Next Article in Journal
Resistance Phenotyping and WGCNA Identify Oxidative-Defense Hub Regulators in Strawberry Challenged by Colletotrichum siamense
Previous Article in Journal
Optimizing Nursery Production of Apple Trees: Assessing the Dose Response to Water and Fertilizer in Two Cultivars
 
 
Search for Articles:
Title / Keyword
Author / Affiliation / Email
Journal
Article Type
 
 
Advanced Search
 
Section
Special Issue
Volume
Issue
Number
Page
 
Journals
Horticulturae
Volume 11
Issue 12
10.3390/horticulturae11121426
horticulturae-logo
Submit to this Journal Review for this Journal Propose a Special Issue
► Article Menu

Article Menu

share Share announcement Help format_quote Cite question_answer Discuss in SciProfiles
first_page
settings
Order Article Reprints
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:
Background:
Article

Genome-Wide Identification and Characterization of Tomato Acyl-CoA Oxidase Family Genes ACX

by
Chunlei Wang
*,
Zesheng Liu
,
Yanlong Gao
,
Qianbing Li
,
Qi Wang
and
Caiting An
College of Horticulture, Gansu Agricultural University, 1 Yinmen Village, Anning District, Lanzhou 730070, China
*
Author to whom correspondence should be addressed.
Horticulturae 2025, 11(12), 1426; https://doi.org/10.3390/horticulturae11121426
Submission received: 23 October 2025 / Revised: 13 November 2025 / Accepted: 22 November 2025 / Published: 25 November 2025
(This article belongs to the Section Biotic and Abiotic Stress)
Browse Figures
Review Reports Versions Notes

Abstract

This study conducted a systematic identification and functional analysis of the SlACX gene family in Solanum lycopersicum. Through genome-wide screening, a total of six SlACX members were identified, and their encoded proteins showed significant differences in physicochemical properties, suggesting potential functional differentiation. Analysis of gene structure and conserved motifs revealed that SlACXs were highly conserved in evolution, but the cis-acting elements in the promoter region were rich and diverse, suggesting that they may integrate multiple signaling pathways. Chromosomal localization and collinearity analysis revealed that gene replication events were the main driving force for family expansion, and there were key interspecific collinearity blocks with Arabidopsis thaliana and Glycine max. Expression analysis showed that SlACXs exhibited remarkable tissue specificity and strong temporal dynamic response patterns to UV, dark, ABA, MeJA, and various abiotic stresses (cold, heat, H2O2, PEG, and NaCl). Several genes (such as SlACX1, SlACX3, SlACX4, and SlACX5) exhibited consistently high expression levels under various stress conditions, underscoring their potential role as central regulatory hubs.

1. Introduction

Tomato (Solanum lycopersicum), one of the most important crops in the world, is also one of the best model plant species for plant research [1]. With the gradual deterioration of the global ecosystem and the increasing demand for high-quality tomatoes, research on tomato resistance and quality has progressively advanced in depth and sophistication.
Acyl-CoA oxidase (ACyl-CoA oxidase, ACX), also known as acyl-CoA oxidase, mainly participates in fatty acid metabolism, the synthesis of various unsaturated fatty acids, and signal transduction [2]. Research indicates that there are various ACX isoenzymes in plants, each with different sizes and subunit compositions, and they possess substrate (short-chain, medium-chain, and long-chain fatty acids) specificity, specifically long-chain ACX (LACX), medium-chain ACX (MACX), and short-chain ACX (SACX) [3]. Research has demonstrated that hydrogen peroxide (H2O2), a terminal product of the β-oxidation pathway mediated by the ACX gene, can induce oxidative reactions and subsequently activate signaling molecules associated with plant stress responses [4]. In addition, the function of β-oxidation not only includes the degradation of fatty acids but also generates various hormone signaling molecules such as IAA (auxin), JA (jasmonic acid), and SA (salicylic acid), thereby activating the hormone metabolic process within plants, inducing the expression of related genes, and directly participating in the response to environmental stress [5]. It was found that the GhACX gene was significantly induced by high temperature, low temperature, salt, and drought stress in Gossypium hirsutum L. [6]. Furthermore, AtACXs can catalyze the degradation of fatty acids. Among them, the AtACX1 and AtACX5 genes are involved in the synthesis of jasmonic acid (JA) to participate in other developmental, pollen fertility, and stress response processes [7].
Although the identification of the ACX gene family has been carried out in model plants such as Arabidopsis thaliana [8], no related reports have been seen in tomatoes. Species such as tomatoes and grapes all belong to the rose group in the middle of the true dicotyledonous plants in evolution, and there is a high degree of genomic collinearity conservation among them. Meanwhile, the functions of the ACX gene family of these species have been well annotated. Therefore, choosing these representative species for cross-phylogenetic scale comparative analysis helps to more accurately infer the evolutionary relationship, functional differentiation process, and conservation characteristics of the SlACX gene in tomatoes in an evolutionary context, thereby providing a theoretical framework for subsequent functional research. This supplement makes the choice of comparative analysis more scientifically logical. This study identified the ACX family genes through the whole genome of tomato and analyzed their physicochemical properties, chromosomal localization, gene structure, gene evolutionary relationships, and cis-acting elements, as well as the expression patterns of ACX genes under different treatments, in order to clarify the function of ACX genes under abiotic stress and provide a basis for subsequent related research and identification.

2. Materials and Methods

2.1. The Genome-Wide Identification of ACX Gene Family Members in Solanum lycopersicum

Genomic reference data for S. lycopersicum were obtained from https://solgenomics.net/ (accessed on 14 August 2025), genome data of soybean were downloaded from https://plants.ensembl.org/index.html (accessed on 14 August 2025), genome data of Oryza sativa were downloaded from http://rice.plantbiology.msu.edu/ (accessed on 17 August 2025), and genome data of Arabidopsis thaliana were downloaded from https://www.arabidopsis.org (accessed on 19 August 2025). Genome data of Zea mays L. were downloaded from https://www.maizegdb.org (accessed on 20 August 2025). The ACX family sequences (PF01756) were downloaded from the Pfam database (http://pfam.xfam.org/) (accessed on 21 August 2025), and a comparative analysis of ACX gene family members was performed using HMMER 3.0 with Hmmsearch. The identified candidates were validated via the SMART online platform (https://smart.embl.de/) (accessed on 21 August 2025) to confirm and finalize the SlACX gene family members [9].

