern in the four treated bamboo shoots mirrored that of SOD activity, initially increasing and then decreasing. APX activity reached its maximum at 7 days of storage. During the middle and late stages of storage, the EFW + MA group showed notably higher APX activity than the other groups.
Bamboo shoots enhance antioxidant enzyme activity, improving their ability to neutralize reactive oxygen species, prevent lipid peroxidation in fruits, and reduce stress from unfavorable conditions. Key enzymatic antioxidants, like CAT, SOD, and APX, mitigate oxidative damage [56]. In this study, the control (CK) group of bamboo shoots showed lower CAT, SOD, and APX activities than the three treated groups. These treatments preserved enzyme activity, reduced oxidative stress, and prolonged storage life, with the EFW + MA group showing the most significant effect.

3.5. Analysis of Differentially Expressed Genes at Different Storage Stages

To elucidate the molecular mechanisms underlying lignin accumulation in bamboo shoots during storage, we employed RNA sequencing to analyze differentially expressed genes between 0 d and 14 d of storage. A substantial number of genes were up-regulated and down-regulated in both the control and various treatment groups (Figure 5A–F), indicating significant physiological and metabolic changes within the bamboo shoots during storage. At 14 d of storage, linoleic acid metabolism, glutathione metabolism, and flavonoid biosynthesis were significantly enriched and down-regulated in CK14 vs. EM14 (EM: EFW + MA treatment group), suggesting that EFW combined with MA treatment effectively maintained the stability of cell membranes in bamboo shoots, delayed senescence and oxidative browning, and inhibited microbial growth during storage. Under the combined treatment of EFW and MA, lignin biosynthesis in bamboo shoots was significantly affected. The markedly enriched pathways included plant-pathogen interaction, phenylpropanoid biosynthesis, tryptophan metabolism, secondary metabolite biosynthesis, amino acid and nucleotide metabolism, plant hormone signal transduction, flavonoid biosynthesis, linoleic acid metabolism, and glutathione metabolism. These findings are consistent with previous research results [5].

3.6. Relative Expression Levels of Lignin Synthesis-Related Genes at Different Storage Stages

Lignin levels in harvested bamboo shoots increase over time, regulated by enzymes like POD, PAL, C3H, and 4CL, along with their gene expressions. The activities of these enzymes and their metabolite levels often vary, typically paralleled by changes in the related gene expression [57,58]. Previous studies have demonstrated that enzymes such as PAL, POD, CAD, 4CL, and C4H, along with their corresponding gene expressions, play a crucial role in regulating lignin synthesis and accumulation [59]. This study’s results indicate that genes such as PedPAL, PedPOD, PedCAD, Ped4CL, PedC3H, PedC4H, PedLAC, PedCCOAOMT, and PedCOMT are likely the main contributors to postharvest lignification in bamboo shoots during storage, especially when treated with MA in conjunction with EFW. In particular, suppressing these genes significantly reduces lignin accumulation and slows lignification.
On the 14th day of storage, the detection data indicated a notable downregulation of gene expression across all bamboo shoot treatment groups compared to the control group (CK) (refer to Figure 6). The gene expression levels of PedPAL1, PedPAL2, PedPOD1, and PedPOD2 in the treatment groups on day 14 matched those at day 0. However, the expression levels of PedCAD1, PedCAD2, Ped4CL2, PedC3H, and PedC4H decreased to below 50% of their initial levels, indicating that EFW with MA significantly inhibited their expression. On the other hand, the relative expression levels of PedPAL3, PedLAC1, PedCCOAOMT, and PedCOMT were elevated on day 14 compared to day 0, likely due to the aging process of the bamboo shoots. Additionally, the activities of PAL and POD enzymes, along with their corresponding gene expression levels and lignin content, showed a positive correlation, suggesting that increased gene expression likely affects enzyme activity during lignification.

