Temperature mediates ROS metabolism and MAPK signaling pathways to influence Scutellaria baicalensis quality

Temperature mediates ROS metabolism and MAPK signaling pathways to influence Scutellaria baicalensis quality | Authorea try { document.documentElement.classList.add('js'); } catch (e) { } var _gaq = _gaq || []; _gaq.push(['_setAccount', 'G-8VDV14Y67G']); _gaq.push(['_trackPageview']); (function() { var ga = document.createElement('script'); ga.type = 'text/javascript'; ga.async = true; ga.src = ('https:' == document.location.protocol ? 'https://ssl' : 'http://www') + '.google-analytics.com/ga.js'; var s = document.getElementsByTagName('script')[0]; s.parentNode.insertBefore(ga, s); })(); Skip to main content Preprints Collections Wiley Open Research IET Open Research Ecological Society of Japan All Collections About About Authorea FAQs Contact Us Quick Search anywhere Search for preprint articles, keywords, etc. Search Search ADVANCED SEARCH SCROLL This is a preprint and has not been peer reviewed. Data may be preliminary. 19 August 2025 V1 Latest version Share on Temperature mediates ROS metabolism and MAPK signaling pathways to influence Scutellaria baicalensis quality Authors : Yiming Lan , Yumu Shen , Yingxin Sun , Hongmei Lin [email protected] , and Limin Yang Authors Info & Affiliations https://doi.org/10.22541/au.175558198.80690692/v1 221 views 132 downloads Contents Abstract Information & Authors Metrics & Citations View Options References Figures Tables Media Share Abstract Extensively applied in traditional Chinese herbal medicine, S cutellaria baicalensis Georgi is a key botanical resource. The synthesis of its main active components, flavonoids, is significantly affected by temperature. In this study, two-year-old Scutellaria baicalensis were subjected to three temperature treatments: low temperature with high diurnal temperature variation (LH), low temperature with low diurnal temperature variation (LL), and high temperature with low diurnal temperature variation (HL). This investigation utilized multi-omics approaches (phenotypic, transcriptomic, metabolomic analyses) combined with biochemical assays to dissect the impacts of temperature regimes on growth traits, transcriptional regulation, metabolic biosynthesis, and pharmaceutical quality in Scutellaria baicalensis. The results showed that the HL treatment extended the root system but reduced the aboveground biomass. Transcriptomic analysis revealed that HL conditions upregulated MAPK signaling pathway - associated genes, thereby enhancing stress resistance, while gene expression patterns were comparable between the LH and LL groups. Metabolomics showed that under HL conditions, the accumulation of key flavonoid compounds such as baicalin was reduced, and the glycosylation process was hindered. Under HL conditions, reactive oxygen species (ROS) accumulated, malondialdehyde (MDA) content increased, superoxide dismutase (SOD) activity decreased, and peroxidase (POD) and catalase (CAT) activities significantly increased. Simultaneously, HL exposure elicited a significant accumulation of osmotic adjustment compounds (proline and soluble sugars), providing evidence for the oxidative stress adaptation strategy employed by Scutellaria baicalensis . These findings reveal the significant impact of temperature on the medicinal quality of Scutellaria baicalensis , and results indicate that reduced temperature conditions facilitate the growth of high - quality Scutellaria baicalensis , thereby establishing the theoretical foundation for achieving both superior quality and high yield in its cultivation. Temperature mediates ROS metabolism and MAPK signaling pathways to influence Scutellaria baicalensis quality Yiming Lan, Yumu Shen, Yingxin Sun, Mei Han, Hongmei Lin11 * Corresponding authors: Hongmei Lin ( [email protected] ); Limin Yang (yanglimin@ jlau.deu.cn), Limin Yang * College of Chinese Medicinal Materials, Jilin Agricultural University, Changchun 130118, China Abstract: Extensively applied in traditional Chinese herbal medicine, S cutellaria baicalensis Georgi is a key botanical resource. The synthesis of its main active components, flavonoids, is significantly affected by temperature. In this study, two-year-old Scutellaria baicalensis were subjected to three temperature treatments: low temperature with high diurnal temperature variation (LH), low temperature with low diurnal temperature variation (LL), and high temperature with low diurnal temperature variation (HL). This investigation utilized multi-omics approaches (phenotypic, transcriptomic, metabolomic analyses) combined with biochemical assays to dissect the impacts of temperature regimes on growth traits, transcriptional regulation, metabolic biosynthesis, and pharmaceutical quality in Scutellaria baicalensis. The results showed that the HL treatment extended the root system but reduced the aboveground biomass. Transcriptomic analysis revealed that HL conditions upregulated MAPK signaling pathway - associated genes, thereby enhancing stress