2.2. Physicochemical Property Analysis of the SlACX Gene Family

The physicochemical properties of the SlACX gene family were completed through the online website ExPASy Proteomics Server (http://web.expasy.org/protparam) (accessed on 21 August 2025). Subcellular localization prediction was conducted using an online website (https://wolfpsort.hgc.jp/) (accessed on 21 August 2025) [10].

2.3. SlACX Family Gene Structure and Motifs Analysis

The conserved motifs of the SlACX family genes were predicted using the online software MEME (http://meme-suite.org/tools/meme) (accessed on 21 August 2025), and the gene structure and conserved motifs of the SlACX gene family were plotted using TBtool v2.0.6 [11].

2.4. Chromosomal Localization Analysis of the SlACX Gene Family

The chromosomal localization of the SlACX gene family members was accomplished through MapChart2.2 software and TBtools [12].

2.5. Phylogenetic Tree and Collinearity Analysis of the SlACX Gene Family

The phylogenetic trees of the SlACX family members of species such as A. thaliana, G. max., and Zea mays L. were constructed using MEGA 7.0 software, while the collinearity analysis was accomplished by the Linux system McscanX software 2012 [13].

2.6. Analysis of Cis-Acting Elements of the SlACX Gene Family

The cis-acting elements of 2000 bp upstream of the promoter of the SlACX gene family were analyzed through PlantCARE 2018, and then the quantities of expression elements such as hormone-induced, environmental adaptation, and adverse stress-induced were subjected to quantitative statistics [12].

2.7. Analysis of Tissue Specificity and Expression Under Different Stress Treatments

Uniform “Micro-Tom” tomato seeds were surface-sterilized with 1% sodium hypochlorite for 10 min and rinsed thoroughly with sterile water. The seeds were then germinated aseptically in a shaker (25 °C, 180 rpm) for three days with daily water changes. Subsequently, the germinated seeds were sown in soil and maintained in a growth chamber under controlled conditions (light: 250 μmol·m−2·s−1, 16/8 h light/dark at 26 ± 2/20 ± 2 °C, 60% RH). After two weeks, seedlings were transplanted to a hydroponic system, and uniform 21-day-old plants were selected for experimentation.
To analyze the expression patterns of SlRBOH genes during vegetative growth, we collected roots, stems, and leaves from untreated 21-day-old seedlings. For the SlACX gene expression analysis during reproductive growth, samples were taken from 55-day-old plants at the flowering stage, including roots, stems, leaves, and flowers, as well as corresponding green fruits (25 days after pollination) and mature fruits (45 days after pollination), with one fruit collected per plant. Each sample type consisted of three biological replicates, each comprising a pool of tissues from eight plants.
For chemical and osmotic stress treatments, seedlings were transferred to a half-strength nutrient solution containing one of the following: 200 mM NaCl, 100 μM ABA, 100 μM MeJA, or 20% (w/v) PEG 6000. For the oxidative stress treatment, seedlings were exposed to a nutrient solution supplemented with 10% (v/v) hydrogen peroxide. For temperature and light stress, seedlings were subjected to either 4 °C (cold) or 40 °C (heat) in incubators. The dark treatment group was placed in complete darkness, while the UV stress group was exposed to UV-C radiation (253.7 nm) under otherwise controlled conditions. The aerial tissues of seedlings from each treatment group, including a zero-hour control, were harvested after 0, 6, 12, and 24 h of exposure. Each sample, representing one biological replicate, was pooled from eight seedlings. All harvested tissues were immediately frozen in liquid nitrogen and stored at −80 °C. For every treatment and time point, three independent biological replicates were collected.

2.8. RNA Extraction and Real-Time Fluorescence Quantitative Analysis

The expression patterns of the SlACX gene family in different tissues and under different treatments were quantitatively analyzed by real-time fluorescence. The specific process was as follows: Total RNA was extracted using TRIzol, and the purity and integrity of RNA were strictly evaluated by gel electrophoresis. Subsequently, high-quality total RNA (1 µg) was reverse-transcribed into first-strand complementary DNA (cDNA) using a PrimeScript RT reagent kit with gDNA Eraser (TaKaRa, Dalian, China) according to the manufacturer’s instructions to provide a stable template for subsequent gene expression analysis [14]. Then, using cDNA as the template, specific primers, SYBR Green fluorescent dye, and premix were added to prepare the qPCR reaction system, and the PCR process was carried out. Ultimately, the change in the expression level of the target gene relative to the SlActin (NC015447.3) was calculated by analyzing the Ct values and using the ΔΔCt method [15]. To ensure the reliability and statistical significance of the experimental results, all experiments in this study were independently repeated three times (Table 1).

2.9. Statistical Analysis

The figure was drawn using Origin software (v10.3, OriginLab, Hampton, MA, USA). The significance of the difference was analyzed using the single-factor analysis method of SPSS 29.0.2 software (p < 0.05).

3. Results

3.1. Genome-Wide Identification of SlACX Genes in Solanum lycopersicum

Six members of the SlACX gene family were finally obtained and named SlACX1–SlACX6, respectively, according to their chromosomal positions. The ORF of SlACX1 was the largest, and that of SlACX6 was the smallest. The same was true for the number of amino acids and molecular weight. The pI distribution of SlACX1–SlACX6 ranged from 6.31 (SlACX3) to 8.74 (SlACX1). The protein domain positions owned by the members of the SlACX gene family were all different, but there was an overlap (Table 2).
The secondary structure of the SlACX family member peptides consists of alpha helix, beta turn, and random coil. The alpha helix accounts for the largest proportion, followed by the random coil, with the beta turn the least prevalent (Table 3). Among them, the proportion of SlACX3 in the alpha helix is 60.09%, the proportion of SlACX6 in the beta turn is 7.01%, and the proportion of SlACX5 in the random coil is 34.85%.
The subcellular localization of SlACX1–SlACX6 genes was determined, and the results indicated that SlACXs are located in the peroxisome, endoplasmic reticulum, plasma membrane, chloroplast, nucleus, vacuole, mitochondria, and cytoplasm. Among them, SlACX2 is distributed in the cytoplasm, SlACX6 is distributed in the peroxisome, and other genes are distributed in all organelle species (Table 4).

3.2. Analysis of Conserved Motifs and Gene Structure of the SlACX Family Genes

The SlACX family gene structure analysis results show that the SlACX1–SlACX6 genes can be classified into three groups (Class I, Class II, and Class III) based on evolutionary tree kinship. Class I includes SlACX3 and SlACX2, Class II includes SlACX6 and SlACX5, and Class III includes SlACX1 and SlACX4 (Figure 1).

