2025年12月9日 (二) 07:04 imported>Olaf8940

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{{Expand language|en|ja|status=yes}}
{{NoteTA
|G1=Lifesciences
|G2=Chemistry
|1=zh-cn:规律间隔成簇短回文重复序列;zh-tw:常間回文重複序列叢集
|2=zh-cn:规律间隔成簇短回文重复序列及其相关蛋白基因;zh-tw:常間回文重複序列叢集關聯蛋白
|3=zh-cn:相关蛋白;zh-tw:關聯蛋白
}}
'''CRISPR'''<small>（[[國際音標|IPA]]：/ˈkrɪspər/；[[DJ音標|DJ]]：/ˈkrispə/；[[KK音標|KK]]：/ˈkrɪspɚ/）</small>是由[[日本]]科學家於1987年在[[大腸桿菌]]的[[基因組]]中發現到特別規律的[[DNA序列]]。即某一小段DNA會一直重複，重複片段之間又有相等長的間隔，此序列就是「'''C'''lustered '''R'''egularly '''I'''nterspaced '''S'''hort '''P'''alindromic '''R'''epeats，簡稱： '''CRISPR'''」亦可稱「'''CRISPR-associated proteins'''，簡稱： '''Cap'''」{{r|sciencedirect-01}}；中文各別譯為「'''常間回文重複序列叢集'''」{{r|scimonth-01}}和「'''常間回文重複序列叢集關聯蛋白'''」{{r|narlabs-01}}。

CRISPR是存在於[[细菌]]、[[古菌]]中的一種[[基因]]，該類基因組中含有曾經攻擊過該細菌的[[病毒]]之基因片段。細菌透過這些基因片段來偵測並抵抗相同病毒的攻擊，並摧毁其DNA。這類基因組是細菌[[免疫系统]]的關鍵組成部分。透過這些基因组，人類可以準確且有效地編輯[[生物體]]内的部分基因，也就是[[CRISPR/Cas9基因編輯技術]]。

== 機制 ==
[[Image:Crispr.png|thumb|350px|right|CRISPR可能的机制示意图{{r|science-03}}]]
'''CRISPR/Cas系統'''，為目前發現存在於多數[[細菌]]與絕大多數的[[古菌]]中的一種[[後天免疫]]系統{{r|annualreviews-01}}，以消滅外來的[[质粒]]或者[[噬菌體]]{{r|science-01|science-02}}，並在自身[[基因组|基因組]]中留下外來基因片段作為“記憶”{{r|nature-01}}。

目前已發現三種不同類型的 CRISPR/Cas系統，存在於大約40%和90%已定序的[[细菌]]和[[古菌]]中{{r|u-psud-01|bmcbioinformatics-01}}。其中第二型的組成較為簡單，以[[Cas9|Cas9蛋白]]以及[[嚮導RNA]]（gRNA）為核心的組成{{r|airitilibrary-01}}。

Cas9是第一個被廣泛應用的CRISPR[[核酸酶]]，其次是Cpf1，其在{{le|新澤西弗朗西斯菌|Francisella novicida}}的[[CRISPR/Cpf1]]系统中被發現{{r|cell-01|nature-02}}，而在其它系統中也被認為是存在的{{r|economist-01}}。

由於其對[[去氧核醣核酸|DNA]]干擾（DNAi）的特性，目前被積極地應用於[[遺傳工程]]中，作為基因體剪輯工具，與[[鋅指核酸酶]]及[[類轉錄活化因子核酸酶]]同樣利用[[非同源性末端接合]]的機制，於基因體中產生[[脫氧核醣核酸|DNA]]的[[双链断裂]]以利剪輯。第二型 CRISPR/Cas 經由遺傳工程的改造應用於[[哺乳類]]細胞及[[斑馬魚]]的基因體剪輯{{r|science-04|nature-03}}。其設計簡單以及操作容易的特性為最大的優點，目前已逐步應用在各種不同的[[模式生物]]當中{{r|airitilibrary-01}}。