3.7. Effect of EFW + MA Treatment on the Lignin Biosynthesis Pathway in Chinese Thorny Bamboo Shoots

This study further investigates the lignin biosynthesis pathway in the Chinese thorny bamboo shoots using transcriptome-related data. The key enzymes involved in lignin biosynthesis are closely associated with lignin synthesis, with the initial three steps of this metabolic pathway typically catalyzed by PAL, C4H, and 4CL [60]. Under the action of these enzymes, phenylalanine is deaminated to form trans-cinnamic acid, which is subsequently hydroxylated into trans-4-coumaric acid. As illustrated in Figure 7, within the lignin biosynthesis metabolic pathway, eight key enzyme genes related to lignin biosynthesis were downregulated in EFW + MA-treated bamboo shoots. EFW + MA treatment downregulated the expression of PedPALs, Ped4CLs, PedC3Hs, PedCCoAOMTs, PedCOMTs, PedCADs, PedPODs, and PedLACs, thereby inhibiting the formation of lignin intermediates and delaying lignification of bamboo shoots, thereby extending their shelf life. The stronger effect of EFW + MA compared with either EFW or MA alone may reflect complementary actions: EFW primarily reduces the initial microbial/oxidative burden at the tissue surface, whereas MA suppresses respiration by limiting O2 availability. When applied separately, only one component of the deterioration process is constrained; when combined, both stress initiation and downstream oxidative/lignification-related metabolism are simultaneously moderated, resulting in a more evident preservation outcome.

4. Conclusions

This study evaluated combining EFW with MA storage to inhibit postharvest maturation and lignification of Chinese thorny bamboo shoots. The treatment that combined EFW with MA effectively inhibited weight loss in thorn bamboo shoots, significantly suppressed the increase in shoot hardness, reduced the accumulation of lignin and cellulose, decreased the decay rate, and inhibited the lignification process by suppressing the activities of PAL, POD, and PPO, thereby maintaining the freshness of the shoots. In addition, the treatment combining electrolyzed functional water with MA reduced MDA levels more effectively than the control, helping to maintain cell structure, prevent shoot epidermis browning, and sustain higher enzymatic activities of CAT, SOD, and APX. This approach minimized oxidative stress and retained the preferred taste of the shoots. EFW and MA treatment reduce lignin synthesis in bamboo shoots during low-temperature storage by lowering enzyme activity and gene expression related to lignin production, including PedPAL, PedPOD, PedCAD, Ped4CL, and PedC3H. This treatment effectively slows the degradation of quality and prevents lignification, preserving freshness and nutritional value.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/horticulturae12030265/s1, Figure S1. Preliminary screening for optimal MA bag thickness, optimal EFW pH, and treatment time.

Author Contributions

Conceptualization and writing—original draft preparation, Y.L. and F.L.; writing—review and editing, B.Z.; supervision, project administration, and funding acquisition, B.Z. and S.H. methodology, Q.G. and M.H.; software, X.X. and Y.T.; validation, X.H. All authors have read and agreed to the published version of the manuscript.

Funding

This work was financially supported by the National Innovation and Entrepreneurship Training Program for Undergraduate (S202410619035), National Key R&D Program of China (2021YFD2200504), the University of Science and Technology of China-Southwest University of Science and Technology Counterpart Cooperation and Development Joint Fund (Grant NO. 24ZXLHJJ03).

Data Availability Statement

The original contributions presented in this study are included in the article/Supplementary Material. Further inquiries can be directed to the corresponding authors.