3.3. Chromosomal Location of Tomato SlACX Genes

Figure 2 shows that the members of the SlACXs gene family are distributed in Chr04 (SlACX1), Chr08 (SlACX2 and SlACX3), and Chr10 (SlACX4, SlACX5, and SlACX6), and SlACX2 and SlACX3 form gene clusters.

3.4. Conserved Motif Analysis of Tomato SlACX Proteins

The motif analysis results show that motif 10, motif 2, motif 3, motif 4, and motif 5 are shared by this family. According to the differences in the number and types of motifs, the SlACX1–SlACX6 genes are classified into three groups, which is consistent with the grouping of gene structures. Most members in the same group have similar motifs, indicating that there are functional similarities among the genes in the same group. From the motif sequences in Table 5, it can be seen that motifs 1, 2, 3, 4, and 7 may jointly constitute the FAD (flavin adenine dinucleotide) binding domain and catalytic core of the ACX enzyme. Motifs 5 and 8 may be involved in the formation of substrate (acyl-CoA) binding channels, responsible for recognizing and binding to aliphatic acyl chains of different lengths. Motifs 6, 9, and 10 are involved in maintaining the tertiary structure of proteins, mediating protein–protein interactions, or undergoing post-transcriptional modifications (such as phosphorylation), thereby precisely regulating enzyme activity (Figure 3).

3.5. Phylogenetic Analysis of SlACX Family Members

An evolutionary tree was constructed based on the ACX gene family members of Arabidopsis thaliana, Glycine max, Oryza sativa, Solanum lycopersicum, and Zea mays (Figure 4). According to the evolutionary relationship, 27 ACX genes can be divided into three major groups (Class I, Class II, and Class III), among which Class I contains the most genes (12 genes), and Class II contains 4 genes. Class III contains 11 genes. SlACX5, SlACX6, and AtACX4 have a relatively close evolutionary relationship, SlACX1 and AtACX2 have a relatively close evolutionary relationship, and SlACX4 has a relatively close evolutionary relationship with AtACX3 and AtACX6. SlACX2 and SlACX3 belong to independent small branches and have a relatively close evolutionary relationship with Glycine max.

3.6. Analysis of Gene Duplication and Collinearity of SlACX Genes

The evolutionary relationships of SlACX family genes between and within species were studied. As shown in Figure 5A, there is a significant collinearity between SlACX5 and AtACX6. There is no collinearity among the other family members. When collinearity was constructed among A. thaliana, S. lycopersicum, and S. tuberosum, it was found that SlACX2 is collinear with At-Chr2, At-Chr4, At-Chr5, and St-Chr08, and SlACX1 is collinear with At-Chr5 and St-Chr04. SlACX4 and SlACX5 are collinear with St-Chr10 (Figure 5B).

3.7. Cis-Acting Element Analysis of Tomato SlACX Genes

The promoter of the SlACXs genes contains various functional elements, such as LTR, MBS, ABRE, TGA-element, and P-box (Figure 6). According to their functional effects, the homeogenic functional elements are classified into adverse response elements, light response elements, hormone response elements, and growth and development response elements (Figure 7). Moreover, there are differences in quantity, indicating that SlACXs can be involved in regulating a series of abiotic stress conditions (Table 6).

3.8. Expression Analysis of SlACX Genes in Different Organs

Unopened flower bud, fully opened flower, mature green fruit, breaker fruit, pimpinellifolium immature green fruit, leaves, pimpinellifolium immature breaker fruit, and root were selected as test materials. As shown in Figure 8, compared with the unopened flower bud, SlACX1, SlACX3, SlACX4, SlACX5, and SlACX6 have higher expression levels in the fully opened flower. Compared with leaves, the expression levels of SlACX1, SlACX2, SlACX4, and SlACX5 are higher in the roots. Compared with mature green fruit, the expression levels of SlACX1, SlACX2, SlACX3, SlACX5, and SlACX6 are higher in breaker fruit. Overall, the SlACX family genes are highly expressed in roots, leaves, and fruits, whether it is green fruit or breaker fruit.

3.9. Expression Analysis of Tomato SlACX Genes Under Hormonal and Abiotic Stress

Analysis of the expression dynamics of the SlACX gene family under UV and dark treatments revealed that the response patterns exhibited significant photodependence and temporal specificity (Figure 9). After 12 h of UV treatment, the expression levels of SlACX1, SlACX2, SlACX4, SlACX5, and SlACX6 rose sharply, and the expression level of SlACX1 continued to increase at 24 h, showing the rapid activation characteristics of core response genes. The expression levels of the SlACX2, SlACX4, SlACX5, and SlACX6 genes showed a downward trend within 24 h. In contrast, only SlACX1 and SlACX5 showed a significant increase at 12 h in the dark treatment, while the gene expression levels of the other family members were relatively low, indicating that the function of this family is mainly concentrated on the response to light stress. It is worth noting that SlACX1 maintained a high expression level under both treatments and may be a key gene under non-stress conditions.
In this study, the expression of the SlACX gene family under ABA and MeJA treatments was analyzed by qRT-PCR, and it was found that the response patterns presented distinct hormone-specific and temporal characteristics. After 6 h of ABA treatment, the expression levels of SlACX1, SlACX2, SlACX4, SlACX5, and SlACX6 decreased. Then, with the extension of stress time, the expression level of SlACXs gradually increased. After 6 h of MeJA treatment, SlACX1, SlACX5, and SlACX6 were induced to respond rapidly, and their expression levels continued to increase after 12 h. After 24 h of stress treatment, their expression levels decreased. In summary, SlACX1/SlACX5 and SlACX6, serving as response modules for ABA and MeJA signals, respectively, may differentially influence the stress resistance, energy metabolism, and secondary synthesis of plants by regulating the rate of fatty acid degradation (Figure 10).
To verify the expression of SlACX under different abiotic stresses, this study conducted quantitative analyses on SlACX under cold, heat, H2O2, PEG, and NaCl treatments. The results showed that the expression level of SlACX1 increased rapidly at 6 h and decreased at 24 h. This is consistent with the expression trends of SlACX2, SlACX4, SlACX5, and SlACX6, while SlACX3 shows typical low-temperature adaptation characteristics, suggesting that it may be involved in the membrane lipid remodeling process in the later stage of low-temperature stress (Figure 11A). Figure 11B shows that heat stress induces a vigorous response. SlACX3 showed explosive upregulation at 6 h, a slow increase at 12 h, and a sharp rise in the expressions of SlACX3, SlACX4, SlACX5, and SlACX6 after 24 h, presenting a typical pattern of acute heat shock response. SlACX2 remained at a persistently low expression throughout the process and might not be sensitive to heat stress.
Under oxidative stress, SlACX1 showed specific high expression at 6 h, which coincides with the time point of the reactive oxygen species outbreak, suggesting that it may directly participate in the clearance of fatty acid peroxides in oxidative damage repair. The remaining SlACX genes did not show a significant expression trend under oxidative stress (Figure 11C). Under PEG conditions, the expression level of SlACX6 reached its peak at 6 h and then began to decline, while the expression levels of SlACX1 and SlACX2 reached their peaks at 12 h. SlACX3, SlACX4, and SlACX5 are not sensitive to PEG processing (Figure 11D). Under salt stress treatment, the expression of the SlACX3 gene was the most significant. From 6 h to 12 h, the expression level of SlACX3 continued to increase but decreased after 24 h. SlACX1, SlACX2, SlACX4, SlACX5, and SlACX6 showed a trend of expression, but it was not obvious (Figure 11E).
Overall, the SlACX gene family exhibits fine functional differentiation under abiotic stress, with SlACX1 showing a rapid response to acute stress (cold, H2O2, and PEG). SlACX3, on the other hand, specifically corresponds to ion stress adaptation. This temporal and stress-specific expression pattern reveals that SlACX plays a multi-faceted core role in energy supply, membrane lipid remodeling, and signal molecule generation by regulating fatty acid degradation fluxes, providing a key target for crop multi-resistance breeding.