== 發現歷史 ==
{{Infobox nonhuman protein
|Name=Cascade<br> (CRISPR相關病毒防禦複合體)
|image=CAS 4qyz.png
|width=300px
|caption =crRNA引導的大腸桿菌串級複合體（藍）與單股DNA（橘）結合的結構
|Organism=大腸桿菌（Escherichia coli）
| TaxID = 511145
| Symbol = CRISPR
| EntrezGene = 947229
|AltSymbols=
|ATC_prefix=
|ATC_suffix=
|ATC_supplemental=
|CAS_number=
|CAS_supplemental=
|DrugBank=
|HomoloGene=
|PDB =4QYZ
| UniProt = P38036
| RefSeqmRNA =
| RefSeqProtein = NP_417241.1
}}
聚簇DNA重複的發現始於世界三個地區的三個獨立地點。現今稱為CRISPR的基因組重複群集，即原核生物擬核DNA鏈中的丛生重复序列，在1987年一篇由[[大阪大学]][[石野良純]]教授領銜的[[大腸桿菌]]研究报告中被首次描述{{r|jb-01}}。2000年，相似的重复序列在其它[[细菌]]和[[古菌]]中被发现，并被命名为「短间隔重复序列（Short Regularly Spaced Repeats，SRSR）」{{r|wiley-01}}。2002年SRSR被重命名为『CRISPR』，其中一部分基因编码的蛋白为[[核酸酶]]和[[解旋酶]]，这些「关联蛋白（associated proteins）」与CRISPR一同组成『CRISPR/Cas系统』{{r|wiley-02}}。

=== Cas9 ===
科学家还研究了来自[[化脓性链球菌]]的更简单的CRISPR系统，其依赖于蛋白[[Cas9]]。Cas9内切核酸酶是包含两个小RNA分子的四组分系统{{r|genomebiology-01}}。[[詹妮弗·杜德纳]]、[[埃马纽埃尔·卡彭蒂耶]]及[[張鋒]]各自独立的探索CRISPR关联蛋白，了解细菌如何在它们的免疫防御使用间隔（spacer）。他们共同研究一个比较简单的依赖于称为[[Cas9|Cas9蛋白]]的CRISPR系统{{r|worldcat-01}}。

=== Cpf1 ===
在2015年，核酸酶Cpf1被发现在{{le|新泽西弗朗西斯菌|Francisella novicida}}的[[CRISPR/Cpf1]]系统{{r|cell-01|nature-02}}。其他这样的系统被认为存在{{r|economist-01}}。Cpf1与Cas9的有几个关键差异，包括：
1. DNA 斷裂方式不同：导致双链DNA中的“交错”或“黏性(sticky end)”切割，而不是由Cas9产生的“钝的 (Blunt end)”切割。
2. {{le|相邻間隔原基序|Protospacer adjacent motif}}(PAM)不同：Cpf1辨認“富含T碱基”的PAM，而 Cas9 辨認 NGG 為PAM，可为 Cas9 提供替代的標靶序列。
3. 仅需要 CRISPR RNA（crRNA）用于成功標定（使用Cas9同时需要crRNA和一个{{le|反向活化crRNA|Trans-activating crRNA}}（tracrRNA））。{{r|geneonline-01}}

=== Cas9n ===
Cas9n是SpCas9的D10A突变体，只保留了SpCas9两个核酸酶结构域（RuvC和HNH）中的一个来产生DNA切口而不是DSB。因此，需要两个靶向相反的Cas9n才能在靶DNA内产生DSB，这种方法大大提高了靶标特异性，因为不太可能产生两个离靶切口。{{r|nih-01}}

== 應用 ==
用於開發[[基因剔除]]和[[標靶治療|標靶]]抗癌藥物治療方法，例如[[囊性纖維化]]和[[鐮刀型紅血球疾病|鐮狀紅血球貧血症]]{{r|scimonth-02}}。

== 評價與獲獎 ==
在2012年和2013年，CRISPR是《[[科学 (期刊)|科学]]》年度突破的第二名{{r|aaas-01}}。在2014年和2016年被《[[麻省理工科技評論]]》评为10项突破技术之一{{r|technologyreview-01|technologyreview-02}}。[[埃玛纽埃勒·沙尔庞捷|埃馬紐埃爾·夏彭蒂耶]]與[[珍妮弗·道德納]]因藉CRISPR-Cas9基因編輯技術的研究成果榮獲2020年[[諾貝爾化學獎]]{{r|cna-01}}。

== 参阅 ==
* [[基因编辑婴儿事件]]
* [[RNA干扰]]
* [[小分子干扰核糖核酸|小分子干扰核糖核酸（siRNA）]]
* [[生物武器]]
* [[合成生物学]]

==參考文獻==
;引用列表
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<ref name="geneonline-01">{{Cite web|url=https://geneonline.news/crispr-cas9-cpf1-next/|title=預測下一個生技世代的風貌(二)： CRISPR 的積極進取 |accessdate=2024-12-06|author=Editor-Jane|date=2016-08-11|publisher=GeneOnline News|language=zh-hant}}</ref>
<ref name="nih-01">{{Cite journal|title=Genome editing using Cas9 nickases|url=https://pubmed.ncbi.nlm.nih.gov/25398340/|last1=Trevino|first1=Alexandro E|last2=Zhang|first2=Feng|date=2014|journal=Methods in Enzymology|issue=|doi=10.1016/B978-0-12-801185-0.00008-8|volume=546|pages=161-174|language=en|pmid=25398340|access-date=2024-12-06|archive-date=2024-12-02|archive-url=https://web.archive.org/web/20241202231841/https://pubmed.ncbi.nlm.nih.gov/25398340/|dead-url=no}}</ref>
<ref name="scimonth-02">{{Cite web|url=https://www.scimonth.com.tw/archives/176|title=物種基因剔除技術爆炸性的新突破─ CRISPR/Cas9技術淺談|accessdate=2024-12-06|author=林翰佐|date=2015-12-01|publisher=科學月刊|language=zh-tw|archive-date=2022-07-04|archive-url=https://web.archive.org/web/20220704085958/https://scimonth.com.tw/archives/176|dead-url=no}}</ref>
<ref name="aaas-01">{{cite news|last1=Cohen|first1=Adam D. |title=AAAS Fellow Receives Nobel Prize in Chemistry for Development of CRISPR|url=https://www.aaas.org/news/aaas-fellow-receives-nobel-prize-chemistry-development-crispr?_gl=1*wt3a14*_gcl_au*MzIzMjQwNTAuMTczMzQ2MTc4OQ..|accessdate=2024-12-06|publisher= American Association for the Advancement of Science|date=2020-10-07|language=en}}</ref>
<ref name="cna-01">{{Cite news|url=https://www.cna.com.tw/news/firstnews/202010075009.aspx|title=諾貝爾化學獎 2女學者開發基因編輯技術享殊榮都曾獲唐獎|accessdate=2024-12-06|author=|date=2020-10-07|publisher=中央社|language=zh-tw}}</ref>
}}

;來源列表
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{{refend}}

== 延伸閱讀 ==
{{scholia|Q412563}}
{{Commons category|CRISPR}}
* [https://fas.org/sgp/crs/misc/R44824.pdf Advanced Gene Editing: CRISPR-Cas9]{{En}}{{Wayback|url=https://fas.org/sgp/crs/misc/R44824.pdf |date=20171205210255 }}
* [https://www.ibiology.org/ibiology_podcasts/jennifer-doudna-genome-engineering-with-crispr-cas9-birth-of-a-breakthrough-technology/ Jennifer Doudna: Genome Engineering with CRISPR-Cas9: Birth of a Breakthrough Technology] {{Wayback|url=https://www.ibiology.org/ibiology_podcasts/jennifer-doudna-genome-engineering-with-crispr-cas9-birth-of-a-breakthrough-technology/ |date=20241207175906 }}{{En}}{{Wayback|url=https://www.ibiology.org/ibiomagazine/jennifer-doudna-genome-engineering-with-crispr-cas9-birth-of-a-breakthrough-technology.html |date=20161125044709 }}

[[Category:基因工程]]
[[Category:免疫學]]
[[Category:非編碼RNA]]
[[Category:重複DNA序列]]
[[Category:分子遺傳學]]

CRISPR - 版本历史

2025年12月9日 (二) 07:04 imported>Olaf8940