Table of Content

28 August 2026, Volume 46 Issue 5

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Review Article



Molecular Mechanisms of Hormone Regulatory Networks and Biosynthesis of Triterpene Saponins in Araliaceae Plants

Xiaotong LIU, Huiting ZHANG, Shilong LIU, Rui GUO, Zheng SHI, Zongqi YANG, Zheyong XUE, Xin HUA

2026, 46(5):  777-790.  doi:10.7525/j.issn.1673-5102.2026.05.001

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Triterpenoid saponins in medicinal plants of the Araliaceae family are important secondary metabolites and key determinants of medicinal quality and pharmacological activity. Their biosynthesis is coordinately regulated by multiple plant hormones and interconnected signaling networks. Jasmonic acid， salicylic acid， auxin， gibberellin， cytokinin， and ethylene can activate or repress transcription factors such as MYC2， WRKY， AP2/ERF， MYB， and bHLH， thereby forming complex regulatory networks that precisely control the expression of key structural genes involved in triterpenoid saponin biosynthesis through synergistic or antagonistic interactions. Current studies indicate that jasmonic acid is the principal positive regulator promoting triterpenoid saponin accumulation， whereas salicylic acid preferentially regulates the downstream modification steps of saponin biosynthesis. In contrast， growth- and development-related hormones mainly influence saponin accumulation indirectly by modulating the growth-defense trade-off. Crosstalk among different hormonal signals further enhances the adaptability of Araliaceae plants to environmental changes and shapes the dynamic patterns of triterpenoid saponin accumulation. This review summarized recent advances in the regulation of triterpenoid saponin biosynthesis by plant hormones in medicinal plants of the Araliaceae family， with the aim of providing a theoretical basis for elucidating the mechanisms underlying active compound accumulation， improving the quality of medicinal plants， and achieving efficient biosynthesis of triterpenoid saponins.



Research Progress on Catalytic Mechanism, Multidimensional Regulation of Squalene Epoxidase in Terpenoid Biosynthesis Pathway

Di ZHAO, Yongchang XUE

2026, 46(5):  791-801.  doi:10.7525/j.issn.1673-5102.2026.05.002

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Terpenoids（TPDs） are of natural products with diverse structures and wide-ranging functions in nature. Their derivatives such as artemisinin， paclitaxel and ginsenosides exhibit anti-inflammatory，anti-cancer， and immunomodulatory activities， and possess significant application value in medicine and agriculture. Squalene epoxidase（SQE） is a key rate-limiting enzyme in TPD biosynthesis. It catalyzes the oxidation of squalene to 2，3-oxidosqualene， which is the core branch node in TPD biosynthesis. It is of great significance to regulate the diversity of terpenoid secondary metabolites. With advances in synthetic biology， SQE research has become increasingly diverse and challenging， broadening its application prospects across various plant species. This paper comprehensively analyzed the biosynthesis pathway， SQE structures and catalytic mechanisms， multi-dimensional regulation and TPD applications. This study aimed to elucidate the key roles of SQE in the TPD biosynthesis pathway， and provide new insights for the subsequent genetic engineering operation of SQE， and the development of plant secondary metabolites.



Ecological Mechanisms of Sexual Dimorphism-Driven Secondary Metabolic Differentiation in Dioecious Medicinal Plants and Their Implications for Medicinal Material Quality