4. Discussion

In recent years, with the gradual improvement and optimization of tomato genome sequencing work, gene family identification has provided important clues for studying gene-related functions [16]. However, there are still many genes that have not been reported, such as the ACX gene. In this study, six SlACX gene family members were identified from the entire tomato genome using bioinformatics analysis methods, distributed on chromosomes Chr04, Chr08, and Chr10. Based on the gene structure and motif structure of the SlACX genes, the six SlACX gene family members were classified into three major categories. Proteins with similar conserved motifs and conserved domains in each major category are clustered together [17], indicating a highly conserved evolution and possibly having similar structures, functions, and evolutionary characteristics [18].
In the systematic analysis of gene families, the integrated study of evolutionary trees, collinearity, and cis-acting elements constitutes the core framework for revealing the functional evolution and regulatory mechanisms of genes [19]. Phylogenetic evolutionary trees construct the kinship among members through sequence alignment and serve as the cornerstone of functional prediction [20]. In this study, the ACX gene family members of species such as soybeans, corn, Arabidopsis thaliana, and rice were used to build an evolutionary tree. The results showed that SlACXs and AtACXs had multiple branches, indicating that the genes between the two species are likely to have similar functions [21]. Collinearity analysis provides insights into the evolutionary history of genes from a macrogenomic perspective [22]. By comparing locus conservation across species, this approach enables precise differentiation between orthologous and paralogous genes [23]. Interspecific and intraspecific collinearity analyses of the SlACX gene family in this study reveal that gene expansion has occurred during tomato evolution, potentially contributing to more sophisticated metabolic regulatory mechanisms in tomatoes [2]. Furthermore, the observed interspecific collinearity with Arabidopsis thaliana and soybean strongly supports the hypothesis that the ACX gene family performs ancient and fundamental physiological roles in plants, with highly conserved functions across lineages [24].
Cis-acting element analysis can infer which signaling pathways a gene may respond to by scanning conserved motifs in the gene promoter region, thereby linking gene function to specific biological processes [25]. In this study, members of the SlACXs gene family were found to not only encode the key rate-limiting enzymes of the fatty acid β-oxidation pathway, but their promoter regions are also more densely distributed with a variety of cis-acting elements, including hormone response elements, abiotic stress response elements, light response elements, and elements related to growth and development. This feature strongly suggests that the SlACX gene may be far from being a simple metabolic enzyme gene, rather serving as a key signal integration node by precisely coordinating the growth and development of tomatoes and their stress defense strategies in complex and variable environments [26].
To verify the above hypotheses and deeply analyze the specific functional differentiation and collaborative mechanisms of the SlACX family in environmental adaptation, this study analyzed the expression patterns of the SlACX gene under different treatments. The expression analysis results clearly reveal that the expression pattern of the SlACX gene family shows extremely significant time dependence and treatment specificity, and the complexity and precision of its regulation far exceed expectations [2]. Under light processing, the responses of SlACX members to UV-B and dark conditions are completely different. Under UV-B irradiation, the family members showed explosive upregulation within 12 h, presenting typical characteristics of an acute stress response. This is likely closely related to the UV-induced reactive oxygen species (ROS) outbreak and the subsequent activation of jasmonic acid signaling [27], aiming to rapidly provide fatty acid degradation products as repair substances or precursors of defense signals [28]. Under dark conditions, only a few family members showed a significant upregulation at 12 h, which might be related to the reprogramming of energy metabolism, that is, maintaining basic energy supply by enhancing the β-oxidation of fatty acids after photosynthesis stops [29]. In hormone treatment, the response by SlACX to ABA and MeJA is particularly remarkable. Most SlACX members were strongly induced to express by ABA at 24 h of treatment, indicating that this family may be a key downstream target of the ABA signaling pathway [30] and may play a core energy supply role in ABA-mediated drought and salt stress responses, such as stomatal closure and osmotic regulation [31]. Particularly crucial is that the response patterns to MeJA showed significant differentiation, with some members reaching their peak expression levels at 12 h. This differentiation highly likely reflects the complex dual role of ACX in JA biosynthesis (as a precursor provider itself) [7] and signal transduction (possibly subjected to feedback suppression), providing a new perspective for understanding the fine regulation of the JA signaling network [30].
Under abiotic stress, the response of the SlACX family demonstrated a more refined functional division of labor. Under cold stress, the expression levels of some members of the SlACX family rose slowly, synchronizing with the process of plant acclimation at low temperatures. High-temperature stress, on the other hand, immediately caused a high level of expression of SlACXs, suggesting its involvement in thermal shock protection. For oxidative stress (H2O2), the transient high expression of a specific SlACX gene (SlACX1) suggests that it is directly involved in the clearance of oxidative damage to lipids. In PEG and NaCl treatments, although both caused osmotic stress, there were significant differences in the response time and intensity spectra of SlACX members, indicating that they could distinguish between drought and salt damage signals and initiate differentiated metabolic adaptation strategies.
In summary, through multi-dimensional expression profiling analysis, this study not only confirmed that the SlACX gene family is a core hub for tomatoes to respond to external signals but also initially mapped out its functional division. These genes, through precise temporal dynamics and specific expression regulation, convert external signals such as light, hormones, and stress into internal metabolic flow reprogramming, thereby precisely regulating the adaptability and stress resistance of plants at the levels of energy supply, signal molecule generation, and osmotic protective substance synthesis. These results have laid a solid theoretical foundation for the subsequent molecular breeding of tomato stress resistance using the key SlACX gene.

