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Article(id=1308842017694769605, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, articleNumber=null, orderNo=null, doi=10.16438/j.0513-4870.2025-1587, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1766937600000, receivedDateStr=2025-12-29, revisedDate=1772899200000, revisedDateStr=2026-03-08, acceptedDate=null, acceptedDateStr=null, onlineDate=1789981854126, onlineDateStr=2026-09-21, pubDate=1783785600000, pubDateStr=2026-07-12, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789981854126, onlineIssueDateStr=2026-09-21, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789981854126, creator=13701087609, updateTime=1789981854126, updator=13701087609, issue=Issue{id=1308841897058198067, tenantId=1146029695717560320, journalId=1189982191388893191, year='2026', volume='61', issue='7', pageStart='2377', pageEnd='2820', issueExtLink='null', onlineDate='null', pubDate='1783785600000', pubDateStr='2026-07-12', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789981825365, creator='13701087609', updateTime=1789984154811, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1308851667525792021, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1308851667525792022, tenantId=1146029695717560320, journalId=1189982191388893191, issueId=1308841897058198067, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=2523, endPage=2532, ext={EN=ArticleExt(id=1308842017875124678, articleId=1308842017694769605, tenantId=1146029695717560320, journalId=1189982191388893191, language=EN, title=The role of TAAR1/5-HT2CR in CUMS-induced anxiety-like behavior in rats and intervention with dual agonists, columnId=1190335348761793317, journalTitle=Acta Pharmaceutica Sinica, columnName=Original Articles, runingTitle=null, highlight=null, articleAbstract=

Anxiety disorders affect over 3.3% of the global population, with limited treatment options available. Both trace amine-associated receptor 1 (TAAR1) and serotonergic 2C receptor (5-HT2CR) are implicated in various emotional activities, including anxiety. This study aims to investigate the role of TAAR1/5-HT2CR in the development of anxiety-like behavior in rats induced by chronic unpredictable mild stress (CUMS), and the intervention effects of a dual agonist. A CUMS-induced rat anxiety model was established. This animal research protocol has been approved by the Yantai University Laboratory Animal Ethics Committee (animal ethics number: YTU20240110). Anxiety-like behavior was assessed using the elevated zero-maze test and open field test. Western blot analysis was employed to detect the expression of TAAR1, 5-HT2CR, and brain-derived neurotrophic factor (BDNF) proteins in the rat amygdala. To observe the effects of Taar1 and Htr2c gene on anxiety-like behavior in rats, we knocked down or overexpressed these genes in the rat basolateral amygdala (BLA). Results demonstrated that CUMS-induced anxiety significantly reduced the expression levels of TAAR1 and 5-HT2CR in the amygdala of rats. Overexpression of either Taar1 or Htr2c in the BLA ameliorated anxiety-like behavior. Conversely, knocking down Taar1 or Htr2c in the BLA exacerbated CUMS-induced anxiety-like behavior. Oral administration of PCC0105004 alleviated anxiety-like behavior in CUMS rats, while knocking down both Taar1 and Htr2c in the BLA completely abolished this effect of PCC0105004. Further investigations revealed that PCC0105004 significantly increased the expression of BDNF in the amygdala of rats. These findings indicate that TAAR1 and 5-HT2CR play crucial roles in the development of anxiety-like behavior in CUMS rats, suggesting that a dual agonist targeting TAAR1/5-HT2CR may offer a novel therapeutic strategy for anxiety disorders.

, authors=Xin-xin MU1, Qian WANG1, Min XU1, Ying-tian ZHANG1, Sheng-min JI1, Jing-wei TIAN1, 2, Chun-mei LI1, 2, *, authorsList=Xin-xin MU, Qian WANG, Min XU, Ying-tian ZHANG, Sheng-min JI, Jing-wei TIAN, Chun-mei LI, authorCompany=null, correspAuthors=Chun-mei LI, authorNote=null, correspAuthorsNote=null, copyrightStatement=Copyright ©2026 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, fund=null), CN=ArticleExt(id=1308842019271827925, articleId=1308842017694769605, tenantId=1146029695717560320, journalId=1189982191388893191, language=CN, title=TAAR1/5-HT2CR在CUMS诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预研究, columnId=1190335348896011050, journalTitle=药学学报, columnName=研究论文, runingTitle=null, highlight=null, articleAbstract=

焦虑症影响着全球超过3.3%的人口, 且治疗选择有限, 而痕量胺相关受体1 (trace amine-associated receptor 1, TAAR1) 与5-羟色胺2C受体(serotonergic 2C receptor, 5-HT2CR) 可参与焦虑等多种情绪活动。本文旨在研究TAAR1/5-HT2CR在慢性不可预见性温和应激(chronic unpredictable mild stress, CUMS) 诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预作用。本研究构建CUMS诱导的大鼠焦虑模型[本实验方案获得烟台大学实验动物管理委员会批准(批准号: YTU20240110)], 采用高架O迷宫试验和旷场试验评价大鼠焦虑样行为; 采用Western blot检测大鼠杏仁核中TAAR1、5-HT2CR和脑源性神经营养因子(brain-derived neurotrophic factor, BDNF) 蛋白的表达; 分别敲低或过表达大鼠基底外侧杏仁核(basolateral amygdala, BLA) 中Taar1和Htr2c基因, 观察其对大鼠焦虑样行为的影响; 大鼠灌胃给予TAAR1/5-HT2CR双重激动剂PCC0105004, 同时敲低BLA中Taar1和Htr2c基因, 观察其对大鼠的焦虑样行为的影响。结果显示, CUMS诱导的焦虑大鼠杏仁核中TAAR1和5-HT2CR表达水平显著降低。过表达BLA中Taar1或Htr2c基因可改善大鼠的焦虑样行为。相反, 敲低BLA中Taar1或Htr2c基因可加剧CUMS诱导的大鼠焦虑样行为。灌胃给予PCC0105004可缓解CUMS大鼠的焦虑样行为, 敲低BLA中Taar1和Htr2c基因可完全消除PCC0105004的上述作用。进一步研究显示, PCC0105004可显著增加杏仁核中BDNF的表达。结果表明, TAAR1和5-HT2CR在CUMS大鼠焦虑样行为发生中发挥重要作用, TAAR1/5-HT2CR双重激动剂可能为焦虑症的治疗提供一种新的治疗策略。