Acknowledgments

We would like to express our sincere gratitude to all those who have contributed to the success of this research.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Effects of different treatments on the external quality of the Chinese thorny bamboo shoot. (A) Visual changes in bamboo shoots during storage under different treatments. Changes in (B) brightness, (C) redness, (D) yellowness, (E) weight loss rate, and (F) decay rate of bamboo shoots are presented. Data are expressed as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
Figure 1. Effects of different treatments on the external quality of the Chinese thorny bamboo shoot. (A) Visual changes in bamboo shoots during storage under different treatments. Changes in (B) brightness, (C) redness, (D) yellowness, (E) weight loss rate, and (F) decay rate of bamboo shoots are presented. Data are expressed as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
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Figure 2. Effects of EFW and MA treatments on texture and respiratory rate of the Chinese thorny bamboo shoots. Changes in (A) hardness, (B) lignin content, (C) cellulose content, and (D) respiratory intensity. Data are presented as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
Figure 2. Effects of EFW and MA treatments on texture and respiratory rate of the Chinese thorny bamboo shoots. Changes in (A) hardness, (B) lignin content, (C) cellulose content, and (D) respiratory intensity. Data are presented as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
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Figure 3. Changes in enzymes involved in lignin biosynthesis during the storage. Changes in (A) POD, (B) PAL, and (C) PPO activities in the Chinese thorny bamboo shoots. Data are presented as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
Figure 3. Changes in enzymes involved in lignin biosynthesis during the storage. Changes in (A) POD, (B) PAL, and (C) PPO activities in the Chinese thorny bamboo shoots. Data are presented as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
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Figure 4. Changes in antioxidant system-related enzymes during the storage of the Chinese thorny bamboo shoots. Changes in (A) MDA content, (B) CAT, (C) SOD, and (D) APX activities. Data are expressed as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
Figure 4. Changes in antioxidant system-related enzymes during the storage of the Chinese thorny bamboo shoots. Changes in (A) MDA content, (B) CAT, (C) SOD, and (D) APX activities. Data are expressed as mean ± standard deviation (SD). Different letters indicate significant differences among treatments at the same storage time (p ≤ 0.05).
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Figure 5. KEGG enrichment analysis revealed DEGs in the Chinese thorny bamboo shoots stored for 0 and 14 days. In the figure, the CK group represents the control group, with CK0 and CK14 indicating storage days 0 and 14, respectively; the EM group represents the treatment group, which underwent combined EFW and MA treatment, with EM0 and EM14 indicating storage days 0 and 14, respectively.
Figure 5. KEGG enrichment analysis revealed DEGs in the Chinese thorny bamboo shoots stored for 0 and 14 days. In the figure, the CK group represents the control group, with CK0 and CK14 indicating storage days 0 and 14, respectively; the EM group represents the treatment group, which underwent combined EFW and MA treatment, with EM0 and EM14 indicating storage days 0 and 14, respectively.
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Figure 6. Relative expression of lignin synthesis-related genes in the Chinese thorny bamboo shoots treated with EFW and MA during storage. Data are presented as mean ± standard deviation (SD). * in the figure indicate significant differences between treatments (p ≤ 0.05).
Figure 6. Relative expression of lignin synthesis-related genes in the Chinese thorny bamboo shoots treated with EFW and MA during storage. Data are presented as mean ± standard deviation (SD). * in the figure indicate significant differences between treatments (p ≤ 0.05).
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Figure 7. Effect of EFW + MA treatment on lignin biosynthesis in the Chinese thorny bamboo shoots.
Figure 7. Effect of EFW + MA treatment on lignin biosynthesis in the Chinese thorny bamboo shoots.
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MDPI and ACS Style

Long, Y.; Li, F.; Gu, Q.; He, M.; Xiao, X.; Tang, Y.; He, X.; Hu, S.; Zhao, B. The Mechanism of Electrolyzed Functional Water Combined with MA on Postharvest Physiology and Lignification of the Chinese Thorny Bamboo Shoot. Horticulturae 2026, 12, 265. https://doi.org/10.3390/horticulturae12030265

AMA Style

Long Y, Li F, Gu Q, He M, Xiao X, Tang Y, He X, Hu S, Zhao B. The Mechanism of Electrolyzed Functional Water Combined with MA on Postharvest Physiology and Lignification of the Chinese Thorny Bamboo Shoot. Horticulturae. 2026; 12(3):265. https://doi.org/10.3390/horticulturae12030265

Chicago/Turabian Style

Long, Yuan, Fei Li, Qing Gu, Miao He, Xinya Xiao, Yan Tang, Xueying He, Shanglian Hu, and Bo Zhao. 2026. "The Mechanism of Electrolyzed Functional Water Combined with MA on Postharvest Physiology and Lignification of the Chinese Thorny Bamboo Shoot" Horticulturae 12, no. 3: 265. https://doi.org/10.3390/horticulturae12030265

APA Style

Long, Y., Li, F., Gu, Q., He, M., Xiao, X., Tang, Y., He, X., Hu, S., & Zhao, B. (2026). The Mechanism of Electrolyzed Functional Water Combined with MA on Postharvest Physiology and Lignification of the Chinese Thorny Bamboo Shoot. Horticulturae, 12(3), 265. https://doi.org/10.3390/horticulturae12030265

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