5. Conclusions

This study systematically revealed the evolutionary conservation and functional diversity of the SlACX gene family in tomatoes. This family not only acts as the executor of fatty acid β-oxidation but, through its complex regulatory element library in the promoter region, also becomes a key signal hub integrating light signals, hormone networks, and diverse stress factors. The expression pattern further confirms that family members exhibit fine functional division of labor and synergy when responding to different environmental challenges. Some genes dominate stress signal transduction, while others may maintain basal metabolic homeostasis. This “functional modularization” feature enables the SlACX family to precisely regulate the direction of carbon source flow, achieving a dynamic balance among energy supply, defense substance synthesis, and system resistance establishment, thereby endowing tomatoes with strong environmental adaptability. The research results provide precise targets and a theoretical basis for molecular design breeding of tomato stress resistance using key SlACX genes.

Author Contributions

Conceptualization, C.W.; data curation, C.W. and Z.L.; resources, C.W. and Z.L.; formal analysis, Y.G.; funding acquisition, C.W.; investigation, Q.L. and Q.W.; methodology, C.A., C.W. and Z.L.; software, Q.L. and Y.G. All authors have read and agreed to the published version of the manuscript.

Funding

This work was supported by the Fuxi Young Talents Fund of Gansu Agricultural University (Gaufx-03Y07); the Supporting Funds for Youth Mentor of Gansu Agricultural University (GAU-QDFC-2024-15); the National Natural Science Foundation of China (32460753); and the Key Project of Gansu Provincial Natural Science Foundation, China (23JRRA1406).

Data Availability Statement

Data are contained within the article.

Conflicts of Interest

The authors declare no conflicts of interest.