resistance, while gene expression patterns were comparable between the LH and LL groups. Metabolomics showed that under HL conditions, the accumulation of key flavonoid compounds such as baicalin was reduced, and the glycosylation process was hindered. Under HL conditions, reactive oxygen species (ROS) accumulated, malondialdehyde (MDA) content increased, superoxide dismutase (SOD) activity decreased, and peroxidase (POD) and catalase (CAT) activities significantly increased. Simultaneously, HL exposure elicited a significant accumulation of osmotic adjustment compounds (proline and soluble sugars), providing evidence for the oxidative stress adaptation strategy employed by Scutellaria baicalensis . These findings reveal the significant impact of temperature on the medicinal quality of Scutellaria baicalensis , and results indicate that reduced temperature conditions facilitate the growth of high - quality Scutellaria baicalensis , thereby establishing the theoretical foundation for achieving both superior quality and high yield in its cultivation. Key words: Scutellaria baicalensis ; different temperature; RNA-seq; Antioxidant system; Medicinal material quality 1. Introduction Scutellaria baicalensis Georgi, a perennial herb in the Lamiaceae family, is widely cultivated in China (Shang et al., 2010; Zhao et al., 2016). Its dried roots, a traditional Chinese medicine with millennia-long application, exhibit notable anti-cancer (Chen et al., 2013; Gao et al., 2011), hepatoprotective (Yang et al., 2012), antibacterial, and antiviral activities. (Huang et al., 2023). The herb’s medicinal value primarily stems from its rich flavonoid content, including baicalein, wogonin, baicalin, and wogonoside (Li-Weber, 2009; Makino et al., 2008; Xu et al., 2020). As specified in the Pharmacopoeia of the People’s Republic of China, baicalin serves as the key marker for evaluating the quality of Scutellaria baicalensis medicinal materials (Commission Chinese Pharmacopoeia, 2020). In China, Scutellaria baicalensis has an extremely wide distribution. Its broad range allows it to grow in a variety of environments, indicating strong ecological adaptability. However, the regions widely recognized for producing high-quality Scutellaria baicalensis are mainly located in northern China, such as Hebei and Shanxi, as well as the high-altitude areas of Gansu in the west. Temperature represents a pivotal ecological factor governing plant growth and development, emerging as a dominant environmental determinant that impacts the yield and quality of numerous economically important crops (Battisti and Naylor, 2009; Lobell et al., 2011). Persistent adverse temperature conditions may exacerbate the impact of other biotic and abiotic stresses, triggering the outbreak of a series of diseases (Cohen and Leach, 2020; Desaint et al., 2021). The synergistic impact of these adverse factors significantly affects plant growth and development, resulting in diminished crop yield and quality (Zhao et al., 2017). Excessively high temperatures can alter the content and stability of many molecules in plants, disrupt the homeostasis of the cellular environment, and impair photosynthetic activity (Hikosaka et al., 2006; Moore et al., 2021). Additionally, elevated temperatures trigger excessive reactive oxygen species (ROS) accumulation, perturb plant hormone biosynthesis and signaling cascades, and thereby induce transcriptomic and metabolomic alterations (Devireddy et al., 2021; Singh et al., 2020). The biosynthesis of secondary metabolites in medicinal plants is intricately associated with temperature. Research evidence indicates that the impact of temperature increase on secondary metabolite upregulation or downregulation is determined by multiple contributing factors (Wang et al., 2025). High temperature can upregulate or downregulate related genes, affecting the accumulation of substances in plants (Liu et al., 2016). Flavonoids are plant metabolites with diverse structures. Widely present across plant kingdoms, they exert roles in multiple plant physiological processes (Wang et al., 2019). Temperature represents a crucial element influencing the biosynthetic process of flavonoids within plants. Research has demonstrated that high temperatures at 30°C cause a notable decrease in the levels of anthocyanins as well as the transcriptional activity of genes involved in the biosynthetic route (Fang et al., 2019). Moreover, flavonoids possess the capacity to eliminate free radicals and are of great significance in plants’ defense against ROS - induced damage (Schmidt et al., 2010). Despite being a critical ecological factor affecting medicinal herb quality, temperature’s regulatory effects on Scutellaria baicalensis have not been Scutellaria baicalensis . Therefore, this study used two-year-old Scutellaria baicalensis as the experimental material and cultivated it under different controlled temperatures to explore the effects of temperature on its quality and yield through the detection of growth and active components. Meanwhile, transcriptomics and metabolomics were employed to analyze the gene responses and metabolite changes of Scutellaria baicalensis under different temperatures. Combined with the ROS system, the molecular mechanism of temperature factors in the quality formation of Scutellaria baicalensis was preliminarily explored, providing a theoretical basis for the high-quality and high-yield production of Scutellaria baicalensis . 