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Article(id=1198628667493548587, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1198628666650493481, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2022-1258, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1669132800000, receivedDateStr=2022-11-23, revisedDate=1674403200000, revisedDateStr=2023-01-23, acceptedDate=null, acceptedDateStr=null, onlineDate=1763704943774, onlineDateStr=2025-11-21, pubDate=1689091200000, pubDateStr=2023-07-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1763704943774, onlineIssueDateStr=2025-11-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1763704943774, creator=13701087609, updateTime=1763704943774, updator=13701087609, issue=Issue{id=1198628666650493481, tenantId=1146029695717560320, journalId=1189982191388893191, year='2023', volume='58', issue='7', pageStart='0', pageEnd='1980', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1763704943573, creator=13701087609, updateTime=1766137716668, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1208832456644490122, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1198628666650493481, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1208832456644490123, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1198628666650493481, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=1851, endPage=1858, ext={EN=ArticleExt(id=1198628668642787887, articleId=1198628667493548587, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=Exploring the protective mechanism of Tibetan medicine Potentilla anserine on cyclophosphamide-induced myelosuppression based on metabonomics technology, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

The study aims to explore the effects and mechanisms of water extract of Potentilla anserina (PA) on myelosuppression mice induced by cyclophosphamide based on metabonomics. The myelosuppressive mouse model was established by injected with cyclophosphamide and treated with water extract of PA. Thymus and spleen indexes, peripheral hemogram and bone marrow nucleated cells of each group was detected. Bone marrow pathology analysis was performed by hematoxylin-eosin staining. The levels of interleukin 3 (IL-3), interleukin 6 (IL-6), erythropoietin (EPO), granulocyte colony stimulating factor (GM-CSF), malondialdehyde (MDA), superoxide dismutase (SOD) and catalase (CAT) in serum were measured. The changes of biomarkers and related metabolic pathways were analyzed by UPLC-Q-TOF/MS-based metabonomics. Animal experiments were approved by the Animal Ethics Committee of Southwest Minzu University. The high doses of PA could significantly improve the decrease of white blood cell (WBC), red blood cell (RBC) counts and hemoglobin (HGB) levels of mice induced by cyclophosphamide (P < 0.05), and significantly increase the number of nucleated cells and the area of hematopoietic tissue in femoral bone marrow. The medium and high doses of PA could significantly improve the serum levels of SOD, CAT, MDA, IL-6 and GM-CSF (P < 0.05), and have no significant effect on the expression of IL-3 and EPO (P > 0.05). Serum metabolomics analysis showed that the aqueous extracts of PA could alleviate myrosuppression by regulating the aminoacyl-tRNA, valine, leucine and isoleucine biosynthesis mediated by 13 different metabolites such as valine, leucine, asparagine and hydroxyisohexic acid. PA improve the inhibition of hematopoietic function in myelosuppression mouse, and its mechanisms may be related to anti-oxidation and promoting the expression of hematopoietic-related cytokines and regulating the related metabolic pathways.

, correspAuthors=Wei LIU, Pu-yang GONG, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2023 Acta Pharmaceutica Sinica. All rights reserved., copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Jing-xian LIU, Xiao-min LUO, Jian GU, Shi-guang HUANG, Qin WANG, Wei LIU, Pu-yang GONG), CN=ArticleExt(id=1198628669884301919, articleId=1198628667493548587, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=基于代谢组学技术探究藏药蕨麻对环磷酰胺致骨髓抑制的改善作用机制, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

探究蕨麻水提物对环磷酰胺诱导骨髓抑制小鼠的影响及其代谢组学机制。采用环磷酰胺建立骨髓抑制小鼠模型并给予蕨麻水提物治疗, 计算各组小鼠胸腺与脾指数, 检测血常规变化, 测定骨髓有核细胞数, 并采用苏木精-伊红(hematoxylin-eosin, HE) 染色进行骨髓病理学分析。测定小鼠血清中白细胞介素3 (interleukin 3, IL-3)、白细胞介素6 (interleukin 6, IL-6)、促红细胞生成素(erythropoietin, EPO)、粒细胞巨噬细胞集落刺激因子(granulocyte macrophage colony stimulating factor, GM-CSF)、丙二醛(malondialdehyde, MDA)、超氧化物歧化酶(superoxide dismutase, SOD) 及过氧化氢酶(catalase, CAT)水平。借助UPLC-Q-TOF/MS进行血清非靶向代谢组学分析, 探析生物标志物变化及所涉及的代谢通路。动物实验操作遵循西南民族大学实验动物伦理委员会规定。蕨麻高剂量组可显著改善环磷酰胺诱导的小鼠外周血中白细胞(white blood cell, WBC)、红细胞(red blood cell, RBC) 计数及血红蛋白(hemoglobin, HGB) 水平下降(P < 0.05), 显著增加股骨骨髓中有核细胞数目及造血组织面积。蕨麻中、高剂量可显著改善小鼠血清中SOD、CAT、MDA、IL-6及GM-CSF水平(P < 0.05), 对IL-3及EPO表达无显著影响(P > 0.05)。血清代谢组学分析表明蕨麻水提物可通过调控缬氨酸、亮氨酸、L-天冬酰胺及羟基异己酸等13个差异代谢物介导的氨酰基-tRNA生物合成、缬氨酸、亮氨酸和异亮氨酸的生物合成等通路改善骨髓抑制。蕨麻可改善环磷酰胺诱导的小鼠骨髓造血功能抑制, 其作用机制可能与抗氧化、促进造血相关细胞因子的表达及调节相关代谢通路有关。