, authors=穆鑫鑫1, 王茜1, 徐敏1, 张颖甜1, 汲胜民1, 田京伟1, 2, 李春梅1, 2, *, authorsList=穆鑫鑫, 王茜, 徐敏, 张颖甜, 汲胜民, 田京伟, 李春梅, authorCompany=null, correspAuthors=李春梅, authorNote=

作者贡献:

李春梅、田京伟负责课题设计、实验指导和文章修订; 穆鑫鑫负责实验实施、结果分析、论文撰写和修改; 王茜、徐敏、张颖甜、汲胜民负责部分实验和结果统计与分析。

, correspAuthorsNote=
李春梅, Tel: 13688696218, E-mail:
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A: Experimental workflow; B-E: Following CUMS modeling, rats exhibited open arm residence time (B) and open arm entries (C) in the elevated zero-maze test, central zone residence time (D) and total locomotor distance (E) in the open field test; F, G: Protein expression levels of TAAR1 (F) and 5-HT2CR (G) in the amygdala of rats 21 days after CUMS modeling. n = 10 (B-E) and 6 (F, G), mean ± SEM. *P < 0.05, **P < 0.01, ***P < 0.001. 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A: Experimental workflow; B-D: Following stereotactic injection of the Taar1 knockdown virus and CUMS modeling, rats exhibited open arm residence time (B) in the elevated zero-maze test and central zone residence time (C) in the open field test, and expression of Taar1 mRNA (D) in each group; E-G: Following stereotactic injection of the Htr2c knockdown virus and CUMS modeling, rats exhibited open arm residence time (E) in the elevated zero-maze test and central zone residence time (F) in the open field test, and expression of Htr2c mRNA (G) in each group. n = 10 (B, C, E, F) and 6 (D, G), mean ± SEM. *P < 0.05, ***P < 0.001. AAV: Adeno-associated virus, figureFileSmall=c4L7Hl9krFENQgHl57X+Ag==, figureFileBig=s7mJ2cyIi2kW9+g1lrjSdQ==, tableContent=null), ArticleFig(id=1308842026620248593, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=EN, label=null, caption=null, figureFileSmall=IeFMvoxGR4I34dRuMdouPQ==, figureFileBig=kV0nadajUkOwup/iKm0siA==, tableContent=null), ArticleFig(id=1308842026704134674, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=CN, label=Fig.4, caption=mRNA and protein expression levels of Taar1 or Htr2c in the amygdala of rats 15 days after viral vector injection. A: Experimental workflow; B, C: Expression of Taar1 (B) and Htr2c (C) mRNA in each group; D-G: Expression of TAAR1 (D, E) and 5-HT2CR (F, G) protein in each group. n = 6, mean ± SEM. **P < 0.01, ***P < 0.001; #P < 0.05, ##P < 0.01, figureFileSmall=IeFMvoxGR4I34dRuMdouPQ==, figureFileBig=kV0nadajUkOwup/iKm0siA==, tableContent=null), ArticleFig(id=1308842027027096083, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=EN, label=null, caption=null, figureFileSmall=LpKmJg7gwad7vaJHXEdSkg==, figureFileBig=AWVhJaSLhfDYfRvfN/PyuA==, tableContent=null), ArticleFig(id=1308842027119370772, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=CN, label=Fig.5, caption=Effects of Taar1 or Htr2c overexpression in the BLA on CUMS-induced anxiety-like behavior in rats. A: Experimental workflow; B-E: Following stereotactic injection of the Taar1 overexpression virus and CUMS modeling, rats exhibited open arm residence time (B) and open arm entries (C) in the elevated zero-maze test, central zone residence time (D) and total locomotor distance (E) in the open field test; F-I: Following stereotactic injection of the Htr2c overexpression virus and CUMS modeling, rats exhibited open arm residence time (F) and open arm entries (G) in the elevated zero-maze test, central zone residence time (H) and total locomotor distance (I) in the open field test. n = 10, mean ± SEM. ***P < 0.001; #P < 0.05, ##P < 0.01, ###P < 0.001. 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A: Experimental workflow; B-E: Following stereotactic injection of the Taar1 knockdown virus and CUMS modeling, rats exhibited open arm residence time (B) and open arm entries (C) in the elevated zero-maze test, central zone residence time (D) and total locomotor distance (E) in the open field test; F-I: Following stereotactic injection of the Htr2c knockdown virus and CUMS modeling, rats exhibited open arm residence time (F) and open arm entries (G) in the elevated zero-maze test, central zone residence time (H) and total locomotor distance (I) in the open field test. n = 10, mean ± SEM. *P < 0.05, ***P < 0.001; #P < 0.05, figureFileSmall=/3yPiWrzTtgbSXJWNf+cRw==, figureFileBig=pJhK3VWfcE5snYVTikrAtA==, tableContent=null), ArticleFig(id=1308842027568161303, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=EN, label=null, caption=null, figureFileSmall=nffD0vwRaOs2Z1mv8u93hg==, figureFileBig=mpRf9zv2239FyTsZu3Au0Q==, tableContent=null), ArticleFig(id=1308842027639464472, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=CN, label=Fig.7, caption=Effects of PCC0105004 on anxiety-like behaviors in CUMS-induced anxious rats following knockdown of Taar1 and Htr2c in the BLA. A: Experimental workflow; B-E: Following stereotactic injection of the Taar1 and Htr2c knockdown virus and CUMS modeling, rats exhibited open arm residence time (B) and open arm entries (C) in the elevated zero-maze test, central zone residence time (D) and total locomotor distance (E) in the open field test; F, G: Protein expression levels of BDNF in the amygdala of rats. n = 10 (B-E) and 6 (F, G), mean ± SEM. ***P < 0.001; #P < 0.05, ###P < 0.001. BDNF: Brain-derived neurotrophic factor, figureFileSmall=nffD0vwRaOs2Z1mv8u93hg==, figureFileBig=mpRf9zv2239FyTsZu3Au0Q==, tableContent=null), ArticleFig(id=1308842027853373977, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
GeneGene sequence (5'-3')
AAV-shTaar1-1CGCCACAAAGCAAGGAAACA
AAV-shTaar1-2GCTGTGTGCGACCTTTAAGA
AAV-shTaar1-3GGAACTCTTCTGCAAACTTC
AAV-shHtr2c-1GCAACATTCTTGTTATCATGG
AAV-shHtr2c-2GCAAAGTGTTCGTGAATAACA
AAV-shHtr2c-3GCCGTCAAACTCTGATGTTAC
AAV-shNCCGCTGAGTACTTCGAAATGTC
), ArticleFig(id=1308842027933065754, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=CN, label=Tab.1, caption=

shTaar, shHtr2c, shNC gene sequences. Taar1: Trace amine-associated receptor 1; Htr2c: Serotonergic 2C receptor

, figureFileSmall=null, figureFileBig=null, tableContent=
GeneGene sequence (5'-3')
AAV-shTaar1-1CGCCACAAAGCAAGGAAACA
AAV-shTaar1-2GCTGTGTGCGACCTTTAAGA
AAV-shTaar1-3GGAACTCTTCTGCAAACTTC
AAV-shHtr2c-1GCAACATTCTTGTTATCATGG
AAV-shHtr2c-2GCAAAGTGTTCGTGAATAACA
AAV-shHtr2c-3GCCGTCAAACTCTGATGTTAC
AAV-shNCCGCTGAGTACTTCGAAATGTC
), ArticleFig(id=1308842028004368923, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=EN, label=null, caption=null, figureFileSmall=null, figureFileBig=null, tableContent=
GeneForward (5'-3')Reverse (5'-3')
Taar1CGCCACAAAGCAAGGAAACACGCCACAAAGCAAGGAAACA
Htr2cATTTGTGCCCCGTCTGGATTATTTGTGCCCCGTCTGGATT
GAPDHGGTCGGAGTCAACGGATTTGGGTCGGAGTCAACGGATTTG
), ArticleFig(id=1308842028067283484, tenantId=1146029695717560320, journalId=1189982191388893191, articleId=1308842017694769605, language=CN, label=Tab.2, caption=

Primer sequences

, figureFileSmall=null, figureFileBig=null, tableContent=
GeneForward (5'-3')Reverse (5'-3')
Taar1CGCCACAAAGCAAGGAAACACGCCACAAAGCAAGGAAACA
Htr2cATTTGTGCCCCGTCTGGATTATTTGTGCCCCGTCTGGATT
GAPDHGGTCGGAGTCAACGGATTTGGGTCGGAGTCAACGGATTTG
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TAAR1/5-HT2CR在CUMS诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预研究
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穆鑫鑫 1 , 王茜 1 , 徐敏 1 , 张颖甜 1 , 汲胜民 1 , 田京伟 1, 2 , 李春梅 1, 2, *
药学学报 | 研究论文 2026,61(7): 2523-2532
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药学学报 |研究论文 2026 , 61 (7) : 2523 -2532
TAAR1/5-HT2CR在CUMS诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预研究
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穆鑫鑫1, 王茜1, 徐敏1, 张颖甜1, 汲胜民1, 田京伟1, 2, 李春梅1, 2, *
作者信息
  • 1烟台大学药学院, 分子药理和药物评价教育部重点实验室(烟台大学), 新型制剂与生物技术药物研究山东省高校协同创新中心, 山东 烟台 264005
  • 2先进药物递释系统全国重点实验室, 山东 烟台 264003
通讯作者:
李春梅, Tel: 13688696218, E-mail:
作者简介:

作者贡献:

李春梅、田京伟负责课题设计、实验指导和文章修订; 穆鑫鑫负责实验实施、结果分析、论文撰写和修改; 王茜、徐敏、张颖甜、汲胜民负责部分实验和结果统计与分析。

The role of TAAR1/5-HT2CR in CUMS-induced anxiety-like behavior in rats and intervention with dual agonists
Xin-xin MU1, Qian WANG1, Min XU1, Ying-tian ZHANG1, Sheng-min JI1, Jing-wei TIAN1, 2, Chun-mei LI1, 2, *
Affiliations
  • 1School of Pharmacy, Key Laboratory of Molecular Pharmacology and Drug Evaluation (Yantai University), Ministry of Education, Collaborative Innovation Center of Advanced Drug Delivery System and Biotech Drugs in Universities of Shandong, Yantai University, Yantai 264005, China
  • 2State Key Laboratory of Advanced Drug Delivery and Release Systems, Yantai 264003, China
出版时间: 2026-07-12 doi: 10.16438/j.0513-4870.2025-1587
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焦虑症影响着全球超过3.3%的人口, 且治疗选择有限, 而痕量胺相关受体1 (trace amine-associated receptor 1, TAAR1) 与5-羟色胺2C受体(serotonergic 2C receptor, 5-HT2CR) 可参与焦虑等多种情绪活动。本文旨在研究TAAR1/5-HT2CR在慢性不可预见性温和应激(chronic unpredictable mild stress, CUMS) 诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预作用。本研究构建CUMS诱导的大鼠焦虑模型[本实验方案获得烟台大学实验动物管理委员会批准(批准号: YTU20240110)], 采用高架O迷宫试验和旷场试验评价大鼠焦虑样行为; 采用Western blot检测大鼠杏仁核中TAAR1、5-HT2CR和脑源性神经营养因子(brain-derived neurotrophic factor, BDNF) 蛋白的表达; 分别敲低或过表达大鼠基底外侧杏仁核(basolateral amygdala, BLA) 中Taar1和Htr2c基因, 观察其对大鼠焦虑样行为的影响; 大鼠灌胃给予TAAR1/5-HT2CR双重激动剂PCC0105004, 同时敲低BLA中Taar1和Htr2c基因, 观察其对大鼠的焦虑样行为的影响。结果显示, CUMS诱导的焦虑大鼠杏仁核中TAAR1和5-HT2CR表达水平显著降低。过表达BLA中Taar1或Htr2c基因可改善大鼠的焦虑样行为。相反, 敲低BLA中Taar1或Htr2c基因可加剧CUMS诱导的大鼠焦虑样行为。灌胃给予PCC0105004可缓解CUMS大鼠的焦虑样行为, 敲低BLA中Taar1和Htr2c基因可完全消除PCC0105004的上述作用。进一步研究显示, PCC0105004可显著增加杏仁核中BDNF的表达。结果表明, TAAR1和5-HT2CR在CUMS大鼠焦虑样行为发生中发挥重要作用, TAAR1/5-HT2CR双重激动剂可能为焦虑症的治疗提供一种新的治疗策略。

焦虑  /  基底外侧杏仁  /  痕量胺相关受体1  /  5-羟色胺2C受体  /  PCC0105004

Anxiety disorders affect over 3.3% of the global population, with limited treatment options available. Both trace amine-associated receptor 1 (TAAR1) and serotonergic 2C receptor (5-HT2CR) are implicated in various emotional activities, including anxiety. This study aims to investigate the role of TAAR1/5-HT2CR in the development of anxiety-like behavior in rats induced by chronic unpredictable mild stress (CUMS), and the intervention effects of a dual agonist. A CUMS-induced rat anxiety model was established. This animal research protocol has been approved by the Yantai University Laboratory Animal Ethics Committee (animal ethics number: YTU20240110). Anxiety-like behavior was assessed using the elevated zero-maze test and open field test. Western blot analysis was employed to detect the expression of TAAR1, 5-HT2CR, and brain-derived neurotrophic factor (BDNF) proteins in the rat amygdala. To observe the effects of Taar1 and Htr2c gene on anxiety-like behavior in rats, we knocked down or overexpressed these genes in the rat basolateral amygdala (BLA). Results demonstrated that CUMS-induced anxiety significantly reduced the expression levels of TAAR1 and 5-HT2CR in the amygdala of rats. Overexpression of either Taar1 or Htr2c in the BLA ameliorated anxiety-like behavior. Conversely, knocking down Taar1 or Htr2c in the BLA exacerbated CUMS-induced anxiety-like behavior. Oral administration of PCC0105004 alleviated anxiety-like behavior in CUMS rats, while knocking down both Taar1 and Htr2c in the BLA completely abolished this effect of PCC0105004. Further investigations revealed that PCC0105004 significantly increased the expression of BDNF in the amygdala of rats. These findings indicate that TAAR1 and 5-HT2CR play crucial roles in the development of anxiety-like behavior in CUMS rats, suggesting that a dual agonist targeting TAAR1/5-HT2CR may offer a novel therapeutic strategy for anxiety disorders.

anxiety disorder  /  basolateral amygdala  /  trace amine-associated receptor 1  /  serotonergic 2C receptor  /  PCC0105004
穆鑫鑫, 王茜, 徐敏, 张颖甜, 汲胜民, 田京伟, 李春梅. TAAR1/5-HT2CR在CUMS诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预研究. 药学学报, 2026 , 61 (7) : 2523 -2532 . DOI: 10.16438/j.0513-4870.2025-1587
Xin-xin MU, Qian WANG, Min XU, Ying-tian ZHANG, Sheng-min JI, Jing-wei TIAN, Chun-mei LI. The role of TAAR1/5-HT2CR in CUMS-induced anxiety-like behavior in rats and intervention with dual agonists[J]. Acta Pharmaceutica Sinica, 2026 , 61 (7) : 2523 -2532 . DOI: 10.16438/j.0513-4870.2025-1587
正文
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焦虑症是全球范围内最常见且最具致残性的精神疾病之一。其高患病率、慢性化及高共病率促使其成为健康相关致残的主要原因之一[1]。在全球范围内, 焦虑症对患者及社会造成严重影响, 占全球疾病负担的3.3%[2]。尽管心理治疗和社会支持在焦虑治疗中至关重要, 但焦虑的治疗仍主要依赖于以下经典抗焦虑药物: 选择性5-羟色胺再摄取抑制剂(selective serotonin reuptake inhibitors, SSRIs)、5-羟色胺去甲肾上腺素再摄取抑制剂(serotonin norepinephrine reuptake inhibitors, SNRIs)、三环类抗抑郁药物(tricyclic antidepressants, TCAs)、苯二氮䓬类(benzodiazepines, BZDs) 及单胺氧化酶抑制剂(monoamine oxidase inhibitors, MAOIs)[3]。由于耐受性、依赖性[3]及药物存在起效慢(2~4周)、有效率低、个体差异大、不良反应大(体重增加、胃肠道刺激等)、认知损伤等问题, 这些药物无法充分发挥治疗作用[4, 5]。尽管如此, 过去5~10年间针对焦虑症的新型治疗药物的研究远远少于针对重度抑郁障碍、双相情感障碍和精神分裂症的药物的研究。因此, 深入揭示抗焦虑的新机制, 寻找新型抗焦虑药物靶点, 是精神科学领域有待解决的关键问题。
焦虑相关信息的处理涉及大脑区域(包括前额叶皮层、边缘系统和中脑边缘系统), 在这些神经结构中, 杏仁核——尤其是基底外侧杏仁核(basolateral amygdala, BLA)——对焦虑调节具有至关重要的作用[6-8]。杏仁核中的神经元对压力和焦虑诱发的刺激高度敏感, 该区域接收来自丘脑和感觉皮层关于感官刺激的兴奋性传入信号, 对焦虑情绪的形成具有决定性影响[9, 10]。
痕量胺相关受体1 (trace amine-associated receptor 1, TAAR1) 是一种G蛋白偶联受体, 在多种哺乳动物脑区广泛表达, 包括前额叶皮层和中脑边缘系统区域(海马体和杏仁核), 这些区域对情绪调节、认知和行为至关重要[11]。此外, 临床前研究表明, TAAR1的内源性配体及该受体的选择性激动剂均表现出抗焦虑样作用[12]。5-羟色胺(serotonergic, 5-HT) 系统是治疗焦虑症最广泛应用的药理靶点[13], 5-HT广泛分布于中枢神经系统, 在调控情绪、认知及记忆等生物学过程中发挥了关键作用[14]。5-羟色胺2C受体(serotonergic 2C receptor, 5-HT2CR) 主要分布在边缘系统和基底节等脑区[14, 15], 在5-HT2CR表达的大脑区域中, 杏仁核被认为是其介导焦虑相关行为的关键神经基质[16]。
(S)-4, 7, 8, 9, 10, 10a-六氢-5H-噻吩并[2', 3': 3, 4]吡啶并[1, 2-a]吡嗪(PCC0105004) 是一种新型的TAAR1/5-HT2CR双重激动剂[17]。药代动力学评估显示, 大鼠给药0.25~1 h内PCC0105004在血浆和脑中的浓度达到峰值, 脑/血浆比值范围为8.45~4.22, 表明其具有良好的药代动力学特性[18]。
在本研究中, 采用遗传学[短发夹RNA (short hairpin RNA, shRNA) 敲低或过表达] 和药理学(TAAR1/5-HT2CR双重激动剂PCC0105004) 方法, 研究TAAR1和5-HT2CR在慢性不可预见性温和应激(chronic unpredictable mild stress, CUMS) 诱导的大鼠焦虑样行为发生中的作用及其双重激动剂的干预作用。
材料与方法
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实验动物  选用Sprague-Dawley (SD) 雄性大鼠(6~7周龄), 平均体重180~220 g, 由北京维通利华实验动物技术有限公司提供, 动物生产许可证号: SCXK (京) 2021-0011。将动物饲养在SPF级标准温控动物房(饲养条件: 20~24 ℃, 40%~60%湿度, 12 h光照/黑暗循环) 内, 自由进食和饮水。动物模型建立及病毒注射在动物适应环境7天后进行。本研究采用的实验方法均已获得烟台大学实验动物伦理委员会批准(批准号: YTU20240110)。
药物与试剂  PCC0105004 (C10H14N2S, 图1) 纯度为95%, 购自天津药明康德新药开发有限公司; 10%、12.5%一步法PAGE彩色凝胶极速配制试剂盒(山东思科捷生物技术有限公司, 货号: EC0023-A、EC0024-A); 磷酸盐缓冲液、TBS干粉缓冲液、Tris-甘氨酸SDS-PAGE SWE高分辨快速电泳缓冲液、Tris甘氨酸转膜缓冲液(干粉) (武汉塞维尔生物科技有限公司, 货号: G4202-100ML、G0001-2L、G2081-1L、G2017-1L); 蛋白标准、RIPA裂解液(强)、BCA蛋白浓度测定试剂盒、鼠抗GAPDH抗体、鼠抗β-actin抗体、山羊抗鼠HRP、山羊抗兔HRP (货号: P0007、P0013B、P0012、AF0006、AF0003、A0216、A0208) (上海碧云天生物技术有限公司); Power SYBRTM Green PCR预混液(美国Applied Biosystems公司, 货号: 4367659); 兔抗TAAR1、兔抗5-HT2CR、兔抗脑源性神经营养因子(brain-derived neuro-trophic factor, BDNF) (美国Immunoway Biotechnology公司, 货号: YN2695、YT4399、YT7890)。
主要仪器  高速冷冻微量离心机(德国Eppendorf公司, 型号: 5425R); 基础电泳+转印系统(美国BIO-RAD公司, 型号: 1645050); 转移脱色摇床(型号: TS-8)、干式恒温器(型号: GL-1900), 海门市其林贝尔仪器制造有限公司; 酶标仪(美国BioTek公司, 型号: Synergy NEO2); 荧光定量PCR系统(美国Applied Biosystems公司, 型号: 7500 Fast); 全自动化学发光凝胶成像系统(北京赛智创业科技有限公司, 型号: ChampChemi 610)。
CUMS模型建立  采用CUMS的方法建立大鼠焦虑模型[19]。除对照组外所有实验组大鼠均接受为期21天的CUMS处理, 具体应激源包括: 束缚(4 h)、笼体倾斜(45°持续24 h)、潮湿垫料(24 h)、禁食/禁水(24 h)、夹尾(距尾端1 cm处, 3 min)、冷水游泳(4 ℃, 3 min) 及昼夜颠倒(24 h)。建模期间, 实验组大鼠单笼饲养, 每日随机接受不同应激刺激; 对照组每笼饲养5只大鼠, 不施加任何刺激。
行为学测试  在所有行为测试过程中, 采用双盲设计, 即实验人员均不了解组别分配及药物处理情况。
旷场试验  采用60 cm×60 cm×50 cm (长×宽×高) 的箱体, 箱内划分为16个等面积区域, 其中央区域为4个中心方格。实验前, 将动物提前放置实验环境中2 h, 以适应环境。测试期间, 大鼠在箱内自由活动5 min, 全程保持安静, 通过TopScan监测系统记录其运动活动。记录指标包括总移动距离及中央区域停留时间。每次测试后, 使用75%酒精喷雾擦拭箱体以清除前一只动物的痕迹。
高架O迷宫测试  采用黑色圆形轨道(直径: 100 cm, 宽度: 10 cm), 距地面70 cm, 设有两条封闭通道和两条开放通道。实验前, 将动物提前放置实验环境中2 h, 以适应环境。实验时, 将大鼠放置在开放臂和封闭臂的交界处, 并且头部朝向开放臂方向, 通过TopScan监测系统记录5 min内开放臂停留时间及进入次数。每次测试后用75%酒精喷雾擦拭迷宫以清除前一只动物的痕迹。
腺相关病毒载体(adeno-associated virus, AAV) 构建  AAV介导的载体购自上海吉凯基因化学有限公司。研究选择易于转染神经系统的AAV9载体, 用于过表达和敲低Taar1 (Taar1基因编码TAAR1蛋白, GenBank: NM_134328.1) 和Htr2c (Htr2c基因编码5-HT2CR蛋白, GenBank: NM_012765.3) 基因。两个基因的编码序列经扩增后整合至GV388载体(CMV bGlobin-MCS-EGFP-3FLAG-WPRE-hGH polyA), 分别命名为AAV-Taar1和AAV-Htr2c。未编码目标基因的质粒作为对照, 命名为AAV-NC。针对大鼠Taar1或Htr2c的shRNA实验, 选用GV478载体(U6-MCS-CAG-EGFP) 构建shRNA。大鼠shRNA介导的Taar1基因敲低(AAV-shTaar1) 序列和大鼠shRNA介导的Htr2c基因敲低(AAV-shHtr2c) 序列如表1所示。以不含Taar1和Htr2c的shRNA的AAV作为对照组, 标记为AAV-shNC, 其序列如表1所示。
腺相关病毒载体注射  病毒注射时, 使用滴度为1012 vg·mL-1的病毒感染大鼠BLA区域。大鼠腹腔注射10%戊巴比妥钠(3.5 mL·kg-1) 麻醉后, 固定于脑立体定位仪上。以每侧2 µL、0.2 µL·min-1的速度缓慢注射到大鼠BLA区域(AP = -2.50 mm, ML = ±5.00 mm, DV = -8.20 mm)。病毒注入完成后, 将微量注射器原位停滞10 min, 随后缓慢回抽以最大限度减少病毒沿注射路径扩散。
Western blot检测  大鼠腹腔注射10%戊巴比妥钠(3.5 mL·kg-1), 麻醉后, 经心脏灌注磷酸盐缓冲液, 直至心脏呈现肉红色, 然后迅速分离取出杏仁核组织, 置于裂解缓冲液中充分研磨, 随后将杏仁核组织样本以12 000 r·min-1, 离心20 min。收集上清液, 使用BCA蛋白浓度测定试剂盒测定蛋白浓度。取50 µg蛋白样本经10%~12.5% SDS-PAGE凝胶电泳分离, 转移至聚偏氟乙烯膜进行免疫印迹分析。在5%脱脂牛奶中封闭2 h后, 与相应一抗4 ℃孵育过夜。经TBST缓冲液洗涤3次后, 加入二抗孵育2 h后, 使用全自动化学发光凝胶成像系统进行蛋白条带检测和成像。采用ImageJ软件量化条带强度, 并以GAPDH和β-actin为内参进行归一化处理。
RNA提取与RT-qPCR  使用RNA提取试剂盒提取RNA后, 采用逆转录试剂盒制备cDNA, 随后按说明书使用SYBR Green法结合Oligo dT引物进行qPCR定量。所用引物序列购自北京擎科生物科技股份有限公司, 如表2所示。每个10 µL荧光定量反应体系包含: 5 µL 2× SYBR qPCR Mix、0.5 µL核酸模板(DNA/cDNA)、各0.2 µL上游引物和下游引物、0.2 µL参比染料、3.9 µL无酶水。采用Power SYBRTM Green PCR预混液和Fast 7500实时PCR系统进行定量PCR, 设置参数为: 95 ℃预变性10 min; 40个循环: 95 ℃ 15 s、60 ℃ 60 s; 随后进行熔解曲线分析95 ℃ 15 s、60 ℃ 60 s、95 ℃ 30 s, 最后37 ℃冷却30 s。样本基因表达水平以GAPDH为内参进行归一化, 采用2-ΔΔCt法量化各组间的相对表达水平。
统计学分析  数据以均值±标准误(SEM) 表示, 采用IBM SPSS 21.0进行分析, 并使用GraphPad Prism 9.0绘制图表。两个独立样本比较采用独立样本t检验进行显著性分析; 多个独立样本比较采用单因素方差分析(ANOVA) 及Bonferroni事后检验进行比较, P < 0.05视为具有统计学意义。
结果
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1　CUMS诱导的焦虑大鼠杏仁核中TAAR1与5-HT2CR表达水平降低
焦虑模型建立、行为学测试及脑组织取材的时间线如图2A所示。如图2B、C所示, 与control相比, CUMS模型大鼠在高架O迷宫的开放臂停留时间显著减少(P < 0.001), 进入开放臂次数无显著差异; 如图2D、E所示, CUMS模型组大鼠在旷场的中心区域停留时间显著减少(P < 0.01), 总运动距离无显著差异。如图2F、G所示, WB结果显示, 与control组相比, CUMS诱导的焦虑大鼠杏仁核中TAAR1和5-HT2CR的表达水平显著降低(P < 0.01, P < 0.05)。上述结果均表明CUMS诱导的大鼠出现焦虑样行为, 且不影响模型组大鼠的自主活动。
2　 Taar1和Htr2c敲低序列选择
AAV-shRNA注射、焦虑模型建立、行为学测试及脑组织取材的时间线如图3A所示。分别设计了3组不同序列的shRNA干扰TAAR1和5-HT2CR的表达。含shTaar1和shHtr2c的AAV-9病毒(AAV-shTaar1-1、AAV-shTaar1-2、AAV-shTaar1-3、AAV-shHtr2c-1、AAV-shHtr2c-2及AAV-shHtr2c-3) 或阴性对照(AAV-shNC) 微量定位注射至大鼠BLA区域。如图3B~D所示, 与空白载体(AAV-shNC) 组相比, 3组shTaar1的大鼠在高架O迷宫开放臂停留时间没有显著变化, 在旷场中心区域停留时间呈缩短趋势, 其中AAV-shTaar1-1组缩短最为显著(P < 0.05); 在mRNA水平上, 各敲低序列均能显著减少Taar1的mRNA含量(P < 0.001)。最终选择AAV-shTaar1-1为最佳敲低序列(记作AAV-shTaar1) 用于后续研究。
如图3E~G所示, 与空白载体(AAV-shNC) 组相比, 3组shHtr2c的大鼠在高架O迷宫试验中开放臂停留时间和旷场中心区域停留时间呈缩短趋势, 其中AAV-shHtr2c-1组缩短最为显著(P < 0.001, P < 0.05); 在mRNA水平上, AAV-shHtr2c-1组显著减少了Htr2c的mRNA含量(P < 0.05)。最终选择AAV-shHtr2c-1 (记作AAV-shHtr2c) 为最佳敲低序列用于后续研究。
3　敲低或过表达Taar1或Htr2c的mRNA含量和蛋白表达水平验证
AAV-shRNA和AAV-RNA注射、焦虑模型建立、行为学测试及脑组织取材的时间线如图4A所示。
qRT-PCR结果(图4B、C) 表明, 与control组相比, 空白载体(AAV-shNC与AAV-NC) 组Taar1和Htr2c的mRNA含量无明显变化; 与空白载体(AAV-shNC) 组相比, AAV-shTaar1和AAV-shHtr2c组的mRNA表达水平显著降低(P < 0.001); 与空白载体(AAV-NC) 组相比, AAV-Taar1和AAV-Htr2c组的mRNA表达水平显著升高(P < 0.05)。
WB (图4D~G) 结果表明, 与control组相比, 空白载体(AAV-shNC与AAV-NC) 组TAAR1和5-HT2CR的蛋白表达无明显变化; 与空白载体(AAV-shNC) 组相比, AAV-shTaar1和AAV-shHtr2c组的蛋白表达水平显著降低(P < 0.001, P < 0.01); 与空白载体(AAV-NC) 组相比, AAV-Taar1和AAV-Htr2c组的蛋白表达水平显著升高(P < 0.01, P < 0.05)。
4　过表达Taar1或Htr2c对CUMS诱导的大鼠焦虑样行为的影响
AAV-RNA注射、焦虑模型建立、行为学测试的时间线如图5A所示。如图5B~E所示, 与空白载体(AAV-NC) 相比, AAV-Taar1组在高架O迷宫开放臂停留时间和旷场中心区域停留时间显著增加(P < 0.001, P < 0.05)。如图5F~I所示, 与空白载体(AAV-NC) 相比, AAV-Htr2c组在高架O迷宫开放臂停留时间和旷场中心区域停留时间显著增加(P < 0.01)。在各项实验中, 各组大鼠的总移动距离无显著差异。上述结果表明, 过表达Taar1或Htr2c可减轻CUMS诱导的大鼠的焦虑样行为, 且不影响其自主活动。
5　敲低Taar1和Htr2c对CUMS诱导的大鼠焦虑样行为的影响
AAV-shRNA注射、焦虑模型建立、行为学测试及脑组织取材的时间线如图6A所示。如图6B~E所示, 与空白载体(AAV-shNC) 组相比, AAV-shTaar1组大鼠在高架O迷宫开放臂停留时间显著减少(P < 0.05); 如图6F~I所示, 与空白载体(AAV-shNC) 组相比, AAV-shHtr2c组大鼠在旷场中心区域停留时间显著降低(P < 0.05)。在各项实验中, 各组大鼠总活动距离无显著差异。上述结果表明, 敲低Taar1和Htr2c可增强CUMS诱导焦虑大鼠的焦虑样行为, 且大鼠自主活动不受影响。
6　 Taar1和Htr2c双敲低后PCC0105004对CUMS诱导的大鼠焦虑样行为的影响
AAV-shRNA注射、焦虑模型建立、行为学测试及脑组织取材的时间线如图7A所示。如图7B、C所示, 与control组相比, CUMS组大鼠在高架O迷宫开放臂停留时间显著减少(P < 0.001), 灌胃(i.g.) 给予PCC0105004 (1 mg·kg-1) 可显著增加大鼠在开放臂停留时间(P < 0.05), 但无法改善AAV-shHT+PCC组大鼠在开放臂的时长。如图7D、E所示, 与control组相比, CUMS组大鼠在旷场中心区域停留时间显著减少(P < 0.001), 灌胃给予PCC0105004可显著增加大鼠在中心区域停留时长(P < 0.001), 但无法改善AAV-shHT+PCC组大鼠在中心区域的时长。上述结果表明, BLA脑区中敲低Taar1和Htr2c后, 1 mg·kg-1 PCC0105004对改善CUMS诱导的大鼠焦虑样行为没有显著影响。
鉴于PCC0105004为TAAR1和5-HT2CR的双重激动剂, 随后通过WB分析其对信号通路BDNF的影响, 如图7F、G所示, 与control组相比, CUMS组大鼠杏仁核中BDNF蛋白表达显著下降(P < 0.001), 灌胃给予1 mg·kg-1 PCC0105004后可显著增加BDNF的表达(P < 0.001)。结果表明, PCC0105004可能通过影响BDNF通路改善大鼠的焦虑样行为。
讨论
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焦虑症是一种以过度担忧、紧张和恐惧为核心症状的精神类疾病。其病理生理机制涉及神经递质失调、脑区功能异常、遗传因素等多层面复杂的相互作用。由于其治疗方法有限, 因此需要寻找新型的抗焦虑靶点。TAAR1广泛分布于大脑组织和周围组织, 与学习、记忆、情绪和成瘾性有关[20]。近年来, 对于TAAR1的研究取得了突破性进展, 大量证据表明, TAAR1已经成为精神类疾病治疗研发的新兴靶点, 与精神分裂症、抑郁症、药物成瘾多种神经精神疾病的病理生理过程密切相关[21]。TAAR1激动剂在动物模型中可减少中枢神经系统5-HT投射区域的异常放电, 同时改善焦虑样行为[22]。5-HT2CR分布在大脑的边缘系统, 与焦虑和抑郁相关, 研究证实, 5-HT2CR激动剂能够改善慢性应激引起的突触数量减少, 调节兴奋-抑制性平衡, 从而发挥抗焦虑作用[23]。
为验证TAAR1和5-HT2CR参与焦虑症发生的生理病理过程, 采用CUMS诱导大鼠焦虑模型。CUMS是采用温和不可预测的刺激来诱导大鼠焦虑的一种方法, 通过模拟人类日常生活中遇到的不良应激, 从而诱发动物的焦虑抑郁状态[24]。焦虑模型采用高架O迷宫和旷场来进行验证。本研究结果表明, 在CUMS诱导的大鼠焦虑模型中, 杏仁核组织中TAAR1和5-HT2CR的表达水平下降, 并且表现出焦虑样行为。过表达或敲低BLA中Taar1和Htr2c基因, 以进一步研究其对CUMS诱导的大鼠焦虑样行为的影响, 实验结果表明, 过表达BLA中Taar1或Htr2c可改善大鼠的焦虑样行为, 而敲低BLA中Taar1或Htr2c可加重大鼠的焦虑样行为。此外, 还通过TAAR1/5-HT2CR双重激动剂PCC0105004来验证TAAR1/5-HT2CR在焦虑大鼠中的作用。结果表明, 灌胃给予PCC0105004能够有效缓解大鼠的焦虑样行为, 但敲低Taar1和Htr2c的焦虑大鼠可逆转PCC0105004的上述作用。
大量证据表明, TAAR1与5-HT系统, 特别是5-羟色胺1A受体和5-羟色胺2A受体亚型之间可能存在功能上的交互作用[25]。更为重要的是, TAAR1和5-HT2CR所调控的信号通路, 最终可能汇聚为一个共同的下游通路——BDNF通路[18, 26]。该通路是调控神经元存活、突触可塑性和情绪行为的核心通路[27, 28]。本研究进一步表明, 焦虑大鼠杏仁核中的BDNF表达水平降低, 而给予药物治疗后可显著逆转上述结果。这表明, PCC0105004可能是通过影响BDNF通路来发挥抗焦虑作用。上述结果为TAAR1和5-HT2CR参与焦虑症发生的病理生理过程提供了重要依据。

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2026年第61卷第7期
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doi: 10.16438/j.0513-4870.2025-1587
  • 接收时间：2025-12-29
  • 首发时间：2026-09-21
  • 出版时间：2026-07-12
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  • 收稿日期：2025-12-29
  • 修回日期：2026-03-08
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    1烟台大学药学院, 分子药理和药物评价教育部重点实验室(烟台大学), 新型制剂与生物技术药物研究山东省高校协同创新中心, 山东 烟台 264005
    2先进药物递释系统全国重点实验室, 山东 烟台 264003

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2种不同金属材料的力学参数
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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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