2. Materials and methods 2.1 Experimental design and treatments Scutellaria baicalensis used here was collected from Jilin Agricultural University’s Medicinal Plant Garden. After field sowing in spring 2022, one-year-old seedlings were harvested in 2023. Prior to sprouting in spring 2023, uniform one-year-old seedlings were chosen and planted in PVC pipes (40 cm × 7.5 cm). The pipes were buried in the soil to allow natural growth, with normal field management carried out during this period. The experiment was conducted during the peak accumulation period of active components in Scutellaria baicalensis . On August 30, 2023, Scutellaria baicalensis with similar growth were transferred to a light incubator for different temperature treatments. Three temperature treatments were established: LH (low temperature with high diurnal variation, 10°C–22°C), LL (low temperature with low variation, 10°C–18°C), and HL (high temperature with low variation, 17°C–25°C), representing suitable and unsuitable growth conditions for Scutellaria baicalensis . The temperatures in parentheses represent the night and day temperatures, respectively, with 12 hours each for day and night. Each treatment included 25 plants of Scutellaria baicalensis , and sampling was conducted after 30 days of treatment. 2.2 Determination of growth indices of Scutellaria baicalensis After harvesting, Scutellaria baicalensis samples were washed under running water, and the following parameters were measured: stem length, stem diameter, root length, root diameter, fresh root weight, and fresh shoot weight. Subsequently, a section was cut from the same position of the main root of each plant, pre-cooled in liquid nitrogen, and then stored in a -80°C freezer for subsequent biochemical, transcriptomic, and metabolomic analyses. The remaining parts were placed in an oven to dry at 45°C, then ground and sieved through a 100-mesh screen for storage. 2.3 Determination of antioxidant enzyme activities and osmotic adjustment substances in Scutellaria baicalensis The activities of superoxide dismutase (SOD, A001-1-1), peroxidase (POD, A084-3-1), catalase (CAT, A007-1-1), and the contents of malondialdehyde (MDA, A003-1-1), proline (PRO, A107-1-1), total soluble protein (TP, A045-2-2), and soluble sugar (SS, A145-1-1) were measured using detection kits from Nanjing Jiancheng Bioengineering Institute, following the manufacturer’s instructions. 2.4 Measurement of the content of active ingredients in Scutellaria baicalensis For each treatment, three replicate samples were prepared. Each sample (0.300 g of dried root powder) was extracted with 9 mL of 70% ethanol using the MARS 6 system at 80°C for 6 minutes. After the extraction process, filter the mixture. Dilute it five - fold with 70% ethanol and then pass it through a 0.22 - mm filter membrane for subsequent use. Baicalin, baicalein, wogonin, oroxylin A, scutellarin, scutellarein, and chrysin contents were determined by HPLC (Agilent 1260, USA). Chromatographic methods and standard curve details are provided in Tables S1 and S2. 2.5 Transcriptome sequencing RNA was retrieved from the roots of Scutellaria baicalensis. A Qubit fluorometer and a Qsep400 high - throughput biofragment analyzer were employed to assess and quantify the total RNA. The intact RNA was split into short - length RNA fragments. These fragments underwent reverse transcription to yield cDNA. The cDNA was purified and amplified by means of polymerase chain reaction (PCR) to complete the establishment of the cDNA library. Finally, the library preparations were sequenced on an Illumina Novaseq platform. Each treatment group was run in triplicate. After sequencing, raw data were obtained and filtered using FastQC 0.23.2 and Trimmatic 0.36.5, then assembled by HISAT2 2.1.0. Differentially expressed genes (DEGs) were identified with the DESeq2 package via negative binomial distribution testing, including count data normalization and FDR < 0.05 with |Log2 (Fold Change)| ≥ 1 as criteria. 2.6 Metabolomics analysis The freeze-dried sample underwent grinding at 30 Hz for 2 minutes using a high-speed grinder. Subsequently, 100 mg of the powdered sample was transferred to a centrifuge tube, to which 1.0 mL of 70% methanol-water solution was added. The mixture was vortexed at 4℃, centrifuged at 10,000×g for 10 minutes, and the resulting supernatant was collected for LC-MS analysis. Each experimental treatment was replicated three times biologically. Metabolite profiling was conducted using a UPLC-ESI-MS/MS system (Metware Biotechnology, China). Differentially accumulated metabolites (DAMs) were identified based on fold-change (≥ 2 or ≤ 0.5), p-value ≤ 0.05, and variable importance in the projection (VIP) ≥ 1. 2.7 Quantitative real-time PCR (qRT-PCR) To authenticate the transcriptomic findings, nine randomly selected differentially expressed genes (DEGs) underwent qRT-PCR validation. Gene-specific primers were designed using Primer Premier 5.0 (Table S3), with 18S rRNA serving as the endogenous control. The 20-μL reaction mixture consisted of 10 μL SYBR Premix Ex Taq (RR820, Takara, China), 1 μL each of forward and reverse primers, 1 μL cDNA template, and 7 μL ddH₂O. The thermal cycling protocol comprised 30 s at 95°C for pre-denaturation, followed by 40 cycles of 5 s at 95°C, 30 s at 56°C, and 15 s at 72°C. Relative gene expression was calculated using the 2 -ΔΔCt normalization method. 2.8 Statistical analysis Statistical analyses were conducted using IBM SPSS Statistics 24.0 (Armonk, NY, USA) via one-way ANOVA with a 95% confidence interval, designating p < 0.05 as the significance threshold. Data visualization was performed in GraphPad Prism 8.0, whereas bioinformatics analyses and figure generation utilized MetWare tools (http://www.metware.cn/). not-yet-known not-yet-known not-yet-known unknown 3. Results 3.1 Differences in phenotypic characteristics of Scutellaria baicalensis under different temperatures Fig. 1: Impacts of various temperatures on the growth of Scutellaria baicalensis . (A) Fresh weight of root, (B) Length of root, (C) Diameter of the thickest part of the root, (D) Total weight of aboveground parts (stem and leaves), (E) Length of stem, (F) Diameter of the thickest part of the stem. The letters presented in the figure signify significant differences among groups ( p <0.05). Under different temperature treatments, there were minor phenotypic differences in the roots of Scutellaria baicalensis , with no significant differences in root fresh weight and root diameter (Fig. 1). However, root length was longest under the HL treatment and shortest under the LH treatment. Conversely, more remarkable differences in the above - ground parts were observed among different treatments. The aerial part fresh weight peaked under LL treatment, showing a 53.3% increase compared to HL. Stem length and stem diameter were highest under the LH treatment, which were 26.9% and 18.2% higher than those under the HL treatment, respectively. 3.2 Transcriptome analysis of Scutellaria baicalensis To explore the effects of temperature on the transcriptional regulation of Scutellaria baicalensis , we performed transcriptome sequencing on Scutellaria baicalensis from the three treatments. A total of 61.56 Gb high-quality data were generated, averaging 6.84 Gb per sample. Q20 and Q30 rates were above 98.57% and 95.55%, respectively. The GC content was approximately 46%, and the overall sequencing error rate was lower than 0.03% (Table S4). Alignment to the reference genome efficiency was 95.90%–96.41%, and uniquely mapped reads represented 89.03%–90.18% (Table S5). PCA revealed that principal components PC1 and PC2 contributed 37.69% and 18.48% to sample variance, respectively. This indicated notable differences in the gene expression of Scutellaria baicalensis cultivated under the three temperatures (Fig. S1). Comparative analyses between LH and HL, LH and LL, and LL and HL identified 2,114 (895 upregulated and 1,219 downregulated), 650 (273 upregulated and 377 downregulated), and 2,091 (1,016 upregulated and 1,075 downregulated) differentially expressed genes (DEGs), respectively (Fig. 2A). A Venn diagram revealed the shared and unique DEGs among them. The most shared DEGs were between LH_vs_HL and LL_vs_HL, with 1,219 DEGs in common, while only 94 DEGs were shared among all three comparison groups (Fig. 2B). DEGs were enriched in KEGG pathways including Plant-pathogen interaction, MAPK signaling, secondary metabolite biosynthesis, and flavonoid biosynthesis, as shown by enrichment analysis (Fig. 2C). Overall, the DEGs between LL and LH were more similar, while HL showed a distinct difference from the other two, indicating that the transcriptional regulation of Scutellaria baicalensis grown under HL conditions was significantly different from that under the other two conditions. Fig. 2: Transcriptomic analysis of Scutellaria baicalensis . (A) Bar chart of DEGs. (B) Venn diagram of DEGs. (C) Enrichment of DEGs in KEGG pathways. Fig. 3: KEGG circle plots of DEGs. (A) KEGG pathways containing DEGs from LH_vs_HL. (B) KEGG pathways containing DEGs from LL_vs_HL. Based on the KEGG circle plots of DEGs from LH_vs_HL and LL_vs_HL (Fig. 3), it can be observed that the KEGG pathways enriched by DEGs in the two groups are highly similar. A great many genes are concentrated in ko01110 (Biosynthesis of secondary metabolites), ko04075 (Plant hormone signal transduction), and ko04016 (MAPK signaling pathway - plant), ko04626 (Plant - pathogen interaction). By examining the proportion of red and blue in the third circle of the figure, it can be seen that the proportion of downregulated DEGs in ko04626, ko04075, and ko04016 is relatively high in both groups, indicating that the gene expression in these three KEGG pathways is lower in LH and LL compared to HL. not-yet-known not-yet-known not-yet-known unknown Fig. 4: DEGs enriched in MAPK signaling pathway - plant (map04016), Plant hormone signal transduction (map04075), and Plant-pathogen interaction (map04626). (A) Network and Venn diagrams of DEGs. (B) Heatmap of DEGs enriched in the MAPK signaling pathway – plant. The network graph of DEGs within the three aforementioned KEGG pathways was constructed (Fig. 4A). Results indicated substantial overlap between DEGs in the MAPK signaling pathway (map04016) and those in the Plant hormone signal transduction (map04075) and Plant-pathogen interaction (map04626) pathways. Specifically, 7 DEGs were present in all three pathways. According to the heatmap of DEGs in map04016 (Fig. 4B), it was found that the genes appearing in map04626 were significantly higher under HL treatment than LL and LH, while most DEGs appearing in map04075 were lower under HL treatment than LL and LH. 3.3 Metabolomics analysis of Scutellaria baicalensis Fig. 5: Metabolomics analysis of Scutellaria baicalensis . (A) Correlation analysis among samples. (B) Analysis of the proportion of metabolite composition. (C) Venn diagram of DAMs in each comparison group. (D) Principal component analysis of samples. A total of 1,620 compounds were detected in the metabolomics analysis, among which 612 were differentially accumulated metabolites (DAMs). The correlation analysis among samples indicated good biological replication within each group (Fig. 5A). Based on the correlation coefficients between different groups, it can be observed that the LL and LH groups had a stronger correlation, while their correlation with the HL group was weaker. This suggests that the metabolites in the former two groups were more similar, whereas HL exhibited greater differences. PCA analysis also confirmed the above results (Fig. 5D). The detected metabolites were mainly divided into 11 categories, with flavonoids accounting for the highest proportion (33.11%), followed by phenolic acids (14.27%) (Fig. 5B). The Venn diagram results showed that there were 83 DAMs shared between LL_vs_HL and LH_vs_HL, while only 10 DAMs were common among all three groups (Fig. 5C). This result also indicates significant differences among the three comparison groups, with the DAMs in LL and LH being more similar. Fig. 6: (A) Enrichment of DAMs in KEGG pathways. (B) DAMs enriched in the Flavonoid biosynthesis pathway. KEGG pathways including Cyanoamino acid metabolism and Glucosinolate biosynthesis were where the DAMs were mainly enriched (Fig. 6A). DAMs were also significantly enriched in Phenylpropanoid and Flavonoid biosynthesis, the key pathways for flavonoid compound biosynthesis and Scutellaria baicalensis active ingredient production. Flavonoids were the most abundant compounds in the detection results. By comparing the DAMs in the Flavonoid biosynthesis pathway, it was found that the LH group had a greater variety of substances with higher content, while the HL group had fewer (Fig. 6B). In addition, the LH group had higher content of compounds of the Amino acids and derivative type, while the HL group had higher content of Saccharides. 3.4 Influence of temperature upon the antioxidant and osmotic adjustment systems within Scutellaria baicalensis Fig. 7: Changes in the antioxidant and osmotic adjustment systems in Scutellaria baicalensis . The bar graphs display the amounts of antioxidant enzymes and osmotic adjustment substances, and the heat map presents the expression levels of genes associated with the synthesis of antioxidant enzymes. In the antioxidant system of Scutellaria baicalensis , the activities of the protective enzymes varied among different treatments (Fig. 7). SOD activity was significantly lower in the HL treatment than in the LL and LH treatments, whereas POD and CAT activities were notably higher in HL compared to the other two treatments. Looking at the genes responsible for the synthesis of these three enzymes, it can be seen that the genes for SOD synthesis were downregulated in HL, while most of the genes for POD and CAT synthesis were upregulated. Enzyme activities correlated with the expression patterns of their encoding genes. For osmotic adjustment, PRO and SS levels in HL were significantly elevated relative to LL/LH. TP content did not differ from LL but was higher than LH, whereas HL exhibited significantly higher MDA content than the other treatments. 3.5 Effects of temperature on the quality of Scutellaria baicalensis medicinal materials Fig. 8: Measurement of the Contents of Seven Active Ingredients in Scutellaria baicalensis via HPLC The contents of active components such as baicalin in Scutellaria baicalensis were quantitatively determined by HPLC (Fig. 8). In the HL condition, baicalin, wogonoside, and scutellarin contents were significantly lower than those in the LH condition. However, the contents of baicalein, wogonin, scutellarein, and oroxylin A showed an inverse situation. Overall, the contents of glycosylated compounds with aglycones were lower in HL than in LH to varying degrees, while their precursor substances showed the opposite trend. Fig. 9: Baicalin biosynthetic pathway-related major metabolite contents and catalytic enzyme gene expression levels. The heatmap modules in the figure, from left to right, represent LL, HL, and LH processing respectively. Baicalin, one of the primary active ingredients in Scutellaria baicalensis , is also a crucial compound that determines its quality. Combining the results of transcriptomics, metabolomics, and HPLC detection, the gene expression levels and compound contents in the baicalin biosynthetic pathway were analyzed (Fig. 9). Within this pathway, enzyme-encoding gene expression was lowest in HL and highest in LH for most genes. Phenylalanine was detected in the metabolomics with lower content in HL, while the following three compounds were not detected in the metabolomics. The contents of pinocembrin, chrysin, and baicalein were higher in HL, while that of baicalin was lower. 3.6 Verification of gene expression via qRT - PCR Fig. 10: Validation of DEGs from the transcriptome by qRT-PCR. qRT-PCR analysis was performed on nine randomly selected DEGs to validate the RNA-seq results (Fig. 10). The outcomes of the nine chosen genes in RNA - seq and qRT-PCR were strikingly alike, suggesting the dependability of the RNA-seq data. 4. Discussion 4.1 Influence of temperature upon the growth of Scutellaria baicalensis Temperature exerts a significant effect on plant root growth. Studies have shown that high temperature enhances root metabolism and enzyme activity, promoting elongation, while low temperature inhibits cell division and elongation (Fuellner et al., 2012; Luo et al., 2020). The notable discrepancy in root length across temperature gradients in this investigation reinforces this inference. By contrast, intergroup comparisons revealed no statistically significant variations in root fresh mass or diameter. This might be attributed to the fact that this stage was already the harvest period of Scutellaria baicalensis , with the obvious changes in the roots having weakened and the focus mainly shifting to internal alterations. Since stems and leaves are directly exposed to the air, while roots have soil as a buffer, the aboveground parts are more sensitive to temperature and more affected than the roots. The aboveground fresh weight was highest in LL and lowest in HL, probably because high temperature accelerates respiration, consuming more photosynthetic products and thus reducing biomass accumulation (Zhu et al., 2021). A larger diurnal temperature variation may promote an increase in internode length and stem thickness by affecting hormones such as gibberellins, which in turn leads to the highest stem length and stem diameter in LH (Hu et al., 2019; Myster and Moe, 1995). 4.2 Transcriptional Regulation of MAPK in Scutellaria baicalensis under Different Temperatures Within plant biology, MAPK plays a critical role across diverse processes, especially in modulating responses to temperature shifts (Li et al., 2025; Mo et al., 2021). Transcriptomic data showed that plant-pathogen interaction-related genes were significantly upregulated in the HL treatment group. (Figure 4). Likely, the high temperature enhanced the quantity of soil pathogens, which in turn induced a succession of disease - resistance reactions in Scutellaria baicalensis . (Delgado-Baquerizo et al., 2020). FLS2/BAK1-FRK1-WRKY33, as a key signaling pathway for plants to respond to external stimuli, is activated at high temperatures. FLS2 forms a complex with BAK1 to initiate the immune signal cascade, stimulating the downstream FRK1 receptor kinase and regulating WRKY33, which is involved in antifungal defense (Zou et al., 2018). On the other hand, BAK1 can form a complex with OST1 to regulate ABA signaling and can also bind to FRK1 to enhance defense responses (Shang et al., 2020). In the HL group, genes involved in this pathway were significantly upregulated, as shown in this study, probably because the relatively higher temperature is more conducive to the reproduction of various pathogens in the soil, thereby stimulating the disease resistance response of Scutellaria baicalensis (Marquez et al., 2021; Saremi et al., 1999). PYL is an ABA receptor that forms a complex with PP2C upon binding to ABA, inhibiting the activity of PP2C and thereby activating ABA signaling (Wasilewska et al., 2008). Additionally, PYL directly modulates MYC2, linking the ABA and JA signaling pathways. (Aleman et al., 2016). In the HL environment, PP2C-related gene downregulation and PYL-related gene upregulation activate the ABA pathway in Scutellaria baicalensis . Concurrently, ERF1 gene upregulation and MYC2 gene downregulation suggest antagonistic ERF-MYC2 regulation of plant stress responses. Studies show MYC2 directly binds to the ERF1 promoter, repressing its expression (Li et al., 2024). 4.3 The Effect of Temperature on the Metabolites of Scutellaria baicalensis The metabolomics analysis revealed that flavonoids are the predominant metabolites in Scutellaria baicalensis (Fig 5), and they also constitute its primary active components. The biosynthesis of plant flavonoids is significantly regulated by temperature (Li et al., 2025). Studies indicate that elevated temperatures lead to a substantial decrease in anthocyanin accumulation and suppress the expression of biosynthetic pathway genes (Lin-Wang et al., 2011). In the flavonoid biosynthesis pathway, LH significantly elevated amino acid levels, whereas HL increased sugar contents (Fig. 6). As precursors for flavonoid synthesis, amino acids supply abundant raw materials for subsequent compound production. Sugars, on the other hand, help maintain cell osmotic potential, and their increased content can assist cells in withstanding adverse external environments (Hasanuzzaman et al., 2013). 4.4 Influence of Temperature upon the Antioxidant System of Scutellaria baicalensis According to Medina et al. (2021), elevated temperature profoundly accelerates reactive oxygen species (ROS) biosynthesis in plant cells, resulting in oxidative stress induction. MAPK involvement accompanies ROS generation. As shown by Montillet et al. (2013), hydrogen peroxide (H₂O₂) functions upstream of the MAPK cascade and contributes to ABA-mediated antioxidant protection. Under high temperature, the accumulated ROS in plant cells can cause lipid peroxidation, leading to an increase in MDA and subsequent cell membrane damage(Wang et al., 2024). At this point, the plant antioxidant system synthesizes protective enzymes to clear ROS from the plant body, preventing ROS toxicity. This investigation revealed that HL exposure induced a significant upregulation of POD and CAT activities, while concurrently suppressing SOD activity compared to LL and LH light treatments (Fig. 7). The observed divergence in SOD activity may be attributed to transcriptional repression of SOD-encoding genes under elevated temperatures. As demonstrated by Wang et al. (2017), SOD exhibits inherent thermolability, resulting in rapid degradation at high temperatures. Moreover, findings from Hasanuzzaman et al. (2019) suggest a shift in ROS detoxification pathways, where high-temperature conditions promote an ascorbate-glutathione cycle- and CAT-driven mechanism, thereby reducing the reliance on SOD-mediated scavenging. High temperature increases the contents of PRO, TP, and SS in plants, thereby maintaining cellular homeostasis and alleviating oxidative damage (Fig. 7). Additionally, PRO functions as an antioxidant among these substances, directly eliminating ROS to alleviate oxidative injury (Hasanuzzaman et al., 2013). Heat shock proteins (HSPs), a group of plant proteins sensitive to temperature, are highly induced under both high and low temperatures to maintain cellular balance. Therefore, the content of TP is relatively high in both LL and HL treatments. 4.5 Molecular Mechanisms Concerning the Influence of Temperature on the Formation of Scutellaria baicalensis Quality Fig. 11: Molecular mechanisms of temperature effects on the quality formation of Scutellaria baicalensis . Flavonoids serve as a crucial material foundation for the pharmacological activities of Scutellaria baicalensis and are also a significant criterion for evaluating its quality. Baicalin, among them, is one of the most vital chemical constituents. Zhao et al. (2016) showed that phenylalanine is transformed into baicalin in Scutellaria baicalensis roots through the phenylpropanoid and flavonoid biosynthetic pathways. Using metabolomics and HPLC techniques, this study quantified baicalin, phenylalanine (the starting material of its synthetic pathway), and intermediate metabolites pinocembrin, chrysin, and baicalein (Fig. 9). The contents of the starting material phenylalanine and the final product baicalin were low in HL, while the contents of the intermediate products were high. However, the expression of all related genes was low. This may be because the last step of glycosylation is significantly affected by temperature, and the process is hindered, leading to the accumulation of intermediate products. The active components in Scutellaria baicalensis are not limited to baicalin; other chemical components such as baicalein also have important functions. Through the detection of the remaining chemical components, we found that the contents of glycosylated components were significantly reduced in HL, while the contents of their non-glycosylated precursors were significantly increased (Fig. 8). This indicates that the activity of glycosyltransferase in this process is significantly regulated by temperature and shows a downward trend at high temperatures. This phenomenon is also observed in tomatoes and Arabidopsis(Alhaithloul et al., 2021; Martz et al., 2009). According to Zhang et al. (2023), freezing following harvest facilitates the transformation of glycosides into aglycones in Scutellaria baicalensis , with a possible parallel effect under high-temperature conditions. Integrating physiological, transcriptomic, and metabolomic analyses indicates that temperature affects the quality of Scutellaria baicalensis by regulating the MAPK transcriptional control and ROS system in Scutellaria baicalensis (Fig. 11). Studies have shown that soil microorganisms significantly affect the quality of Scutellaria baicalensis , especially the increase in stress-resistant microorganisms, which can significantly reduce the flavonoid compounds in Scutellaria baicalensis (Yang et al., 2024). Under HL treatment, the MAPK transcriptional regulation was significantly activated, enhancing the disease resistance of Scutellaria baicalensis to counteract the increase in soil pathogens at high temperatures. Meanwhile, the accumulated ROS in the body stimulated the entire protective enzyme system, changing the clearance strategy from indirect removal of O 2- to direct removal, thereby increasing the efficiency of ROS clearance(Hasanuzzaman et al., 2019). In addition, osmotic adjustment substances significantly increased to maintain cell osmotic potential, and PRO also participated in ROS clearance, reducing the toxicity of ROS to cells. Through internal adjustments, Scutellaria baicalensis ensured its survival under HL conditions, which in turn led to the downregulation of gene expression in the baicalin biosynthetic pathway, ultimately affecting the formation of its quality. 5. Conclusion Our findings highlight the profound influence of temperature on the medicinal quality of Scutellaria baicalensis , with lower temperature conditions (LH and LL) being more conducive to the accumulation of key medicinal compounds such as baicalin, while high temperature (HL) may reduce medicinal value by activating stress responses and altering metabolic pathways. In particular, the enhancement of plant defense mechanisms under high temperature conditions (such as the MAPK signaling pathway and ROS system) comes at the expense of flavonoid synthesis. These results underscore the pivotal role of temperature in determining the yield and quality of Scutellaria baicalensis , elucidating the preference for northern China as its optimal growth region. Therefore, temperature conditions should be emphasized in the selection of cultivation sites for Scutellaria baicalensis , with priority given to areas with lower temperatures and relatively larger diurnal temperature variations. This not only helps to increase the aboveground biomass but also ensures the accumulation of medicinal components in the roots, thereby achieving the goal of high yield and high quality. CRediT authorship contribution statement Yiming Lan: Writing – original draft, Methodology, Investigation, Data curation, Visualization. Yumu Shen: Investigation, Validation. Yingxin Sun: Investigation. Hongmei Lin : Writing – review & editing, Supervision, Supervision, Funding acquisition, Conceptualization. Mei Han: Writing – review & editing, Funding acquisition, Project administration. Limin Yang: Funding acquisition, Project administration. Funding This study was supported by the Key Research and Development Program of Jilin Provincial Science and Technology Development Plan (20240305001YY); Declaration of Competing Interest The authors declare no competing financial interests. not-yet-known not-yet-known not-yet-known unknown Acknowledgment This study was supported by the Key Research and Development Program of Jilin Provincial Science and Technology Development Plan (20240305001YY); Data availability The raw sequence data reported in this paper have been deposited in the Genome Sequence Archive (Genomics, Proteomics & Bioinformatics 2021) in National Genomics Data Center (Nucleic Acids Res 2025), China National Center for Bioinformation / Beijing Institute of Genomics, Chinese Academy of Sciences (GSA: CRA028578) that are publicly accessible at https://ngdc.cncb.ac.cn/gsa. 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Information & Authors Information Version history V1 Version 1 19 August 2025 Copyright This work is licensed under a Non Exclusive No Reuse License. Keywords scutellaria baicalensis antioxidant system different temperature medicinal material quality metabalome transcriptome Authors Affiliations Yiming Lan Jilin Agricultural University View all articles by this author Yumu Shen Jilin Agricultural University View all articles by this author Yingxin Sun Jilin Agricultural University View all articles by this author Hongmei Lin [email protected] Jilin Agricultural University View all articles by this author Limin Yang Jilin Agricultural University View all articles by this author Metrics & Citations Metrics Article Usage 221 views 132 downloads .FvxKWukQNSOunydq8rnd { width: 100px; } Citations Download citation Yiming Lan, Yumu Shen, Yingxin Sun, et al. Temperature mediates ROS metabolism and MAPK signaling pathways to influence Scutellaria baicalensis quality. Authorea . 19 August 2025. 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