, correspAuthors=刘伟, 龚普阳, authorNote=null, correspAuthorsNote=
*刘伟, E-mail: ;
龚普阳, E-mail:
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A: PCA score plots in positive ion mode; B: PLS-DA score plots in positive ion mode; C: PLS-DA validation plot in positive ion mode; D: PCA score plots in negative ion mode; E: PLS-DA score plots in negative ion mode; F: PLS-DA validation plot in negative ion mode. PCA: Principal component analysis; PLS-DA: Partial least squares discriminant analysis , figureFileSmall=5FnWEZDLjdRSXk2tKKPmrA==, figureFileBig=Q6I0sxoKTojw+YK2LzfSyg==, tableContent=null), ArticleFig(id=1198960097079689628, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=EN, label=null, caption=null, figureFileSmall=1hmGDOq5hjMpsnc0nWiJ5w==, figureFileBig=34VqOYjYlOQ8bqCSgnxKtQ==, tableContent=null), ArticleFig(id=1198960097293599147, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Figure 3, caption= Metabolic pathway enrichment analysis of the differential metabolites , figureFileSmall=1hmGDOq5hjMpsnc0nWiJ5w==, figureFileBig=34VqOYjYlOQ8bqCSgnxKtQ==, tableContent=null), ArticleFig(id=1198960097432011189, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Group Thymus index/mg·g-1 Spleen index/mg·g-1
CON 3.14 ± 0.80 4.32 ± 0.56
MOD 1.55 ± 0.39# 3.23 ± 0.60#
PA-L 2.06 ± 0.56 4.13 ± 1.14
PA-M 1.99 ± 0.70 3.82 ± 0.72
PA-H 1.69 ± 0.56 3.77 ± 0.83
rhG-CSF 2.06 ± 0.39 5.36 ± 0.74**
), ArticleFig(id=1198960097599783366, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 1, caption=

Effects of Potentilla anserine (PA) on the thymus and spleen indexes of myelosuppression mice induced by cyclophosphamide. n = 10, $\bar{x}$ ± s. #P < 0.05 vs CON; **P < 0.01 vs MOD. CON: Control; MOD: Model; PA-L: Potentilla anserine extract in low dose, 1 g·kg-1; PA-M: Potentilla anserine extract in middle dose, 2 g·kg-1; PA-H: Potentilla anserine extract in high dose, 4 g·kg-1; rhG-CSF: Recombinant human granulocytes colony stimulating factor, 10 μg·kg-1

, figureFileSmall=null, figureFileBig=null, tableContent=
Group Thymus index/mg·g-1 Spleen index/mg·g-1
CON 3.14 ± 0.80 4.32 ± 0.56
MOD 1.55 ± 0.39# 3.23 ± 0.60#
PA-L 2.06 ± 0.56 4.13 ± 1.14
PA-M 1.99 ± 0.70 3.82 ± 0.72
PA-H 1.69 ± 0.56 3.77 ± 0.83
rhG-CSF 2.06 ± 0.39 5.36 ± 0.74**
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Group WBC/×109·L-1 RBC/×109·L-1 HGB/g·L-1 PLT/×109·L-1
CON 5.53 ± 1.33 7.78 ± 0.37 131.2 ± 8.00 1200.44 ± 138.66
MOD 1.97 ± 0.85## 6.37 ± 0.53## 107.3 ± 10.60## 976.88 ± 122.03#
PA-L 2.86 ± 0.58 6.78 ± 0.54 111.4 ± 7.62 894.9 ± 154.59
PA-M 3.43 ± 0.70 7.11 ± 0.59* 120.1 ± 9.15* 927.8 ± 202.59
PA-H 4.05 ± 1.13* 7.01 ± 0.68* 122.7 ± 12.53** 963.22 ± 154.53
rhG-CSF 11.66 ± 2.35** 6.79 ± 0.42 119.7 ± 8.38* 959.3 ± 163.20
), ArticleFig(id=1198960097985659365, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 2, caption=

Effects of PA on the peripheral hemogram of myelosuppression mice. n = 10, $\bar{x}$ ± s. #P < 0.05, ##P < 0.01 vs CON; *P < 0.05, **P < 0.01 vs MOD. WBC: White blood cell; RBC: Red blood cell; HGB: Hemoglobin; PLT: Platelet

, figureFileSmall=null, figureFileBig=null, tableContent=
Group WBC/×109·L-1 RBC/×109·L-1 HGB/g·L-1 PLT/×109·L-1
CON 5.53 ± 1.33 7.78 ± 0.37 131.2 ± 8.00 1200.44 ± 138.66
MOD 1.97 ± 0.85## 6.37 ± 0.53## 107.3 ± 10.60## 976.88 ± 122.03#
PA-L 2.86 ± 0.58 6.78 ± 0.54 111.4 ± 7.62 894.9 ± 154.59
PA-M 3.43 ± 0.70 7.11 ± 0.59* 120.1 ± 9.15* 927.8 ± 202.59
PA-H 4.05 ± 1.13* 7.01 ± 0.68* 122.7 ± 12.53** 963.22 ± 154.53
rhG-CSF 11.66 ± 2.35** 6.79 ± 0.42 119.7 ± 8.38* 959.3 ± 163.20
), ArticleFig(id=1198960098124071413, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Group Bone marrow nucleated cells
CON 8.62 ± 1.41
MOD 4.5 ± 1.31##
PA-L 4.38 ± 1.06
PA-M 6.25 ± 0.89*
PA-H 6.50 ± 1.77*
rhG-CSF 10.1 ± 1.36**
), ArticleFig(id=1198960098258289159, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 3, caption=

Effects of PA on the bone marrow nucleated cells of myelosuppression mice. n = 10, $\bar{x}$ ± s. ##P < 0.01 vs CON; *P < 0.05, **P < 0.01 vs MOD

, figureFileSmall=null, figureFileBig=null, tableContent=
Group Bone marrow nucleated cells
CON 8.62 ± 1.41
MOD 4.5 ± 1.31##
PA-L 4.38 ± 1.06
PA-M 6.25 ± 0.89*
PA-H 6.50 ± 1.77*
rhG-CSF 10.1 ± 1.36**
), ArticleFig(id=1198960098505753119, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Group SOD/U·mL-1 MDA/nmol·mL-1 CAT/U·mL-1
CON 152.52 ± 7.07 6.35 ± 1.73 6.02 ± 0.73
MOD 117.01 ± 18.84## 17.55 ± 3.58## 3.29 ± 0.59##
PA-L 135.18 ± 11.46* 12.45 ± 2.94** 4.35 ± 0.84
PA-M 141.18 ± 15.31** 12.28 ± 3.28** 5.31 ± 1.08**
PA-H 142.52 ± 11.75** 10.46 ± 1.31** 5.51 ± 0.64**
rhG-CSF 138.83 ± 6.20* 10.97 ± 2.37** 4.99 ± 0.48**
), ArticleFig(id=1198960098698691119, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 4, caption=