References

  1. Li, Y.; Zhang, F.; Chen, B.; Du, W.; Fang, S.; Wang, G.; Hu, J.; Li, L.; Hu, Z.; Jia, Z.; et al. Identification of MFS gene family in tomato and functional characterization of SlZIF1/SlMFS4 under salt stress. Plant Physiol. Biochem. PPB 2025, 229, 110554. [Google Scholar] [CrossRef]
  2. He, R.; Ma, W.; Zhou, F.; Cao, H.; Kang, Z.; Dong, J.; Xing, J. Molecular characterization of acyl-CoA oxidase (ACX) family genes in maize reveals their role in disease resistance. Genes 2025, 16, 486. [Google Scholar] [CrossRef]
  3. Xin, Z.; Chen, S.; Ge, L.; Li, X.; Sun, X. The involvement of a herbivore-induced acyl-CoA oxidase gene, CsACX1, in the synthesis of jasmonic acid and its expression in flower opening in tea plant (Camellia sinensis). Plant Physiol. Biochem. 2019, 135, 132–140. [Google Scholar] [CrossRef] [PubMed]
  4. Liu, W.; Han, T.; Yuan, H.; Yu, Z.; Zhang, L.; Zhang, B.; Zhai, S.; Zheng, S.; Lu, Y. CATALASE2 functions for seedling postgerminative growth by scavenging H2O2 and stimulating ACX2/3 activity in Arabidopsis. Plant Cell Environ. 2017, 40, 2720–2728. [Google Scholar] [CrossRef] [PubMed]
  5. Xi, W.; Zhang, B.; Liang, L.; Shen, J.; Wei, W.; Xu, C.; Allan, A.; Ferguson, I.; Chen, K. Postharvest temperature influences volatile lactone production via regulation of acyl-CoA oxidases in peach fruit. Plant Cell Environ. 2012, 35, 534–545. [Google Scholar] [CrossRef] [PubMed]
  6. Jia, D.; Bin, L.; Yan, H. Molecular cloning and characterization of three novel genes related to fatty acid degradation and their responses to abiotic stresses in Gossypium hirsutum L. J. Integr. Agric. 2012, 12, 582–588. [Google Scholar]
  7. Rafeiza, B.; Raquel, A.; Zolman, B.K. Peroxisomal acyl-CoA oxidase 4 activity differs between Arabidopsis accessions. Plant Mol. Biol. 2012, 78, 45–58. [Google Scholar]
  8. Yang, H.; Bai, X.; Muhammad, T.; Jia, C.; Baike, W.; Yu, Q.; Wang, J. GHMP gene family: Identification, evolutionary and expression analysis under various exogenous hormones and abiotic stress in tomato. BMC Plant Biol. 2025, 25, 850. [Google Scholar] [CrossRef]
  9. Mo, F.; Xue, X.; Wang, J.; Mozhen, C.; Liu, S.; Zhao, L.; Chen, X. Genome-wide analysis of KCS genes in tomato and functional characterization of SlKCS8 and SlKCS10 in drought tolerance. Plant Physiol. Biochem. 2025, 222, 109783. [Google Scholar] [CrossRef]
  10. Can, H.; Dogan, I.; Uras, E.M. Genome-wide screening of mitogen-activated protein kinase (MAPK) gene family and expression profile under heavy metal stress in Solanum lycopersicum. Biotechnol. Lett. 2025, 47, 27. [Google Scholar] [CrossRef]
  11. Cui, X.; Gu, J.; Liu, P.; Zhen, R.; Zhang, Y.; Wang, F.; Qi, M.; Liu, Y.; Li, T. Genome-wide identification and characterization of the thioredoxin (TRX) gene family in tomato (Solanum lycopersicum) and a functional analysis of SlTRX2 under salt stress. Plant Physiol. Biochem. 2025, 220, 109478. [Google Scholar] [CrossRef]
  12. Sun, H.; Yang, J.; Fan, B.; Wang, Y.; Chen, G.; Cheng, G. Genome-wide analysis of BURP domain-containing gene family in Solanum lycopersicum and functional analysis of SlRD1 under drought and salt stresses. Int. J. Mol. Sci. 2024, 25, 12539. [Google Scholar] [CrossRef]
  13. Xu, J.; Cui, J.; He, Q.; Liu, Y.; Lu, X.; Qi, J.; Xiong, J.; Yu, W.; Li, C. Genome-wide identification of HIPP and mechanism of SlHIPP4/7/9/21/26/32 mediated phytohormones response to Cd, osmotic, and salt stresses in tomato. Plant Physiol. Biochem. PPB 2024, 217, 109220. [Google Scholar] [CrossRef] [PubMed]
  14. Wang, C.; Zhang, J.; Li, J. Genome-wide identification and functional analysis of the trehalose-6-phosphate phosphatase (TPP) gene family in tomato (Solanum lycopersicum L.) and the role of SlTPP3 under NaCl stress. Sci. Hortic. 2025, 345, 114161. [Google Scholar] [CrossRef]
  15. Almeida, J.; Fons, P.L.; Drapal, M.; Kit, L.; Eugenia, M.; Paul, D. The FIBRILLIN multigene family in tomato, their roles in plastoglobuli structure and metabolism. Plant J. Cell Mol. Biol. 2025, 123, e70447. [Google Scholar] [CrossRef] [PubMed]
  16. Wang, X.; Dai, L.; Yang, Y.; Zhu, Y. Identification and expression pattern analysis of the PP2C gene family response to salt stress in tomato. Russ. J. Plant Physiol. 2025, 72, 122. [Google Scholar] [CrossRef]
  17. Cao, H.; Wang, D.; Li, X.; Zhang, Y.; Su, D.; Wang, L.; Xu, K.; Li, Z. Genome-wide identification and expression profiling of SlGeBP gene family in response to hormone and abiotic stresses in Solanum lycopersicum L. Int. J. Mol. Sci. 2025, 26, 6008. [Google Scholar] [CrossRef]
  18. Arif, M.; Abbas, H.; Mahmood, N.; Muhammad, U.; Muhammad, A.; Shahbaz, A.; Yao, X.; Xu, R.; Li, L. Genome-Wide analysis of the NRAMP gene family in Arabidopsis thaliana: Identification, expression and response to multiple heavy metal stresses and phytohormones. BMC Plant Biol. 2025, 25, 1305. [Google Scholar] [CrossRef]
  19. Zhang, H.; Yu, Z.; Yao, X.; Chen, J.; Chen, X.; Zhou, H.; Lou, Y.; Ming, F.; Jin, Y. Genome-wide identification and characterization of small auxin-up RNA (SAUR) gene family in plants: Evolution and expression profiles during normal growth and stress response. BMC Plant Biol. 2021, 21, 4. [Google Scholar] [CrossRef]
  20. Cheng, S.; Hao, L.; Sun, J.; Ren, H.; Song, H. Genome-wide analysis of the SnRK gene family in Caragana Korshinskii and their expression profiling under drought and nitrogen deposition. BMC Genom. 2025, 26, 838. [Google Scholar] [CrossRef]
  21. Fu, G.; Yang, Y.; Mahmood, T.; Liu, X.; Xie, Z.; Zhao, Z.; Dong, Y.; Tian, Y.; Jehanzeb, F.; Iram, S.; et al. Genome-Wide Characterization of the ABI3 Gene Family in Cotton. Genes 2025, 16, 854. [Google Scholar] [CrossRef]
  22. Martina, J.; Thomas, D. Phylogenetic analysis of the expansion of the MATH-BTB gene family in the grasses. Plant Signal. Behav. 2014, 9, e28242. [Google Scholar]
  23. Yue, H.; Yang, Y.; Cha, G. Genomic colinearity and transcriptional regulatory networks of BES1 gene family in horticultural Plants particularly kiwifruit and peach. Horticulturae 2023, 9, 971. [Google Scholar] [CrossRef]
  24. Lv, M.; Lan, X.; Zhang, D.; Juye, Z.; Zhi, L.; Wei, Q.; Cui, Y. Evolution and diversification of adenine phosphoribosyltransferase (APT) gene family revealed their roles on flowering time in Brassica napus L. Ind. Crops Prod. 2025, 234, 121627. [Google Scholar] [CrossRef]