Ting XIE, Jian GENG, Haojie GAO, Zeping WANG, Juan CHEN, Zhichao XIA

2026, 46(5):  802-812.  doi:10.7525/j.issn.1673-5102.2026.05.003

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Sexual dimorphism in dioecious plants reflects divergent reproductive strategies and profoundly influences resource allocation and secondary metabolism. However， the effects of sexual dimorphism on the chemical composition， medicinal quality， and underlying ecological mechanisms of medicinal plants remain insufficiently understood. This review summarized recent advances regarding sexual dimorphism in the secondary metabolites of dioecious medicinal plants. This paper focused on the ecological and molecular mechanisms driving sex-specific metabolic differentiation， including resource allocation strategies， phytohormone regulation， transcriptional networks， and environmental interactions. Furthermore， this paper explored the implications of plant sex for medicinal material quality control and the formation of bioactive constituents. Current evidence indicates that plant sex significantly influences the composition and accumulation of diverse secondary metabolites， promoting sex-specific chemotypes through coordinated regulation of resource acquisition and allocation， environmental adaptation， and metabolic reprogramming. Plant sex represents a critical biological factor influencing medicinal quality and contributing to chemical variation in medicinal materials. However， sex-related chemical variations exhibit strong species-specificity and environmental dependence， and their direct effects on pharmacological efficacy and clinical applications lack sufficient empirical support. Future research should foster interdisciplinary integration among plant ecology， chemical ecology， plant metabolic biology， and pharmaceutical sciences. By combining multi-omics approaches， long-term ecological observations， and standardized bioactivity evaluations， future studies should establish a conceptual framework linking plant sex， resource acquisition and allocation， secondary metabolic networks， and medicinal quality. Such efforts will elucidate the ecological mechanisms underlying sex-driven medicinal quality formation and provide theoretical foundations for the precise utilization of dioecious medicinal plant resources， quality control of traditional Chinese medicines， and their modernization.

Original Paper



Multidimensional Analysis of the Dynamic Evolution of Volatile Components during the Formation of ‘Worm-Like Rings’ in Notopterygium incisum and Identification of Key Markers

Wenlong ZHAO, Jiayi CAO, Yulu NIU, Jie GAO, Jinbao ZHANG, Honggang CHEN, Wen LI, Ling JIN

2026, 46(5):  813-826.  doi:10.7525/j.issn.1673-5102.2026.05.004

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To investigate the dynamic changes in volatile organic compounds（VOCs） during the formation of ‘worm-like rings’ in annual Notopterygium incisum seedlings and identify key odor markers， this study employed an integrated analytical strategy. This approach combined electronic nose technology with headspace solid-phase microextraction coupled with gas chromatography-mass spectrometry（HS-SPME-GC-MS） to profile N. incisum plants multidimensionally before and after ‘worm-like rings’ formation. Electronic nose analysis revealed that the characteristic odor of N. incisum primarily originated from the rhizome. Response values of sensors sensitive to terpenes， such as W5S， increased significantly after the formation of ‘worm-like rings’， indicating that the characteristic odor is predominantly generated concomitantly with ‘worm-like rings’ development， with terpenes serving as a key material basis. GC-MS analysis further identified terpenoids as the main components constituting the characteristic odor. Combined with relative odor activity value（ROAV） analysis， significantly upregulated compounds after ‘worm-like rings’ formation were identified. Notably， ethyl 2-propenoic acid， ethyl ester and propane， 2，2'-［1，2-ethanediylbis（thio）］bis［2-methyl-， jointly contributed to the pungent odor. Meanwhile， terpenoids including eucalyptol and butanal， 2-methyl-， along with esters and aromatic compounds， provided herbal-floral and woody aromas that balanced the overall odor profile. The strong correlation between electronic nose responses and GC-MS data validated the reliability of electronic nose for rapidly monitoring complex VOC dynamics. This study clarifies that the formation of ‘worm-like rings’ in N. incisum represents a critical juncture for the emergence of its characteristic odor， elucidates its material basis and key odor markers， and provides a scientific foundation for understanding the odor formation mechanism and quality evaluation of N. incisum.



Differential Analysis of Volatile Compounds in Different Organs of Apocynum venetum Based on HS-SPME-GC-MS