Effects of PA on the SOD, MDA and CAT levels of myelosuppression mice. n = 10, $\bar{x}$ ± s. ##P < 0.01 vs CON; *P < 0.05, **P < 0.01 vs MOD. SOD: Superoxide dismutase; MDA: Malondialdehyde; CAT: Catalase

, figureFileSmall=null, figureFileBig=null, tableContent=
Group SOD/U·mL-1 MDA/nmol·mL-1 CAT/U·mL-1
CON 152.52 ± 7.07 6.35 ± 1.73 6.02 ± 0.73
MOD 117.01 ± 18.84## 17.55 ± 3.58## 3.29 ± 0.59##
PA-L 135.18 ± 11.46* 12.45 ± 2.94** 4.35 ± 0.84
PA-M 141.18 ± 15.31** 12.28 ± 3.28** 5.31 ± 1.08**
PA-H 142.52 ± 11.75** 10.46 ± 1.31** 5.51 ± 0.64**
rhG-CSF 138.83 ± 6.20* 10.97 ± 2.37** 4.99 ± 0.48**
), ArticleFig(id=1198960098920989243, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
Group GM-CSF/ng·L-1 IL-6/ng·L-1 EPO/ng·L-1 IL-3/ng·L-1
CON 284.28 ± 29.26 17.04 ± 2.68 26.14 ± 2.23 116.41 ± 7.31
MOD 175.36 ± 37.20## 9.91 ± 1.90## 29.63 ± 4.74 101.09 ± 11.17
PA-L 203.04 ± 40.65 10.92 ± 2.08 27.12 ± 3.72 102.87 ± 20.38
PA-M 232.74 ± 19.80* 14.07 ± 2.82* 30.21 ± 4.47 117.46 ± 12.84
PA-H 225.11 ± 33.00* 13.52 ± 1.27* 29.69 ± 5.30 107.49 ± 13.60
rhG-CSF 229.36 ± 23.56* 15.47 ± 2.75** 25.76 ± 4.81 114.69 ± 22.31
), ArticleFig(id=1198960099118121549, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 5, caption=

Effects of PA on the hematopoiesis-related cytokines in the serum of myelosuppression mice. n = 10, $\bar{x}$ ± s. ##P < 0.01 vs CON; *P < 0.05, **P < 0.01 vs MOD. GM-CSF: Granulocyte macrophage colony stimulating factor; IL-6: Interleukin 6; EPO: Erythropoietin; IL-3: Interleukin 3

, figureFileSmall=null, figureFileBig=null, tableContent=
Group GM-CSF/ng·L-1 IL-6/ng·L-1 EPO/ng·L-1 IL-3/ng·L-1
CON 284.28 ± 29.26 17.04 ± 2.68 26.14 ± 2.23 116.41 ± 7.31
MOD 175.36 ± 37.20## 9.91 ± 1.90## 29.63 ± 4.74 101.09 ± 11.17
PA-L 203.04 ± 40.65 10.92 ± 2.08 27.12 ± 3.72 102.87 ± 20.38
PA-M 232.74 ± 19.80* 14.07 ± 2.82* 30.21 ± 4.47 117.46 ± 12.84
PA-H 225.11 ± 33.00* 13.52 ± 1.27* 29.69 ± 5.30 107.49 ± 13.60
rhG-CSF 229.36 ± 23.56* 15.47 ± 2.75** 25.76 ± 4.81 114.69 ± 22.31
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No. Metabolite Formula RT/s VIP Fold change Experimental m/z ESI mode Trend
MOD/CON PA-H/MOD
1 Hydroxyisocaproic acid C6H12O3 148.36 2.90 0.44 131.07 - ↑## ↓**
2 N-Acetylcytidine C11H15N3O6 164.28 3.19 0.40 284.09 - ↑## ↓**
3 Pseudouridine C9H12N2O6 237.94 2.57 0.70 243.06 - ↑# ↓**
4 Asparagine C4H8N2O3 237.73 1.05 0.72 153.03 - ↑# ↓**
5 N-Acetyltryptophan C13H14N2O3 189.52 2.84 0.63 245.09 - ↑# ↓**
6 Valine C5H11NO2 293.15 1.88 1.32 116.07 - ↓## ↑*
7 PE(18:0/18:2(9Z, 12Z)) C41H78NO8P 146.28 4.64 1.29 742.54 - ↓## ↑**
8 Phenylalanine C9H11NO2 250.52 2.74 1.26 164.07 - ↓# ↑*
9 PE(P-18:0/22:6(4Z, 7Z, 10Z, 13Z, 16Z, 19Z)) C45H78NO7P 141.93 4.26 1.47 774.55 ↓# ↑*
10 LysoPE(18:2(9Z, 12Z)/0:0) C23H44NO7P 195.15 5.74 1.51 476.28 - ↓# ↑**
11 Leucine C6H13NO2 256.91 2.12 1.42 132.10 + ↓## ↑*
12 2-Amino-1-phenylethanol C8H11NO 250.37 3.63 1.29 120.08 + ↓## ↑*
13 Ectoine C6H10N2O2 328.23 2.13 2.02 143.08 + ↓# ↑*
), ArticleFig(id=1198960099503997547, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1198628667493548587, language=CN, label=Table 6, caption=

Differential metabolites in plasma after treated with PA-H. RT: Retention time; VIP: Variable importance in projection; ESI: Electron spray ionization. #P < 0.05, ##P < 0.01; *P < 0.05, **P < 0.01