  25. Bhar, S.; Prajapati, V.D.; Gonzalez, S.M.; Yoon, C.-S.; Kevin, M.; Laura, S.; Kari, B.B.; Rebecca, A.B. Comparative metabolomics identifies the roles of acyl-CoA oxidases in the biosynthesis of ascarosides and a complex family of secreted n-Acylethanolamines. ACS Chem. Biol. 2025, 20, 1298–1308. [Google Scholar] [CrossRef]
  26. Zhang, L.; Li, H.; Gao, L.; Qi, Y.; Fu, W.; Li, X.; Zhou, X.; Gao, Q.; Gao, Z.; Jia, H. Acyl-CoA oxidase 1 is involved in γ-decalactone release from peach (Prunus persica) fruit. Plant Cell Rep. 2017, 36, 829–842. [Google Scholar] [CrossRef]
  27. Zhou, Z.; Naoufal, L.; Dounya, K.; Cao, H.; Zhang, K.; Dong, J.; Xing, J. Genome-wide identification and analysis of soybean acyl-ACP thioesterase gene family reveals the role of GmFAT to improve fatty acid composition in soybean seed. Theor. Appl. Genet. 2021, 134, 3611–3623. [Google Scholar] [CrossRef]
  28. Rylott, E.; Rogers, C.; Gilday, A.; Edgell, T.; Larson, T.; Graham, I. Arabidopsis mutants in short- and medium-chain acyl-CoA oxidase activities accumulate acyl-CoAs and reveal that fatty acid β-Oxidation is essential for embryo development. J. Biol. Chem. 2003, 278, 21370–21377. [Google Scholar] [CrossRef]
  29. Zhu, X.; Hou, Z.; Xu, X.; Xiong, Z.; Chen, Y.; Yang, L.; Liu, Z.; Fang, Z. TMT-based comparative proteomics reveals the role of acyl-CoA oxidase 4 in enhancing the drought stress tolerance in common buckwheat (Fagopyrum esculentum). Int. J. Biol. Macromol. 2022, 215, 262–271. [Google Scholar] [CrossRef]
  30. Pedersen, L.; Henriksen, A. Acyl-CoA Oxidase 1 from Arabidopsis thaliana. Structure of a key enzyme in plant lipid metabolism. J. Mol. Biol. 2004, 345, 487–500. [Google Scholar] [CrossRef]
  31. Kim, C.M.; Kim, H.T.; Park, H.J.; Lee, C.; Cho, S. Expression of rice acyl-CoA oxidase isoenzymes in response to wounding. J. Plant Physiol. 2006, 164, 665–668. [Google Scholar] [CrossRef]
Horticulturae 11 01426 g001
Figure 1. Gene structure of tomato SlACX genes. The evolutionary tree was constructed based on the full length of tomato SHMT protein sequences using MEGA7.0. The exon–intron graph of tomato SHMT genes was drawn using TBtools v1.0986.
Figure 1. Gene structure of tomato SlACX genes. The evolutionary tree was constructed based on the full length of tomato SHMT protein sequences using MEGA7.0. The exon–intron graph of tomato SHMT genes was drawn using TBtools v1.0986.
Horticulturae 11 01426 g001
Horticulturae 11 01426 g002
Figure 2. The distribution of SlACX gene family members on chromosomes in Solanum lycopersicum.
Figure 2. The distribution of SlACX gene family members on chromosomes in Solanum lycopersicum.
Horticulturae 11 01426 g002
Horticulturae 11 01426 g003
Figure 3. The motif composition and distribution of tomato SlACX proteins. (A) Colored boxes represent different conserved motifs. (B) Motif1–Motif10.
Figure 3. The motif composition and distribution of tomato SlACX proteins. (A) Colored boxes represent different conserved motifs. (B) Motif1–Motif10.
Horticulturae 11 01426 g003
Horticulturae 11 01426 g004
Figure 4. The unrooted phylogenetic tree of the ACX gene family in Solanum lycopersicum, Arabidopsis thaliana, Glycine max, Oryza sativa, and Zea mays. The maximum likelihood method was used to construct a phylogenetic tree containing 6 Solanum lycopersicum (Sl), 6 Arabidopsis thaliana (At), 4 Oryza sativa (Os), 6 Zea mays (Zm), and 5 Glycine max ACX proteins. The three subgroups are colored differently. The three differently colored shapes represent ACX proteins from five species. The red triangle, black circle, matcha circle, light blue pentagram, and blue triangle represent Arabidopsis thaliana, Glycine max, Oryza sativa, Solanum lycopersicum, and Zea mays ACX proteins, respectively.
Figure 4. The unrooted phylogenetic tree of the ACX gene family in Solanum lycopersicum, Arabidopsis thaliana, Glycine max, Oryza sativa, and Zea mays. The maximum likelihood method was used to construct a phylogenetic tree containing 6 Solanum lycopersicum (Sl), 6 Arabidopsis thaliana (At), 4 Oryza sativa (Os), 6 Zea mays (Zm), and 5 Glycine max ACX proteins. The three subgroups are colored differently. The three differently colored shapes represent ACX proteins from five species. The red triangle, black circle, matcha circle, light blue pentagram, and blue triangle represent Arabidopsis thaliana, Glycine max, Oryza sativa, Solanum lycopersicum, and Zea mays ACX proteins, respectively.
Horticulturae 11 01426 g004
Horticulturae 11 01426 g005
Figure 5. Gene duplication and collinearity analysis of ACX genes. (A) Intraspecies collinearity analysis of SlACXs. (B) Interspecies collinearity analysis of SlACXs, AtACXs, and StACXs.
Figure 5. Gene duplication and collinearity analysis of ACX genes. (A) Intraspecies collinearity analysis of SlACXs. (B) Interspecies collinearity analysis of SlACXs, AtACXs, and StACXs.
Horticulturae 11 01426 g005
Horticulturae 11 01426 g006
Figure 6. Cis-element distribution in SlACX genes of Solanum lycopersicum.
Figure 6. Cis-element distribution in SlACX genes of Solanum lycopersicum.
Horticulturae 11 01426 g006
Horticulturae 11 01426 g007
Figure 7. The number of cis-acting elements in SlACX genes.
Figure 7. The number of cis-acting elements in SlACX genes.
Horticulturae 11 01426 g007
Horticulturae 11 01426 g008
Figure 8. Expression levels of SlACX genes in different tissues.
Figure 8. Expression levels of SlACX genes in different tissues.
Horticulturae 11 01426 g008
Horticulturae 11 01426 g009
Figure 9. Expression levels of SlACX genes under (A) UV and (B) dark conditions. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Figure 9. Expression levels of SlACX genes under (A) UV and (B) dark conditions. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Horticulturae 11 01426 g009
Horticulturae 11 01426 g010
Figure 10. Expression levels of SlACX genes under (A) ABA and (B) MeJA. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Figure 10. Expression levels of SlACX genes under (A) ABA and (B) MeJA. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Horticulturae 11 01426 g010