Xueqing ZHAO, Li JIANG, Bing ZHAO, Rong CONG, Xiaoxu DONG

2026, 46(5):  827-837.  doi:10.7525/j.issn.1673-5102.2026.05.005

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To clarify the differences in composition and aroma characteristics of volatile substances among different tissues and organs of Apocynum venetum and to elucidate the key sites and material basis for floral scent emission， this study used the stamens， petals， receptacles， stems， and leaves of A. venetum as experimental materials. Volatile compounds from these tissues and organs were extracted， identified， and quantitatively analyzed using headspace solid-phase microextraction coupled with gas chromatography-mass spectrometry（HS-SPME-GC-MS）. A systematic comparison was performed by integrating principal component analysis（PCA）， hierarchical cluster analysis（HCA）， and aroma profile analysis. The results revealed a total of 55 major volatile compounds， which were classified into eight chemical categories： alcohols（8 compounds）， aldehydes（11 compounds）， ketones（4 compounds）， esters（5 compounds）， alkanes（2 compounds）， siloxanes （5 compounds）， terpenes（16 compounds）， and others（4 compounds）. Among them， alcohols and ketones constituted the highest proportions in stamens and petals， with phenylethanol and acetophenone as the predominant volatiles. Aldehydes exhibited the highest relative content in the receptacle（22.58%）， primarily consisting of （2E）-2-nonenal and （2E， 6Z）-2，6-nonadienal. Terpenes accounted for 56.60% of the volatile compounds in stems， whereas alcohols（44.51%） and terpenes（21.16%） were dominant in leaves. PCA and HCA indicated a significant divergence in volatile profiles between reproductive organs（floral tissues） and vegetative organs （stems and leaves）， with stamens and petals showing the highest similarity. Aroma analysis further demonstrated that floral tissues were the primary sites responsible for the emission of A. venetum floral scent， which was characterized predominantly by floral and fruity notes， whereas stems and leaves exhibited woody and herbal aroma tones. In summary， the volatile composition in different tissues and organs of A. venetum displays pronounced organ-specific patterns. These findings provide a scientific foundation for understanding the mechanisms underlying floral fragrance biosynthesis and for the targeted utilization of its various organs.



Cloning, Expression Analysis and Functional Verification of the PrGSTF12 Gene from Paeonia rockii

Sitong CHEN, Sha LUO, Qianru ZHANG, Jiahui LI, Chen TAO, Qianqian SHI

2026, 46(5):  838-850.  doi:10.7525/j.issn.1673-5102.2026.05.006

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Glutathione S-transferase（GST） genes play a crucial role in plant anthocyanin accumulation. To clarify the roles of GSTs in anthocyanin transport and accumulation in the petal blotches of Paeonia rockii， color indices， flavonoid components and contents were determined in petals at different developmental stages. The results indicated that peonidin and cyanidin were the major pigments responsible for the formation of petal blotches. Subsequently， a differentially expressed GST gene， PrGSTF12， was identified and cloned from the petal transcriptome of P. rockii. Subcellular localization， expression analysis， and virus-induced gene silencing（VIGS） were performed to investigate its biological function. Phylogenetic analysis revealed that PrGSTF12 had a close genetic relationship with known GSTs involved in anthocyanin transport. Subcellular localization assays indicated that PrGSTF12 was targeted to the tonoplast， suggesting its function in this cellular compartment. Quantitative real-time PCR analysis showed that the expression level of PrGSTF12 was significantly higher in the basal blotch site than that in the non-blotch site of petals， and its expression level was positively correlated with local anthocyanin accumulation. The VIGS assay demonstrated that silencing PrGSTF12 resulted in noticeable fading of the petal basal blotches， accompanied by a significant reduction in anthocyanin content. Meanwhile， the expression levels of anthocyanin synthesis-related genes， including PrCHS， PrCHI， PrF3H， PrF3'H， PrDFR and PrANS， were markedly decreased. In summary， these results demonstrate that PrGSTF12 plays important roles in anthocyanin transport and accumulation in the petal basal blotches of P. rockii and contributes to basal blotch formation.



Genome-wide Identification of WRKY Transcription Factors and Key Genes for Abscisic Acid-mediated Flavonoid Synthesis in Sour Jujube (Ziziphus jujuba var. spinosa)

Chuxing DONG, Mengyuan ZHANG, Xinbo SHI, Gang ZHANG, Zhongxing SONG, Nan WANG, Honghong JIAO

2026, 46(5):  851-865.  doi:10.7525/j.issn.1673-5102.2026.05.007

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WRKY transcription factors play important roles in flavonoid biosynthesis and plant growth and development； however， their functional roles in the medicinal plant sour jujube （Ziziphus jujuba var. spinosa） remain largely unexplored. In this study， we conducted a comprehensive genome-wide identification of the WRKY gene family in sour jujube using bioinformatics approaches. We systematically analyzed their phylogenetic relationships， chromosomal localization， conserved domains， and cis-acting elements in promoter regions. Furthermore， we performed exogenous abscisic acid（ABA） treatment on sour jujube seeds， combined with flavonoid content determination and gene expression analysis， to screen candidate genes significantly associated with flavonoid accumulation. The tissue-specific expression patterns of these candidate genes were subsequently validated by quantitative real-time PCR. A total of 47 ZjWRKY genes were identified in the sour jujube genome， which were classified into three subfamilies（Group Ⅰ， Group Ⅱ， and Group Ⅲ） based on phylogenetic analysis. The ZjWRKY genes were unevenly distributed across 11 chromosomes， and all encoded proteins contained the conserved WRKYGQK domain， predominantly localized in the nucleus. Notably， 89% of ZjWRKY genes contained ABA-responsive elements（ABREs） within their promoter regions. Exogenous ABA treatment significantly promoted flavonoid accumulation in germinated sour jujube seeds， and the flavonoid content was correlated with the expression patterns of 12 ZjWRKY genes. Among them， ZjWRKY10， ZjWRKY30， and ZjWRKY45 showed a strong correlation with flavonoid content and were predominantly and highly expressed in seeds and pulp tissues. These results suggest that the three ZjWRKY genes may act synergistically in regulating fruit and seed development as well as ABA-induced flavonoid biosynthesis in sour jujube. This study provides important clues for further elucidating the ABA signaling-WRKY transcription factor-flavonoid biosynthesis regulatory network.



High-concentration CO₂ Treatment Inhibits the Accumulation of Anthocyanins in the Bulbs of Lilium davidii var. unicolor

Wenguang FAN, Hengyu XIAN, Peng BAI, Bolu SUN, Haiwei REN, Yaqin TIAN

2026, 46(5):  866-879.  doi:10.7525/j.issn.1673-5102.2026.05.008

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The aim of this study was to explore the physiological effects and molecular mechanisms of high-concentration CO₂-induced inhibition on the purple-red coloration in Lilium davidii var. unicolor， clarifying its regulatory effects on anthocyanin accumulation and related metabolic pathways. Bulbs were used as experimental materials and subjected to a controlled atmosphere treatment with 15% CO₂， 10% O₂， and 75% N₂. Bulb surface color parameters， anthocyanin content， metabolite（e.g. soluble protein） levels， and oxidase activities were determined after 0， 3， 6， and 9 days of storage. Transcriptome sequencing was further performed， and differentially expressed genes were screened through GO and KEGG enrichment analyses. The expression of key genes was validated by qRT-PCR. The results showed that high-concentration CO₂ significantly delayed the deepening of purple-red coloration during storage， maintained higher L* values， and markedly inhibited anthocyanin accumulation. At the physiological level， high-concentration CO₂ reduced the activities of polyphenol oxidase（PPO） and peroxidase（POD）， indicating its inhibitory effects on both enzymatic browning and anthocyanin biosynthesis. Transcriptomic analysis showed that the differentially expressed genes were mainly enriched in the phenylpropanoid biosynthesis and flavonoid biosynthesis pathways. Most key structural genes associated with anthocyanin biosynthesis were downregulated under high-concentration CO₂ treatment. These findings indicate that high-concentration CO₂ inhibits anthocyanin accumulation by suppressing gene expression in the phenylpropanoid and flavonoid biosynthetic pathways， thereby delaying the development of purple-red coloration and improving the postharvest storage quality of the bulbs. This study provides a theoretical basis for the development of controlled-atmosphere preservation technology for Lilium davidii var. unicolor.



Effects of Methyl Jasmonate on Growth, Bioactive Metabolites Accumulation and Key Genes Expression in Suspension Cells of Angelica sinensis

Huaiyi MA, Dan QIU, Yidi WANG, Mengfei LI

2026, 46(5):  880-888.  doi:10.7525/j.issn.1673-5102.2026.05.009

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To screen an optimal methyl jasmonate （MeJA） concentration suitable for growth and primary bioactive metabolite accumulation of Angelica sinensis suspension cells， the cell biomass， primary bioactive metabolite contents， in vitro antioxidant activities， and expression levels of genes related to ferulic acid and flavonoids biosynthesis were measured to the suspension cells inoculated into the liquid medium containing different concentrations of MeJA（CK， 25， 50， 100， 150， and 200 μmol⋅L^-1） for 9 days. The results showed that the cell growth was enhanced at a lower concentration of MeJA（25 μmol⋅L^-1） while inhibited at higher concentrations. The contents of primary bioactive metabolites （ferulic acid， soluble sugar， total flavonoids and phenolic） and in vitro antioxidant activities showed increasing trends with an increase in MeJA concentration， while the values tended to stabilize or decrease at the higher MeJA concentrations（150-200 μmol⋅L^-1）. Compared with CK， the expression levels of genes（e.g.， PAL2， 4CLL6， and HCT4） related to ferulic acid biosynthesis were enhanced at lower concentrations of MeJA and inhibited at higher concentrations； whereas the expression levels of genes（e.g.， CHS2， CHI2， and F3H-3） related to flavonoids biosynthesis were enhanced at the higher MeJA concentrations. These findings provide references for regulating suspension cell growth and metabolite accumulation of A. sinensis by using elicitors.



Effects of Different Culture Modes on Growth and Bioactive Metabolite Accumulation in Callus of Fritillaria cirrhosa

Jingyi KE, Mengfei LI

2026, 46(5):  889-898.  doi:10.7525/j.issn.1673-5102.2026.05.010

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The aim of this study was to screen an optimal culture method for the growth and bioactive metabolite accumulation of Fritillaria cirrhosa D. Don embryogenic callus， thereby providing a reference for the rapid proliferation and efficient enrichment of its secondary metabolites within a short period. Three culture methods， namely “solid-solid”， “solid-liquid” and “solid-liquid-solid”， were adopted to culture the embryogenic callus of F. cirrhosa. After 40 days of culture， structural differentiation， growth yield， in vitro antioxidant capacity， and bioactive metabolite accumulation were compared between the treatments. Significant differences were observed among the three culture methods for the embryogenic callus of F. cirrhosa， with the “solid-liquid-solid” mode showing the best overall performance. （1）Structural differentiation： The callus cultured by “solid-liquid-solid” mode showed uniform morphology， compact texture， and bright color. The cell nucleus was regular in shape and evenly distributed. The procambial bundle structure was well-defined， and the cells exhibited typical embryogenic characteristics. （2）Proliferation coefficient： The proliferation multiple of the callus under this system was 1.45， which was 2.84 and 1.93 times that of the “solid-solid” and “solid-liquid” cultures， respectively. （3）In vitro antioxidant capacity： The callus from the “solid-liquid-solid” culture exhibited strong DPPH radical scavenging activity（47.55%） and a high FRAP value（720.95 μmol⋅L^-1）， both of which were significantly superior to those of the “solid-solid” culture （40.41%， 482.86 μmol⋅L^-1） and the “solid-liquid” culture（44.15%， 694.68 μmol⋅L^-1）. （4）Bioactive metabolite accumulation： The embryogenic callus in this group exhibited the highest levels of soluble sugar（56.15 mg⋅g^-1）， total flavonoids（2.06 mg⋅g^-1）， total phenolic（1.44 mg⋅g^-1）， imperialine（0.25 mg⋅g^-1）. These values were 1.07， 1.15， 1.66， and 1.39 times as high as those of the “solid-solid” culture， and 1.23， 1.11， 1.13， and 1.19 times as high as those of the “solid-liquid” culture， respectively. This study provides a theoretical basis and technical support for the large-scale production of secondary metabolites and the establishment of a plant regeneration system for F. cirrhosa.



Expression Characteristics and Functional Analysis of the Jujube Gene ZjEB1c in Fruit Shape Regulation

Wenqing FU, Rongshen MA, Yue REN, Zimeng LI, Xu WANG, Wenhao BO

2026, 46(5):  899-912.  doi:10.7525/j.issn.1673-5102.2026.05.011

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To clarify the role of the jujube（Ziziphus jujuba） EB1 family member ZjEB1c in fruit shape formation， EB1 family members in the Jingzao 39 genome were identified， and their expression patterns were analyzed across seven developmental stages in seven jujube cultivars. Meanwhile， an overexpression vector of ZjEB1c was constructed and heterologously transformed into tomato（Solanum lycopersicum）， and its regulatory effects were investigated through fruit phenotypic observation and transcriptome sequencing. The results showed that two EB1 family members， ZjEB1a and ZjEB1c， were identified in jujube. ZjEB1c was upregulated during the middle stage of fruit development（50-78 days post-anthesis）， and its expression level was higher in long-fruit cultivars. Compared with the wild type， the average fruit shape index of three independent transgenic tomato lines increased from 0.928 to 1.117-1.168， and the number of mesocarp cells per unit area increased significantly at the longitudinal base. 744 commonly differentially expressed genes were identified through comparative transcriptome analysis， which were mainly enriched in hormone signal transduction， cell wall and pectin metabolism-related pathways. Comprehensive analysis indicated that ZjEB1c was closely associated with fruit longitudinal elongation and suggested that it participated in fruit shape formation through the processes related to hormone signal transduction and cell division.



Effects of Brassinolide and Gibberellic Acid on the Growth and Quality of Blue Honeysuckle

Jieru ZHAO, Yan LIU, Jing CHEN, Dong QIN, Junwei HUO

2026, 46(5):  913-926.  doi:10.7525/j.issn.1673-5102.2026.05.012

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Application of plant growth regulators is an effective strategy for regulating fruit tree growth and improving fruit quality. A field experiment was conducted to investigate the effects of brassinolide （EBR） and gibberellic acid（GA₃） by foliar spraying on blue honeysuckle （Lonicera caerulea ‘Lanjingling’） during the flowering stage. The results showed that both growth regulators effectively promoted plant growth and fruit quality. Specifically， 30 mg⋅L^-1 GA₃ significantly promoted new shoot growth， with a 72.95% increase compared with the control（CK）； 50 mg⋅L^-1 GA₃ was most effective for increasing fruit flavonoid content and hydroxyl radical scavenging rate， which rose by 63.06%and 14.67%， respectively， relative to CK. In contrast， 75 mg⋅L^-1 GA₃ maximized ascorbic acid content and DPPH radical scavenging rate， with increases of 27.10% and 11.52%， respectively， compared with CK. Application of 0.1 mg⋅L^-1 EBR significantly increased leaf chlorophyll content and fruit firmness by 30.55% and 27.02%， respectively， relative to CK. The combined treatment of 0.3 mg⋅L^-1 EBR and 75 mg⋅L^-1 GA₃ yielded the optimal results， increasing leaf area by 72.44%， total anthocyanin content by 78.54%， and total phenolic content by 45.40%， respectively， compared with CK； this combination also maximized the total antioxidant capacity of the fruit. Therefore， spraying appropriate concentrations of EBR and GA₃ during flowering is an effective cultivation practice for improving the growth and fruit quality of blue honeysuckle. This study provides a theoretical basis for the application of brassinolide and gibberellic acid in blue honeysuckle cultivation.



Structural Analysis of Nectar-Secreting Sites in the Bracts of Paeonia lactiflora

Weina GUO, Gang LI, Xiaoli XU, Li ZHAO, Lei CHENG

2026, 46(5):  927-937.  doi:10.7525/j.issn.1673-5102.2026.05.013

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Although the bracts of Paeonia lactiflora secrete nectar during the bud stage， their secretory sites lack the external morphological characteristics of typical nectaries. To clarify the structural basis of this phenomenon， this study employed semi-thin and ultra-thin sectioning techniques， combined with light microscopy， transmission electron microscopy， and scanning electron microscopy， to observe the microstructural and ultrastructural features of P. lactiflora bracts at different developmental stages. The results showed that： （1）The bracts comprise the epidermis， hypodermis， ground parenchyma， and vascular bundles. During the leaf-expansion stage， the epidermis， some hypodermal cells， and a few parenchyma cells stained deeply with toluidine blue； at the firm stage， staining in these cells became lighter as nectar was secreted. （2）At the leaf-expansion stage， the internal tissues of the bracts exhibited the ultrastructural features of secretory cells： abundant organelles， dense cytoplasm， and well-developed plasmodesmata， which constitute the structural basis for nectar secretion. （3）Nectar is primarily secreted through stomata， and the secretion process may involve symplastic and apoplastic pathways as well as exocytosis. In conclusion， P. lactiflora bracts do not differentiate into morphologically independent and functionally specialized nectary tissues during development， rather， they achieve nectar secretion through the functional specialization of existing tissues， thereby enriching the diversity of nectary structures in angiosperms.



Effects of Substrate Irrigation with Growth Retardants on Rooting of Cerasus humilis Softwood Cuttings

Xiaoyan WANG, Haonan WANG, Qianqian HE, Luting JIA, Shuai ZHANG, Pengfei WANG

2026, 46(5):  938-947.  doi:10.7525/j.issn.1673-5102.2026.05.014

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To investigate the mechanism of different growth retardants applied through substrate drenching in regulating the rooting of Cerasus humilis softwood cuttings， semi-lignified shoots of C. humilis ‘Nongda No.6’ were used as cuttings in this study. Suitable concentrations and application timings of plant growth regulators were screened， and the dynamic changes of nutrients and endogenous hormones during the rooting process were analyzed under an optimal treatment. The results demonstrated that substrate drenching with different concentrations of paclobutrazol（PP333） and uniconazole（S3307） both promoted rooting. Among them， the treatment with 100 mg⋅L^-1 PP333 yielded the highest rooting rate（60.83%）， which was significantly higher than that of the control. Moreover， the treatment with 50 mg⋅L^-1 S3307 exhibited the optimal comprehensive performance in terms of rooting rate， cutting survival， and root system development， indicating its suitability for the softwood cutting propagation of C. humilis. When 50 mg⋅L^-1 S3307 was applied at 7， 14， and 21 days after cutting， the highest rooting rate （57.22%） was observed in 14-d treatment （S-14 d）. During the rooting process， the S-14 d treatment significantly enhanced the accumulation of soluble sugars， starch， and soluble proteins in the stem segments of cuttings， resulting in a higher carbon-to-nitrogen ratio that accelerated rooting speed and enhanced root quality. Meanwhile， the S-14 d treatment increased the contents of indole-3-acetic acid（IAA） and zeatin （ZT）， inhibited the accumulation of gibberellic acid（GA₃） and abscisic acid（ABA）， which optimized the hormone balance to facilitate cutting rooting. In conclusion， substrate drenching with 50 mg⋅L^-1 S3307 at 14 days after cutting effectively promoted the rooting of C. humilis softwood cuttings by regulating nutrient metabolism and endogenous hormone levels．



Water-carbon Physiological Mechanism of Alternate Fruiting of Chinese Sea Buckthorn

Yongxin DAI, Chenkai NING, Peng TIAN, Zeqi LI, Xinli YANG, Lin WANG

2026, 46(5):  948-959.  doi:10.7525/j.issn.1673-5102.2026.05.015

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To explore the physiological mechanisms underlying alternate fruiting in Chinese sea buckthorn （Hippophae rhamnoides subsp. sinensis）， on-year female， off-year female， and male plants were selected as materials. Branch water potential， branch percentage loss of conductivity（PLC） and leaf net photosynthetic rate（P_n） were measured in the current year（2022）. Additionally， nonstructural carbohydrate （NSC） content in branches and leaves， branch growth， fruiting traits， and fruit nutrient contents were assessed in both the current year and the subsequent year（2023）. The results indicated that： （1）No significant differences were observed in branch predawn or midday water potential among the three plant types. Branch predawn PLC values remained below 20.0%， whereas midday PLC values ranged from 29.9% to 33.5%. （2） Significant differences in P_n were found among the groups. The P_n of on-year female plants was 55.7% and 20.0% higher than that of off-year female and male plants， respectively. In September of the current year， starch content in the branches and leaves of on-year female plants was 26.2%-42.2% lower than that of off-year female and male plants， with NSC content decreasing by 4.6%-29.7%. In April of the subsequent year， starch content in on-year female plants was 36.3%-60.5% lower， and NSC content decreased by 23.5%-42.1% compared to off-year female and male plants. （3）In the current year， growth indices of leaves， branches， and buds in off-year female plants were on average 19.1% higher than those in on-year female plants. In the subsequent year， growth indices in on-year female plants were on average 6.6% higher than those in off-year female plants. （4）In the current year， the fruit number per 10 cm of branch in on-year female plants was 2.1 times that of off-year female plants； however， longitudinal and transverse fruit diameters， mass per 100 fruits， and contents of soluble solids， vitamin C， and total flavonoids were significantly lower than those in off-year female plants. In the subsequent year， the fruit number per 10 cm of branch in off-year female plants was 1.3 times that of on-year female plants， with no significant differences in fruit nutrient contents. These findings suggest that branch， leaf， and bud growth in on-year female plants of sea buckthorn is significantly inhibited. The high carbohydrate demand for fruit development reduces carbon storage， leading to the decreased fruit and flower bud quality， which triggers alternate fruiting in the subsequent year. Water status is not the primary driver of alternate fruiting in Chinese sea buckthorn； rather， the storage and competition for photosynthetic products play a dominant role.



Effects of Successive Subculture on the Field Colonization Adaptability of Brachythecium plumosum in the Qinling Mountains

Yuhan YAN, Qiuxia MO, Chongfeng BU, Yakun TANG

2026, 46(5):  960-970.  doi:10.7525/j.issn.1673-5102.2026.05.016

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To investigate the effects of long-term artificial subculture on the field colonization adaptability of bryophyte crust seed sources， this study conducted indoor successive subculture， outdoor colonization， and artificial rainfall simulation scouring experiments on the saxicolous moss Brachythecium plumosum from the Qinling Mountains. The differences in field colonization adaptability among seed sources with different successive subculture generations were analyzed by measuring relevant growth and physiological indicators. The results showed that： （1）The initial generation（S0）， the first successive subculture generation（S1）， and the second successive subculture generation（S2） all colonized rapidly， with coverages exceeding 90% after 32 d， among which S0 exhibited the best overall growth performance. （2）After rainfall scouring， S0 showed the smallest reduction in coverage， and possessed the strongest crust stability and resistance to water erosion. （3） After 32 d of colonization， the photosynthetic pigment density of S0 remained stable， while the malondialdehyde（MDA） and soluble substance contents gradually decreased； S0 alleviated oxidative damage by increasing peroxidase（POD） activity and maintaining catalase（CAT） activity. In contrast， the photosynthetic pigment contents of S1 and S2 decreased significantly， while their MDA and soluble substance contents increased markedly. The antioxidant enzyme-responsive patterns of S1 were similar to that of S0， whereas S2 exhibited a distinct pattern， characterized primarily by a significant increase in CAT activity， with a limited increase in POD activity. （4）Redundancy analysis（RDA） showed that the physiological response patterns of S2 were significantly different from those of S0 and S1， with photosynthetic pigment content and antioxidant enzyme activities being the key indicators driving this differentiation. In conclusion， successive subculture weakens the adaptability of B. plumosum seed sources to natural environments. In engineering applications， the number of subculture generations should be reasonably controlled to ensure the ecological function and field colonization adaptability of bryophyte crust seed sources.