, figureFileSmall=null, figureFileBig=null, tableContent=
No. Metabolite Formula RT/s VIP Fold change Experimental m/z ESI mode Trend
MOD/CON PA-H/MOD
1 Hydroxyisocaproic acid C6H12O3 148.36 2.90 0.44 131.07 - ↑## ↓**
2 N-Acetylcytidine C11H15N3O6 164.28 3.19 0.40 284.09 - ↑## ↓**
3 Pseudouridine C9H12N2O6 237.94 2.57 0.70 243.06 - ↑# ↓**
4 Asparagine C4H8N2O3 237.73 1.05 0.72 153.03 - ↑# ↓**
5 N-Acetyltryptophan C13H14N2O3 189.52 2.84 0.63 245.09 - ↑# ↓**
6 Valine C5H11NO2 293.15 1.88 1.32 116.07 - ↓## ↑*
7 PE(18:0/18:2(9Z, 12Z)) C41H78NO8P 146.28 4.64 1.29 742.54 - ↓## ↑**
8 Phenylalanine C9H11NO2 250.52 2.74 1.26 164.07 - ↓# ↑*
9 PE(P-18:0/22:6(4Z, 7Z, 10Z, 13Z, 16Z, 19Z)) C45H78NO7P 141.93 4.26 1.47 774.55 ↓# ↑*
10 LysoPE(18:2(9Z, 12Z)/0:0) C23H44NO7P 195.15 5.74 1.51 476.28 - ↓# ↑**
11 Leucine C6H13NO2 256.91 2.12 1.42 132.10 + ↓## ↑*
12 2-Amino-1-phenylethanol C8H11NO 250.37 3.63 1.29 120.08 + ↓## ↑*
13 Ectoine C6H10N2O2 328.23 2.13 2.02 143.08 + ↓# ↑*
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基于代谢组学技术探究藏药蕨麻对环磷酰胺致骨髓抑制的改善作用机制
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刘静贤 , 罗晓敏 , 顾健 , 黄时光 , 王琴 , 刘伟 * , 龚普阳 *
药学学报 | 研究论文 2023,58(7): 1851-1858
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药学学报 | 研究论文 2023, 58(7): 1851-1858
基于代谢组学技术探究藏药蕨麻对环磷酰胺致骨髓抑制的改善作用机制
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刘静贤, 罗晓敏, 顾健, 黄时光, 王琴, 刘伟* , 龚普阳*
作者信息
  • 西南民族大学药学院, 四川 成都 610041

通讯作者:

*刘伟, E-mail: ;
龚普阳, E-mail:
Exploring the protective mechanism of Tibetan medicine Potentilla anserine on cyclophosphamide-induced myelosuppression based on metabonomics technology
Jing-xian LIU, Xiao-min LUO, Jian GU, Shi-guang HUANG, Qin WANG, Wei LIU* , Pu-yang GONG*
Affiliations
  • College of Pharmacy, Southwest Minzu University, Chengdu 610041, China
出版时间: 2023-07-12 doi: 10.16438/j.0513-4870.2022-1258
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摘要
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探究蕨麻水提物对环磷酰胺诱导骨髓抑制小鼠的影响及其代谢组学机制。采用环磷酰胺建立骨髓抑制小鼠模型并给予蕨麻水提物治疗, 计算各组小鼠胸腺与脾指数, 检测血常规变化, 测定骨髓有核细胞数, 并采用苏木精-伊红(hematoxylin-eosin, HE) 染色进行骨髓病理学分析。测定小鼠血清中白细胞介素3 (interleukin 3, IL-3)、白细胞介素6 (interleukin 6, IL-6)、促红细胞生成素(erythropoietin, EPO)、粒细胞巨噬细胞集落刺激因子(granulocyte macrophage colony stimulating factor, GM-CSF)、丙二醛(malondialdehyde, MDA)、超氧化物歧化酶(superoxide dismutase, SOD) 及过氧化氢酶(catalase, CAT)水平。借助UPLC-Q-TOF/MS进行血清非靶向代谢组学分析, 探析生物标志物变化及所涉及的代谢通路。动物实验操作遵循西南民族大学实验动物伦理委员会规定。蕨麻高剂量组可显著改善环磷酰胺诱导的小鼠外周血中白细胞(white blood cell, WBC)、红细胞(red blood cell, RBC) 计数及血红蛋白(hemoglobin, HGB) 水平下降(P < 0.05), 显著增加股骨骨髓中有核细胞数目及造血组织面积。蕨麻中、高剂量可显著改善小鼠血清中SOD、CAT、MDA、IL-6及GM-CSF水平(P < 0.05), 对IL-3及EPO表达无显著影响(P > 0.05)。血清代谢组学分析表明蕨麻水提物可通过调控缬氨酸、亮氨酸、L-天冬酰胺及羟基异己酸等13个差异代谢物介导的氨酰基-tRNA生物合成、缬氨酸、亮氨酸和异亮氨酸的生物合成等通路改善骨髓抑制。蕨麻可改善环磷酰胺诱导的小鼠骨髓造血功能抑制, 其作用机制可能与抗氧化、促进造血相关细胞因子的表达及调节相关代谢通路有关。

蕨麻  /  环磷酰胺  /  骨髓抑制  /  代谢组学

The study aims to explore the effects and mechanisms of water extract of Potentilla anserina (PA) on myelosuppression mice induced by cyclophosphamide based on metabonomics. The myelosuppressive mouse model was established by injected with cyclophosphamide and treated with water extract of PA. Thymus and spleen indexes, peripheral hemogram and bone marrow nucleated cells of each group was detected. Bone marrow pathology analysis was performed by hematoxylin-eosin staining. The levels of interleukin 3 (IL-3), interleukin 6 (IL-6), erythropoietin (EPO), granulocyte colony stimulating factor (GM-CSF), malondialdehyde (MDA), superoxide dismutase (SOD) and catalase (CAT) in serum were measured. The changes of biomarkers and related metabolic pathways were analyzed by UPLC-Q-TOF/MS-based metabonomics. Animal experiments were approved by the Animal Ethics Committee of Southwest Minzu University. The high doses of PA could significantly improve the decrease of white blood cell (WBC), red blood cell (RBC) counts and hemoglobin (HGB) levels of mice induced by cyclophosphamide (P < 0.05), and significantly increase the number of nucleated cells and the area of hematopoietic tissue in femoral bone marrow. The medium and high doses of PA could significantly improve the serum levels of SOD, CAT, MDA, IL-6 and GM-CSF (P < 0.05), and have no significant effect on the expression of IL-3 and EPO (P > 0.05). Serum metabolomics analysis showed that the aqueous extracts of PA could alleviate myrosuppression by regulating the aminoacyl-tRNA, valine, leucine and isoleucine biosynthesis mediated by 13 different metabolites such as valine, leucine, asparagine and hydroxyisohexic acid. PA improve the inhibition of hematopoietic function in myelosuppression mouse, and its mechanisms may be related to anti-oxidation and promoting the expression of hematopoietic-related cytokines and regulating the related metabolic pathways.

Potentilla anserina  /  cyclophosphamide  /  bone marrow suppression  /  metabonomics
刘静贤, 罗晓敏, 顾健, 黄时光, 王琴, 刘伟, 龚普阳. 基于代谢组学技术探究藏药蕨麻对环磷酰胺致骨髓抑制的改善作用机制. 药学学报, 2023 , 58 (7) : 1851 -1858 . DOI: 10.16438/j.0513-4870.2022-1258
Jing-xian LIU, Xiao-min LUO, Jian GU, Shi-guang HUANG, Qin WANG, Wei LIU, Pu-yang GONG. Exploring the protective mechanism of Tibetan medicine Potentilla anserine on cyclophosphamide-induced myelosuppression based on metabonomics technology[J]. Acta Pharmaceutica Sinica, 2023 , 58 (7) : 1851 -1858 . DOI: 10.16438/j.0513-4870.2022-1258
正文
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蕨麻为蔷薇科植物鹅绒委陵菜Potentilla anserina L.的干燥块根, 又名戳玛、人参果或长寿果等, 在我国主产于西藏、青海、甘肃和宁夏等地[1]。蕨麻使用历史悠久且为兼具食用和药用价值的传统藏药材之一, 藏语称作“卓老洒曾”。藏医典籍《四部医典》、《月王药诊》及现代著作《中国藏药》等均记载蕨麻具有益气补血、生津利痰及收敛止血等传统功效[2]。蕨麻的现代植物化学与药效学研究已有诸多报道, 其具有显著的抗乙肝病毒、神经保护、抗氧化及抗缺氧等功能特征, 且主要活性物质包括多糖、黄酮及皂苷等[3, 4]。益气补血是蕨麻的主要传统功效之一, 已有研究显示其提取物可增加正常小鼠外周血中红细胞数目及血红蛋白含量[5]。然而, 蕨麻对造血功能障碍相关病理状态的改善作用及机制如何仍鲜见报道。
来源于天然产物的传统草药因其多成分及整体调节的作用特点, 使得阐释其复杂的生物学作用机制成为研究难点。代谢组学技术是系统生物学的重要组成部分, 其可在定量及定性水平上表征体液、组织与器官等生物体样本中代谢物的变化及特征[6]。内源性小分子代谢物作为机体各种病理生理过程的终产物, 可为病理机制和药物作用机制研究提供线索[7]。因此, 代谢组学技术可为蕨麻的改善造血功能障碍的作用机制研究提供有效工具。
为进一步明确蕨麻益气补血的功效内涵, 本研究采用环磷酰胺致骨髓抑制小鼠模型, 评价蕨麻水提取对模型小鼠造血功能的改善作用, 开展非靶向血清代谢组学分析探讨其可能的作用机制, 以期为蕨麻这一传统藏药材的药用价值挖掘提供一定的科学依据。
材料与方法
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实验材料与试剂   蕨麻药材(批号: 20200608) 采自青海省玉树藏族自治州, 经西南民族大学龚普阳博士鉴定为蔷薇科委陵菜属植物鹅绒委陵菜Potentilla anserina L. 的干燥膨大块根。注射用环磷酰胺(CTX, 德国Baxter Oncology GmbH公司, 批号: 9J338A); 重组人粒细胞刺激因子注射液(recombinant human granulocytes colony stimulating factor, rhG-CSF, 齐鲁制药有限公司, 批号: 202005011KBB); 丙二醛(MDA, 批号: 20210722)、超氧化物歧化酶(SOD, 批号: 20210223) 及过氧化氢酶(CAT, 批号: 20210520) 购自南京建成生物工程研究所; 白细胞介素6 (IL-6, 批号: 20210344)、白细胞介素3 (IL-3, 批号: 20210514)、粒细胞巨噬细胞集落刺激因子(GM-CSF, 批号: 20210508)及促红细胞生成素(EPO, 批号: 20210505) 酶联免疫吸附实验(enzyme-linked immuno sorbent assay, ELISA) 试剂盒购自上海科鉴生物科技有限责任公司。
仪器   AB Triple TOF6600质谱仪(美国AB SCIEX公司); Agilent 1290 Infinity LC超高效液相色谱(美国Agilent公司); HH-2数显恒温水浴锅(常州智博瑞仪器制造有限公司); CYT5M酶标仪(美国BioTek公司); BX53显微镜(日本奥林巴斯公司); BC-2800Vet兽用全自动血液细胞分析仪(深圳迈瑞生物医疗电子股份有限公司); 2000A旋转蒸发仪(上海贤德实验仪器有限公司); 5430R低温高速离心机(德国Eppendorf公司)。
药物配置   称取100 g蕨麻药材, 加入1 000 mL蒸馏水浸泡1 h, 然后加热回流提取2次, 每次1 h, 过滤并合并提取液, 减压浓缩, 得到1 g (生药量)·mL-1浓缩液, 而后分别配制为含生药量0.1、0.2、0.4 g·mL-1三种浓度的溶液。
实验动物   SPF级雄性ICR小鼠60只, 体质量20 ± 2 g, 购于成都达硕实验动物有限公司, 生产许可证号: SCXK (川) 2020-030。实验动物置于室温25 ± 1 ℃, 65%湿度环境中饲养, 光周期为12 h光照与黑暗交替, 适应性饲养1周, 对实验动物所有操作及处理均获得西南民族大学实验动物伦理委员会批准。
动物的分组、模型建立及给药   小鼠根据体质量随机分为6组, 设置为空白组(control, CON)、模型组(model, MOD)、蕨麻提取物低(Potentilla anserine extract in low dose, PA-L)、中(Potentilla anserine extract in middle dose, PA-M)、高剂量(Potentilla anserine extract in high dose, PA-H) 组及阳性对照组(rhG-CSF), 每组10只。除空白组外, 各组小鼠连续5天腹腔注射环磷酰胺40 mg·kg-1[8, 9]。期间按各给药组剂量进行给药, 模型组给予等量的生理盐水。依照蕨麻临床推荐用药剂量, 采用体表面积法换算得到小鼠蕨麻低、中、高剂量组剂量分别为1、2、4 g·kg-1。阳性对照组皮下注射rhG-CSF, 参考相关文献及根据前期研究结果设定为10 μg·kg-1 [9, 10], 各组给药共9天。
样本采集   各组小鼠于末次给药后禁食不禁水12 h, 采用1.5%异氟烷麻醉, 称重, 眼底静脉丛取血, 将200 μL全血收集至预先加入EDTA-2Na的EP管中, 采用全自动血细胞分析仪进行血常规分析。收集小鼠剩余血液, 室温下静置1 h, 3 500 r·min-1离心10 min, 收集血清置于-80 ℃储存, 用于血清生化及代谢组学分析。收集小鼠胸腺和脾脏并称重, 计算胸腺指数和脾指数: 胸腺/脾指数＝胸腺/脾质量(mg)/小鼠体重(g)。另取小鼠双侧股骨用于HE染色及骨髓有核细胞数分析。
血清生化指标检测   测定各组小鼠血清中造血功能相关细胞因子(IL-3、IL-6、EPO及GM-CSF) 及氧化应激相关指标(SOD、MDA及CAT) 水平, 具体实验操作步骤遵照试剂盒说明书进行。
骨髓组织病理学分析   分离小鼠左侧股骨, 剔除附着的肌肉和结缔组织, 置于4%的多聚甲醛中固定3天, 采用梯度酒精脱水, 石蜡包埋, 切片机切片(厚度5 μm)。切片采用苏木精染色3 min, 使用盐酸水溶液分化, 氨水返蓝, 水洗, 切片使用85%及95%梯度酒精脱水, 采用伊红染色液进行染色5 min, 脱水, 使用中性树胶进行封片, 最后利用显微镜采集组织病理图像进行分析。
骨髓有核细胞数检测   小鼠取血后, 脱颈椎处死小鼠, 于75%酒精中浸泡10 min后置于超净台中, 取右侧股骨并剥离附着肌肉组织, 剪开股骨两端, 暴露骨髓腔。使用2 mL PBS溶液将骨髓腔中细胞冲出, 用细胞筛过滤多余杂质, 1 500 r·min-1离心5 min, 过4号针头, 制备骨髓单细胞悬液。将制备好的骨髓单细胞悬液转移至无菌EP管中, 1 500 r·min-1离心5 min, 弃上清, 加入红细胞裂解液, 重复离心, 弃上清液, 加入PBS稀释100倍后, 吸取10 μL注入细胞计数板, 计数骨髓有核细胞数。
血清代谢组学样本的制备与分析   取空白组、模型组及蕨麻提取物高剂量组各6个小鼠血清样本, 分别加入预冷的甲醇/乙腈/水(2∶2∶1) 溶液, 涡旋混匀后低温超声30 min, 于-4 ℃下14 000 ×g离心20 min, 真空干燥, 加入50%乙腈水溶液复溶, -4 ℃下14 000 ×g离心20 min, 取上清液进行检测分析。取各组待测样品等量混合, 制得质量控制(quality control, QC) 样本。在Agilent 1290 Infinity LC超高效液相色谱系统HILIC色谱柱进行分离, 流动相A为含25 mmol·L-1乙酸铵及25 mmol·L-1氨水的水溶液, 流动相B为乙腈, 梯度洗脱程序为: 0~0.5 min, 95% B; 0.5~7 min, 95%~65% B; 7~8 min, 65%~40% B; 8~9 min, 40% B; 9~9.1 min, 40%~95% B; 9.1~12 min, 95% B。进样量2 μL, 柱温25 ℃, 流速0.5 mL·min-1。采用电喷雾电离(ESI) 正离子和负离子模式检测, 参数如下: 辅助气1 (Gas1) 和辅助气2 (Gas2) 均为60; 气帘气(CUR), 30 psi (1 psi ≈ 6.9 kPa); 离子源温度, 600 ℃; 喷雾电压(ISVF), 5 500 V; 一级质荷比检测范围为m/z 60~1 000; 二级质荷比检测范围为m/z 25~1 000。
统计学方法   所有药效评价数据采用Graphpad Prism 8.0软件进行分析, 以mean ± SD表示。采用单因素方差分析, 其中三组以及三组以上的数据比较分析采用Dunnett's检验。P < 0.05表明两组间数据具统计学差异。数据通过ProteoWizard将UPLC-QTOF/MS获得原始转化为.mzXML格式, 借助XCMS进行峰对齐、保留时间校正和峰面积提取, 从而进行代谢物鉴定。将所得数据导入SIMCA 14.0软件中进行主成分分析(principal component analysis, PCA) 及正交偏最小二乘判别分析(orthogonal partial least squares discriminant analysis, OPLS-DA)。以OPLS-DA中变量投影重要性值(variable importance in projection, VIP) 大于1及P < 0.05为标准筛选差异代谢标志物。借助MetaboAnalyst 5.0平台(https://www.metaboanalyst.ca/) 进行代谢物通路富集分析。
结果
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1 蕨麻提取物对模型小鼠表观情况及脏器指数的影响
空白组小鼠行为活动正常, 毛发浓密且有光泽, 无明显脱毛现象。观察发现模型组小鼠活动比空白组有所减少, 精神萎靡, 且有普遍地脱毛现象; 各给药组小鼠精神状态及脱毛情况具有一定改善。解剖取各组小鼠胸腺及脾并测定脏器指数结果如表 1所示。与空白组比较, 模型组小鼠胸腺和脾指数显著降低(P < 0.05)。与模型组比较, 蕨麻各剂量组小鼠胸腺指数、脾指数无显著性差异(P > 0.05), 阳性对照组脾指数较模型组明显增高(P < 0.05)。
2 蕨麻水提物对骨髓抑制模型小鼠血常规参数的影响
如表 2所示, 与空白组比较, 模型组小鼠外周血中WBC、RBC、HGB、及血小板(platelet, PLT)水平显著降低(P < 0.05), 表明腹腔注射环磷酰胺导致了外周血细胞的减少。与模型组比较, 蕨麻水提物高剂量及阳性药物可显著升高小鼠外周血中WBC、RBC和HGB水平(P < 0.05), 而对PLT水平无显著影响, 表明蕨麻对骨髓抑制小鼠外周血中红细胞、白细胞及血红蛋白的形成具有一定的促进作用。
3 蕨麻提取物对模型小鼠骨髓有核细胞数的影响
如表 3所示, 与空白组比较, 模型组小鼠股骨骨髓的有核细胞数显著减少(P < 0.05), 表明环磷酰胺可显著抑制骨髓有核细胞产生。与模型组比较, 蕨麻中、高剂量组及阳性对照组小鼠骨髓真核细胞数明显增多(P < 0.05), 表明蕨麻可促进化疗药物导致骨髓造血功能损伤的修复。
4 蕨麻提取物对骨髓抑制小鼠骨髓病理变化的影响
如图 1所示, 空白组小鼠股骨骨髓腔中可见处于不同分化期的血细胞、间充质细胞和丰富的血窦, 未见显著的造血组织减少或增加。连续腹腔注射给予环磷酰胺后, 小鼠骨髓腔血细胞减少, 表明其造血功能受到抑制。模型小鼠灌胃给予蕨麻提取物中剂量和高剂量对骨髓腔造血细胞减少具有显著的改善作用, 表明蕨麻水提物可缓解化疗药物环磷酰胺导致的骨髓组织损伤。
5 蕨麻对骨髓抑制小鼠血清中SOD、CAT及MDA的影响
如表 4所示, 与空白对照组比较, 模型对照组小鼠血清中SOD及CAT水平显著降低(P < 0.01), MDA水平显著增高(P < 0.01), 提示环磷酰胺导致了小鼠的过氧化损伤。与模型组比较, 蕨麻中、高剂量组小鼠血清SOD、CAT水平显著增高, MDA水平显著降低(P < 0.05), 表明蕨麻可能通过调节氧化与抗氧化平衡发挥促造血功能恢复作用。
6 蕨麻对骨髓抑制小鼠血清中造血相关细胞因子的影响
各组小鼠血清中GM-CSF、IL-6、EPO及IL-3水平检测结果如表 5所示。与空白组比较, 模型组小鼠血清中GM-CSF、IL-6水平显著降低(P < 0.01), EPO、IL-3水平无显著性差异(P > 0.05)。与模型对照组比较, 阳性对照组和蕨麻中、高剂量组小鼠血清中GM-CSF、IL-6水平显著升高(P < 0.05), EPO、IL-3水平无显著性变化(P > 0.05), 提示蕨麻的促造血活性可能与调节部分血细胞分化刺激因子有关。
7 代谢组学分析
7.1 代谢轮廓分析
上述药效研究结果显示, 蕨麻高剂量对环磷酰胺诱导的骨髓抑制小鼠的外周血象、氧化应激及造血相关细胞因子均有显著的改善作用。因此, 选择蕨麻高剂量组、空白组和模型组样本开展非靶向血清代谢组学分析。借助PCA和PLS-DA对各组样本开展代谢轮廓分析。如图 2所示, PCA结果表明, 在正、负离子模式下QC样本聚集度较高, 说明本实验所建立的方法稳定性和重复性较好, 且各实验组间均呈现区分趋势, 组内各样本在一定范围内呈现聚集。同时, PLS-DA分析结果同样提示三组间血清代谢轮廓明显分离。对数据模型进行200次置换检验, 得出正离子模式下R2 = 0.681及Q2 = -0.112, 负离子模式下R2 = 0.719及Q2 = -0.092 1, 表明模型未产生过度拟合。
7.2 差异代谢物的筛选
蕨麻提取物改善环磷酰胺诱导的骨髓抑制小鼠的主要差异代谢物如表 6所示。以VIP > 1同时P < 0.05为差异代谢物筛选条件, 通过代谢物一级、二级质谱数据, 结合HMDB数据库及相关文献进行血清代谢物定性鉴别。在正、负离子模式下, 模型组与空白组相比显著上调24个、下调35个; 蕨麻高剂量组与模型组相比显著上调40个、下调15个。将模型组与空白组相比有显著变化且可被蕨麻提取物显著回调的视为差异代谢物, 共计13个。这些结果提示, 蕨麻可引起骨髓抑制病理状态下的多种代谢物向正常水平回调, 提示其可能干预相关代谢物水平及代谢途径改善小鼠骨髓造血功能。
7.3 代谢通路富集分析
将上述筛选得到的13个差异代谢物导入MetaboAnalyst 5.0平台, 进一步分析蕨麻改善骨髓抑制小鼠可能涉及的代谢通路。如图 3所示, 差异代谢物主要涉及氨酰-tRNA生物合成、缬氨酸, 亮氨酸和异亮氨酸的生物合成及降解、苯丙氨酸, 酪氨酸和色氨酸的生物合成、糖基磷脂酰肌醇-anchor的生物合成等通路, 提示蕨麻提取物可能通过调控上述代谢通路改善环磷酰胺诱导的骨髓抑制。
讨论
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环磷酰胺作为一种烷化剂类广谱抗肿瘤药, 常用于多种恶性肿瘤临床化疗, 其抑制骨髓造血系统功能使得患者外周血细胞数目下降[11]。因此, 环磷酰胺诱导的骨髓抑制动物模型常用于药物对造血系统功能障碍的影响与机制研究。本研究中, 连续腹腔注射环磷酰胺可导致小鼠外周血中WBC、RBC、HGB及PLT等显著下降, 蕨麻高剂量显著改善模型小鼠外周血中WBC、RBC及HGB水平。骨髓有核细胞数检测可反映骨髓中粒细胞、巨核细胞、淋巴细胞、单核细胞等具有细胞核的相关细胞数目, 可用于衡量骨髓增生程度[12]。蕨麻中、高剂量可显著改善模型小鼠的骨髓有核细胞数降低。同时, 骨髓组织病理学分析显示蕨麻提取物可改善环磷酰胺诱导的小鼠骨髓造血组织面积减少和造血细胞损伤。综合以上药效学数据, 提示了蕨麻对环磷酰胺诱导的小鼠骨髓造血功能障碍具有改善作用。
环磷酰胺体内可诱导线粒体氧化应激, 继而导致活性氧产生, 启动下游多级反应, 致使氧化与抗氧化失衡, 最终导致细胞凋亡[13]。氧化应激损伤是环磷酰胺诱发骨髓抑制的重要生物学基础之一。前期研究表明了蕨麻多酚及多糖等主要成分具有显著的抗自由基氧化作用[14, 15]。本研究结果显示蕨麻中、高剂量组可以明显提升骨髓抑制小鼠血清中SOD、CAT活力, 并降低MDA水平, 提示其促进骨髓造血功能恢复作用可能与调节机体氧化应激有关。此外, 骨髓造血细胞增殖、分化、成熟与多种促造血生长因子调控有关, 主要包括GM-CSF、EPO、IL-3、IL-6和IL-1β等[16]。GM-CSF主要由骨髓基质细胞分泌产生, 对早期髓样祖细胞具有广泛的促增殖作用, 并刺激髓样祖细胞分化为中性粒细胞和单核细胞[17]。GM-CSF还可诱导IL-3分泌, 二者协同促进嗜碱性细胞生成, 进而提升中性粒细胞与单核细胞数量[18]。IL-6同样作为促造血细胞因子, 亦可由骨髓基质细胞分泌, 促进巨核细胞分化成熟为血小板[19]。本研究结果显示, 模型小鼠血清中GM-CSF及IL-6水平明显降低, EPO及IL-3无显著变化, 表明环磷酰胺导致了部分造血细胞因子生成抑制。蕨麻中、高剂量组小鼠血清中GM-CSF及IL-6水平得到显著升高, 提示蕨麻可能通过促造血生长因子表达从而起到正向调节外周血细胞生成作用。
血清非靶向代谢组学结果表明蕨麻提取物可调控缬氨酸、天冬酰胺、苯基丙氨酸、PE(18∶0/18∶2(9Z, 12Z))、苯丙氨酸等内源性物质参与骨髓造血功能损伤相关的氨基酸代谢、甘油磷脂代谢和氨酰-tRNA生物合成等代谢途径。已有研究表明, 缬氨酸可增强造血祖细胞集落形成, 增加小鼠骨髓和人脐带血中造血祖细胞的体外存活率[20]。同时, 缬氨酸饮食限制可清空小鼠骨髓微环境, 从而为供体造血干细胞移植提供了无需化学和放射清髓的骨髓抑制状态[21]。这些研究结果均表明了缬氨酸对于骨髓造血干/祖细胞的增殖和功能维持起到重要作用。亮氨酸作为一种必需氨基酸, 在蛋白合成中具有重要作用。饮食补充亮氨酸可显著提高先天性纯红细胞再生障碍性贫血小鼠的红细胞数目和血红蛋白含量, 其作用机制可能与降低造血祖细胞中p53活力相关[22]。临床研究表明亮氨酸可升高老年骨髓异常增生综合征患者的绝对网织红细胞数目, 进而促进造血功能恢复[23]。L-天冬酰胺主要涉及天冬氨酸代谢通路, 研究表明造血干细胞可通过增加天冬氨酸合成, 使得天冬酰胺和嘌呤合成, 进而增强造血干细胞功能[24]。本研究结果表明蕨麻提取物可显著逆转环磷酰胺诱导的骨髓抑制小鼠血清中缬氨酸、亮氨酸和L-天冬酰胺水平, 提示了蕨麻可能通过调控多种氨基酸合成和代谢促进造血功能修复。此外, 蕨麻提取物亦可调控模型小鼠血清中多种磷脂酰乙醇胺及溶血磷脂酰乙醇胺含量, 该类物质所介导的甘油磷脂代谢途径在蕨麻补血功能中的作用机制仍有待进一步探索。
综上所述, 本文证实了藏药蕨麻水提物可体内改善化疗药物环磷酰胺诱导的骨髓抑制, 显著促进造血系统功能修复, 其作用机制可能与蕨麻中多种成分的协同抗氧化和促造血生长因子分泌及调控相关氨基酸代谢有关。本研究为蕨麻补血功效的现代科学内涵提供了一定的数据支持, 并为其进一步的物质基础和作用机制研究提供科学依据。
作者贡献: 龚普阳负责实验设计并参与论文撰写; 刘静贤和黄时光负责实验操作和论文初稿撰写; 罗晓敏和刘伟负责数据处理和图表制作; 顾健和王琴负责指导论文修改。
利益冲突: 本文无利益冲突。
基金
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  • 西南民族大学中央高校基本科研业务费(2020NQN38)
  • 西南民族大学大学生创新创业训练计划项目(202210656042)
文献
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2023年第58卷第7期
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doi: 10.16438/j.0513-4870.2022-1258
  • 接收时间：2022-11-23
  • 首发时间：2025-11-21
  • 出版时间：2023-07-12
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  • 收稿日期：2022-11-23
  • 修回日期：2023-01-23
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西南民族大学中央高校基本科研业务费(2020NQN38)
西南民族大学大学生创新创业训练计划项目(202210656042)
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    西南民族大学药学院, 四川 成都 610041

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*刘伟, E-mail: ;
龚普阳, E-mail:
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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