Horticulturae 11 01426 g011
Figure 11. Expression levels of SlACX genes under (A) Cold, (B) Heat, (C) H2O2, (D) PEG, and (E) NaCl. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Figure 11. Expression levels of SlACX genes under (A) Cold, (B) Heat, (C) H2O2, (D) PEG, and (E) NaCl. The different lowercase letters in the figure indicate significant differences in processing times (p < 0.01).
Horticulturae 11 01426 g011
Table 1. qRT-PCR primers for expression analysis of the SlACX gene family.
Table 1. qRT-PCR primers for expression analysis of the SlACX gene family.
Gene NameSequence (F)Sequence (R)
SlACX1AGAACACTTACGCAACCCTAACTTCTATTCCAAGCACCAAATCCTCCAAG
SlACX2CTCAAGAGGAAGCCATATGTTAAGGCTGCTGTTTATCTGTGCCCTGTC
SlACX3CAACCACTGCCACTGCTGATGCTGAAGTAGTAGCACGACGTTATCTC
SlACX4AAAAGGAGTAAGAAACCGGGCAGGTCTCGTTCCCGTAGGC
SlACX5CAACACCAGCGTCCGTCTTTCTTTCCATGCACTCTCTCACTTTCAG
SlACX6TGTCACAGGTACTTGAAGGAGAGGATAGCCGCCAACCAACAAGAATC
Table 2. Genomic information on members of the SlACX family genes in Solanum lycopersicum.
Table 2. Genomic information on members of the SlACX family genes in Solanum lycopersicum.
GeneGene IDGene LocusORF
(bp)		Amino AcidMolecular
Weight		pIACX Domain Locations
SlACX1Solyc04g054890.3.1.ITAG4.0Chr04206768877,398.588.74176–292; 323–484; 529–686
SlACX2Solyc08g078390.4.1.ITAG4.0Chr08199567875,624.548.3517–132; 134–247; 479–656
SlACX3Solyc08g078400.3.1.ITAG4.0Chr08195064972,643.346.3117–132; 134–246; 479–597
SlACX4Solyc10g008110.4.1.ITAG4.0Chr10205268376,734.678.15191–300; 331–489; 529–669
SlACX5Solyc10g076600.2.1.ITAG4.0Chr10132043948,316.048.2256–167; 171–263; 277–423
SlACX6Solyc10g085200.2.1.ITAG4.0Chr10128742847,117.457.5546–156; 160–252; 266–412
Table 3. The secondary structure of the SlACX gene family in Solanum lycopersicum.
Table 3. The secondary structure of the SlACX gene family in Solanum lycopersicum.
ProteinAlpha Helix (%)Beta Turn (%)Random Coil (%)
SlACX155.095.8128.05
SlACX259.594.7227.14
SlACX360.094.3127.27
SlACX453.945.2027.24
SlACX546.015.2434.85
SlACX649.777.0129.91
Table 4. Subcellular localization prediction of the SlACX gene family in Solanum lycopersicum.
Table 4. Subcellular localization prediction of the SlACX gene family in Solanum lycopersicum.
GenePeroxisomeEndoplasmic ReticulumPlasma MembraneChloroplastNucleusVacuoleMitochondriaCytoplasm
SlACX1—3141.5211
SlACX2———121—10
SlACX37321—1——
SlACX4——154—3—
SlACX51551—2——
SlACX612——————2
Table 5. Details of the 10 conserved motifs of SlACX proteins.
Table 5. Details of the 10 conserved motifs of SlACX proteins.
MotifWidth (aa)Motif Sequence
Motif 129CYALTELGHGSNVQGLETTATFDPGTDEF
Motif 234LNGVDNGVJLFDNVRIPRDDLLPRVADVSKDGKY
Motif 328DYQTQQQKLVPLLASTYAFRFVGWRLKK
Motif 421SGAVDIATRYSAVRKQFGAPN
Motif 521EPIYTFEGDNDVLLLQVARFL
Motif 629IHSPTLTASKWWPGGLGKVSTHAIVYARL
Motif 729NAFIVQJRSLEDHKPAPGVQVTDIGNKIG
Motif 841PNCTSDYYQLDDLLTPEEKAIRLKVRECMEKEIAPIMTKYW
Motif 950DVTQRLKANDFSTLPEVHACTAGLKSLTTSATADGIEECRKLCGGHGYLC
Motif 1031HSGLFIPAIKLQGSEMQKEKWLPSAYDMQII
Table 6. Summary of cis-acting elements of SlACX genes.
Table 6. Summary of cis-acting elements of SlACX genes.
ElementSequenceDescription
LTRCCGAAAcis-acting element involved in low-temperature responsiveness
MBSCAACTGMYB binding site involved in drought inducibility
TC-rich repeatsATTCTCTAAC/GTTTTCTTACcis-acting element involved in defense and stress responsiveness
AREAAACCAcis-acting regulatory element essential for the anaerobic induction
Box IICCACGTGGCpart of a light-responsive element
ACECTAACGTATT/GACACGTATGcis-acting element involved in light responsiveness
AE-boxAGAAACTT/AGAAACAApart of a module for light response
AT1-motifAATTATTTTTTATTpart of a light-responsive module
ATC-motifAGTAATCTpart of a conserved DNA module involved in light responsiveness
ATCT-motifAATCTAATCCpart of a conserved DNA module involved in light responsiveness
Box 4ATTAATpart of a conserved DNA module involved in light responsiveness
chs-CMA1aTTACTTAApart of a light-responsive element
GATA-motifGATAGGG/AAGGATAAGG/AAGATAAGATTpart of a light-responsive element
G-boxCCACGTAA/TACGTG/TAACACGTAG/GCCACGTGGA/CACGTC/TCCACATGGCA/CACGTG/CACGTTcis-acting regulatory element involved in light responsiveness
GT1-motifGGTTAA/GGTTAATlight-responsive element
I-boxAAGATAAGGCT/AGATAAGGpart of a light-responsive element
LAMP-elementCTTTATCApart of a light-responsive element
MREAACCTAAMYB binding site involved in light responsiveness
TCT-motifTCTTACpart of a light-responsive element
ABREACGTG/CACGTG/TACGTGTCcis-acting element involved in the abscisic acid responsiveness
CGTCA-motifCGTCAcis-acting regulatory element involved in the MeJA responsiveness
GARE-motifTCTGTTGgibberellin-responsive element
TGACG-motifTGACGcis-acting regulatory element involved in the MeJA responsiveness
TGA-elementAACGACauxin-responsive element
P-boxCCTTTTGgibberellin-responsive element
TATC-boxTATCCCAcis-acting element involved in gibberellin responsiveness
TCA-elementCCATCTTTTTcis-acting element involved in salicylic acid responsiveness
CAT-boxGCCACTcis-acting regulatory element related to meristem expression
GCN4_motifTGAGTCAcis-regulatory element involved in endosperm expression
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Wang, C.; Liu, Z.; Gao, Y.; Li, Q.; Wang, Q.; An, C. Genome-Wide Identification and Characterization of Tomato Acyl-CoA Oxidase Family Genes ACX. Horticulturae 2025, 11, 1426. https://doi.org/10.3390/horticulturae11121426

AMA Style

Wang C, Liu Z, Gao Y, Li Q, Wang Q, An C. Genome-Wide Identification and Characterization of Tomato Acyl-CoA Oxidase Family Genes ACX. Horticulturae. 2025; 11(12):1426. https://doi.org/10.3390/horticulturae11121426

Chicago/Turabian Style

Wang, Chunlei, Zesheng Liu, Yanlong Gao, Qianbing Li, Qi Wang, and Caiting An. 2025. "Genome-Wide Identification and Characterization of Tomato Acyl-CoA Oxidase Family Genes ACX" Horticulturae 11, no. 12: 1426. https://doi.org/10.3390/horticulturae11121426

APA Style

Wang, C., Liu, Z., Gao, Y., Li, Q., Wang, Q., & An, C. (2025). Genome-Wide Identification and Characterization of Tomato Acyl-CoA Oxidase Family Genes ACX. Horticulturae, 11(12), 1426. https://doi.org/10.3390/horticulturae11121426

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop