细胞穿膜肽


穿透细胞膜进入细胞内是许多作用靶点在细胞内的生物大分子发挥作用的先决条件，然而生物膜的生物屏障作用阻止了许多高分子物质进入细胞内，从而很大程度地限制了这些物质在治疗领域的应用。因此，如何引导这些物质穿透细胞膜是一个迫切需要解决的问题，目前介导生物大分子穿透细胞膜的方法主要包括细胞穿透肽（cell penetrating peptides，CPPs）、脂质体、腺病毒、纳米颗粒、影细胞等，而CPPs是一类以非受体依赖方式，非经典内吞方式直接穿过细胞膜进入细胞的多肽，它们的长度一般不超过30个氨基酸且富含碱性氨基酸，氨基酸序列通常带正电荷。


1型人免疫缺陷病毒转录激活因子TAT（human immunodeficiency virus-1 transcription activator， HIV-1 TAT）是第一个被发现的细胞穿透肽，它凭借一种无毒的、高效的方式进入细胞。细胞穿透肽（cell penetrating peptides，CPPs）的一个重要特点是可以携带多种不同大小和性质的生物活性物质进入细胞，包括小分子化合物、染料、多肽、多肽核酸（peptide nucleo acid, PNA）、蛋白质、质粒DNA、siRNA、200nm的脂质体、噬菌体颗粒和超顺磁性粒子等，这一性质为其成为靶向药物的良好载体提供了可能。CPPs作为载体的优势在于低毒性和无细胞类型的限制，尽管CPPs可输送不同类型的物质进入细胞，但其实际应用多集中于寡肽、蛋白质、寡聚核苷（oligonucleotides，ONs)或类似物的细胞转运。


跨膜机理

不同的细胞穿透肽（CPP）跨膜机制不同，一个细胞穿透肽（CPP）的具体机制有赖于几个参数，如分子大小（携带物质）、温度、细胞类型和细胞内外的稳定性等。细胞穿透肽（CPP）进入细胞的具体机制目前还不清楚，比较流行的推测包括以下三种：
A: 倒置胶粒模型（inverted micelle model），CPPs通过细胞膜上磷脂分子的移动形成倒置胶粒结构，而进入胞浆。
B: 直接穿透，即孔隙结构模型 (pore formation model)，CPPs在细胞膜上组成跨膜的孔隙结构而进入胞浆 。
C: 内吞方式进行细胞摄取。

细胞穿透肽 HIV TAT

细胞穿透肽（如HIV TAT）可以以直接穿透和内吞两种方式进入细胞。HIV TAT或者简单的多聚精氨酸可被设计作为有效的药物载体，但CPP（如HIV TAT）是如何实现胞膜转运，目前仍不清楚。简单的HIV TAT是如何促进象直接穿透和内吞作用的入胞机制的呢？来自Gerard Wong实验室的研究人员研究了在不同的条件下，HIV TAT是如何与细胞质膜、细胞骨架、特异的胞膜受体相互作用，从而诱导了多重转运途径。有趣的是，TAT在不同条件下可与同一序列发生多种不同的反应，因而与胞膜、细胞骨架、特异受体相互作用可产生多种转运途径。CPP的跨膜机制与多肽序列存在很敏感的关系，如果在一个纯亲水性的CPP中增加一个疏水残基，就能彻底地改变其转运机制，例如，最简单的CPP原型-多聚精氨基（polyR），可以诱导细胞膜上形成跨膜的孔隙结构。疏水氨基酸通过插入胞膜来形成正曲率，精氨酸可同时形成正曲率和负曲率，赖氨酸只能沿一个方向形成负曲率，这就意味着在精氨酸与赖氨酸/疏水物之间存在补偿关系。

如果疏水性有助于形成负高斯曲率（Gaussian curvature），那为什么TAT肽中的疏水含量相对较低呢？其原因是CPPs都是利用尽可能少的疏水基去形成saddle-splay curvature。序列上的差异很可能只会在膜上诱导短暂的类似孔隙的跨膜结构，从而形成对CPP来说更短的孔隙寿命。由于CPP的氨基酸组成不同，TAT肽在有或无受体情况下都可以介导细胞内吞作用。


下面是一些已知的穿膜肽序列：

名称 序列 名称 序列
2 DSLKSYWYLQKFSWR (Kondo et al. 2012) MPG GALFLGFLGAAGSTMGA cysteamide (Morris et al. 1997)
18A DWLKAFYDKVAEKLKEAF (Datta et al. 2000) MPG GALFLGFLGAAGSTMGASQPKKKRKV cycteamide (Deshayes et al. 2005)
α1H KSKTEYYNAWAVWERNAP (Gomarasca et al. 2017) MPG-8 AFLGWLGAWGTMGWSPKKKRK (Crombez et al. 2009b)
α2H GNGEQREMAVSRLRDCLDRQA (Gomarasca et al. 2017) mRVG YTIWMPENPRPGTPCDIFTKSRGKRASNGGGRRRRRRRRR(Villa-Cedillo et al. 2017)
A22p HTPGNSNKWKHLQENKKGRPRR (Shin et al. 2014) MT23 LPKQKRRQRRRM (Zhou et al. 2017)
Ac-18A-NH2 DWLKAFYDKVAEKLKEAF) (Wimley and White 2000) mtCPP1 r-Dmt-OF (Cerrato et al. 2015)
aCPP Typical sequence R9GPLGLAGE8 (Li et al 2015) MTM AAVALLPAVLLALLAP (Fletcher et al. 2010)
AdVpVI(33-55) Ac-GAFSWGSLWSGIKNFGSTVKNYG (Murayama et al. 2016) MTD84 AVALVAVVAVA (Lim et al. 2014)
AIP6 RLRWR (Wang et al. 2011) MTP MLSLRQSIRFFK (Chuah et al. 2015a, b)
all-d DsC18 Glrkrlrkfrnkikek (Bergmann et al. 2017) MTS KGEGAAVLLPVLLAAPG (Zhao et al. 2001)
αgliadin(31-43) LGQQQPFPPQQPY (Paolella et al. 2018) MTS1 AAVLLPVLLAAP (Rojas et al. 1998)
Alyteserin-2a ILGKLLSTAAGLLSNL (Conlon et al. 2013) Mut3DPT-C9h VKKKKIKAEIKIYVETLDDIFEQWAHSEDL (de la Torre et al. 2017)
ANG TFFYGGSRGKRNNFKTEEY (Demeule et al. 2008) Myr-ApoE Myr-LRKLRKRLLR (Tajik-Ahmadabad et al. 2017)
ApoE(141–150) Ac-LRKLRKRLLRX-Bpg-G (Shabanpoor et al. 2017) New modalities Polycyclic, hairpin, stapled peptides for delivery (Valeur et al. 2017, Waldmann et al. 2017)
ApoE-derived Ac-LRKLRKRLLR (Tailhades et al. 2017) NF1 Stearyl-AGY(PO3)LLGKTNLKALAALAKKIL (Arukuusk et al. 2013)
Arf(1-22) MVRRFLVTLRIRRACGPPRVRV (Johansson et al. 2008) NF51 δ-(Stearyl-AGYLLG)OINLKALAALAKKIL (Arukuusk et al. 2013)
AT1002 FCIGRL (Gopalakrishnan et al. 2009) NF55 δ-(Stearyl-AGYLLG)OINLKALAALAKAIL (Freimann et al. 2016)
AT1AR(304-318) FLGKKFKKYFLQLLK (Östlund et al. 2005) NLS PKKKRKV (Yoneda et al. 1992).
Bac7 RRIRPRPPRLPRPRPRPLPFPRPGPRPIPRPL (Sadler et al. 2002) NLS-StAx-h stapled RRWPRXILDXHVRRVWR (Dietrich et al. 2017)
BGPC7-FHV RRRRNRTRRNRRRVR-RRFYGPV (Wongso et al. 2017) NoLS KKRTLRKNDRKKRC (Yao et al. 2015)
Bim EIWIAQELRRIGDEFNAYYARLLC (Kim et al. 2017) Novicidin KNLRRIIRKGIHIIKKYF (Milosavljevic et al. 2016)
BP16 KKLFKKILKKL (Soler et al. 2014) NPFSD VLTNENPFSDP (Gong et al. 2016)
BP100 KKLFKKILKYL (Eggenberger et al. 2009) NYAD-1 stapled ITFEDLLDYYGP (Zhang et al. 2008)
BPP13a GGWPRPGPEIPP (Sciani et al. 2017) Oct4-PTD DVVRVWFCNRRQKGKR (Adachi et al. 2017)
bPrPp(1-30) MVKSKIGSWILVLFVAMWSDVGLCKKRPKP (Magzoub et al. 2006) P007 Ac-(RAhxR)4-Ahx-βAla (Greer et al. 2014)
BR2 RAGLQFPVGRLLRRLLR (Lim et al. 2013) P1 LRRWSLG (Peng et al. 2017b)
Buforin II TRSSRAGLQFPVGRVIIRLLRK (Park et al.1998) P2 WKRTLRRL (Peng et al. 2017b)
Buforin IIb RAGLQFPVG[RLLR]3 (Lee et al. 2008) P3 YGRKKRRQR (Tan et al. 2006)
C6M1 RLWRLLWRLWRRLWRLLR (Jafari et al. 2014) P7 RRMKWKK (Watson et al. 2017)
C105Y CSIPPEVKFNKPFVYLI (Rhee and Davis 2006) P11 YGRKKRRQRRR (Zhao et al. 2011)
CADY GLWRALWRLLRSLWRLLWRA cycteamide (Crombez et al. 2009a) P11 HSDVHK (Bang et al. 2011)
CAR CARSKNKDC (Toba et al. 2014) P11LRR P11LRR (Li et al. 2010)
CA-Tat KWKLFKKYGRKKRRQRRR (Lv et al. 2017) P14LRR (PLPRPR)4 (Brezden et al. 2016)
CB5005 M KLKLALALALA (Zhang et al. 2016) p18 LSTAADMQGVVTDGMASG (Taylor et al. 2009)
CDB3 REDEDEIEW (Issaeva et al. 2003) P21 KRKKKGKGLGKKRDPCLRKYK (Dixon et al. 2016)
CendR RPARPAR (Hu et al. 2014) P28 LSTAADMQGVVTDGMASGLDKDYLKPDD, Leu50-Asp77 of azurin (Yamada et al. 2016)
cFΦR4 cyclic FΦRRRRQ (Qian et al. 2014) p28 FLHSGTAVTCTYPALTPQWEGSDCTHRL (Signorelli et al. 2017)
CGKRK CGKRK (Griffin et al. 2017) p53 peptide MO6 Stapled TSF*EYWYLL* (Chee et al. 2014)
CIGB-300 cyclic CWMSPRHLGTC-Tat (Perera et al. 2012) PAF26 Ac-rkkwfw (Lopez-Garcia et al. 2002)
CIGB-552 Ac-HARIKpTFRRlKWKYKGKFW (Fernandez Masso et al. 2013) PAS GKPILFF (Woldetsadik et al. 2017)
CLIP6 KVRVRVRVpPTRVRERVK (Soudah et al. 2017) pCLIP6 KVRVRVRVpP(pT)RVRERVK (Chen et al. 2017b)
CooP ACGLSGLGVA (Hyvonen et al. 2014) pD-SP5 riPRPSPKMGV(pS)VS (Chen et al. 2017b)
CpMTP ARLLWLLRGLTLGTAPRRA (Jain and Chugh 2016) PenetraMax KWFKIQMQIRRWKNKR, L- and D- (Khafagy el et al. 2015)
CPNT STSGTGKMTRAQRRAAARRNRA (Qi et al. 2011) Penetratin RQIKIWFQNRRMKWKK (Derossi et al. 1994)
CPP1 (KFF)3K (Patel et al. 2017) Pep-1 KETWWETWWTEWSQPKKKRKV cysteamide (Morris et al. 1997)
CPP33 RLWMRWYSPRTRAYG (Lin et al. 2018) pepM KLFMALVAFLRFLTIPPTAGILKRWGTI (Freire et al. 2014)
CPP-C PIEVCMYREP (Nakayama et al. 2011) pepR LKRWGTIKKSKAINVLRGFRKEIGRMLNILNRRRR (Freire et al. 2014)
CPPecp NYRWRCKNQN (Fu et al. 2017) Pept1 PLILLRLLRGQF (Marks et al. 2011)
C-peptide GPGLWERQAREHSERKKRRRESECKAA (Fan et al. 2016) Peptide 599 GLFEAIEGFIENGWEGMIDGWYGGGGRRRRRRRRRK(Alexander-Bryant et al. 2015)
CRGDK CRGDK (Zhao et al. 2018) Pep42 Cyclic CTVALPGGYVRVC (Kim et al. 2006)
Crotamine YKQCHKKGGHCFPKEKICLPPSSDFGKMDCRWRWKCCKKGSG(Rodrigues et al. 2012) PepNeg SGTQEEY (Neves-Coelho et al. 2017)
cSN50 AAVALLPAVLLALLAPVQRKRQKLMP (Torgerson et al. 1998) PepFect6 Stearyl-AGYLLGK(εTMQ)INLKALAALAKKIL, PF6 (El-Andaloussi et al. 2011)
65-2CTS CPYVNQRPQKARYRNG (Percipalle et al. 2003) PepFect14 Stearyl- AGYLLGKLLOOLAAAALOOLL (Ezzat et al. 2011)
CWR8K CWR8K (Sasaki et al. 2008) PG1 RGGRLCYCRRRFCVCVGR (Liu et al. 2013b)
CyLoP-1 CRWRWKCCKK (Ponnappan et al. 2017) pHLIP AEQNPIY-WARYADWLFTTPLLLLDLALLV-DADEGT (Andreev et al. 2010)
Cyt c(77–101) GTKMIFVGIKKKEERADLIKKA (Howl and Jones 2015) PHPs H6-H10 peptides (Kimura et al. 2017)
DAG cyclic CDAGRKQKC (Mann et al. 2017) PIP1 RXRRXRRXRIKILFQNRRMKWKK (Ivanova et al. 2008)
D-JNKI-1 RPKRPTTLNLFPQVPRSQDT (Bonny et al. 2001) Pip5e RXRRBRRXRILFQYRXRBRXRB (Betts et al. 2012)
DK17 DRQIKIWFQNRRMKWKK (Bera et al. 2016) Pip6a Ac-RXRRBRRXRYQFLIRXRBRXRB (Lehto et al. 2014)
DLP ACKTGSHNQCG (Kumar et al. 2015) POD CGGG(ARKKAAKA)4 (Dasari et al. 2017)
DMBT1-derived GRVEVLYRGSW and GRVRVLYRGSW (Tuttolomondo et al. 2017) PR9 FFLIPKG-R9 (Liu et al. 2013a)
dNP2 KIKKVKKKGRK-KIKKVKKKGRK (Lim et al. 2015) PTD YARVRRRGPRRR (Dong et al. 2016)
DPV3 RKKRRRESRKKRRRES (Tacken et al. 2008) PTD3 R9-ETWWETWWTEW (Kizaka-Kondoh et al. 2009)
DPV1047 CVKRGLKLRHVRPRVTRMDV (De Coupade et al. 2005) PTD4 YARAAARQARA (McCusker et al. 2007)
DRTTLTN DRTTLTN (Gennari et al. 2016) Poly-Arg Most popular R7 - R12 (Mitchell et al. 2000, Futaki, 2006)
DS4.3 RIMRILRILKLAR (Jeong et al. 2014) pVEC LLIILRRRIRKQAHAHSK (Elmquist et al. 2001)
Dynorphin A YGGFLRRIRPKLKWDNQ (Marinova et al. 2005) Pyrrhocoricin VDKGSYLPRPTPPRPIYNRN (Otvos et al. 2000)
EA GLKKLAELAHKLLKLGC (Yang et al. 2014) R4K1 Stapled Ac-RRRRKS*LHRS*LQDS (Speltz et al. 2018)
EB1 LIRLWSHLIHIWFQNRRLKWKKK (Lundberg et al. 2007) R6dGR R6dGR (Wang et al. 2017)
EF GLKKLAELFHKLLKLGC (Yang et al. 2014) R8 R8 (Wender et al. 2001)
EHB RCSHYTGIRCSHMAATTAGIYTGIRCQHVVL-C6H (Cao et al. 2018) R8-dGR R8dGR (Liu et al. 2016b)
EPRNEEK EPRNEEK (Orihuela et al. 2009) R9-H4A2 Ac-YR9-HAHAHH (Okitsu et al. 2017)
F3** diphosphorylated dipeptide (Miao et al. 2016) R6W3 R6W3 (Bechara et al. 2013)
G4R9L4 G4R9L4 (Ramakrishna et al. 2014) R10W6 R10W6 (Bechara et al. 2013)
GALA WEAALAEALAEALAEHLAEALAEALEALAA (Li et al. 2004) RA9 RRAARRARR (Alhakamy et al. 2013)
GeT KIAKLKAKIQKLKQKIAKLK (Rakowska et al. 2014) RALA WEARLARALARALARHLARALARALRACEA (McCarthy et al. 2014)
gH625 HGLASTLTRWAHYNALIRAF (Galdiero et al. 2015) RDP CKSVRTWNEI IPSKGCLRVG GRCHPHVNGG GRRRRRRRRC(Xiao et al. 2017)
Gi3α(346-355) KNNLKECGLY (Jones et al. 2005) REDV REDV (Yang et al. 2016)
Glu-Lys EEEAAKKK (Lewis et al. 2010) RF GLKKLARLFHKLLKLGC (Yang et al. 2014)
GV1001 EARPALLTSRLRFIPK (Kim et al. 2016a) cRGDfC Cyclic RGDfC (Wada et al. 2017)
GWH1 GYNYAKKLANLAKKFANALW (Serna et al. 2017) iRGD Cyclic CRGDKGPDC (Peng and Kopecek, 2015)
H2A derived SGRGKQGGKARAKAKTRSSRAGLQFPVGRVHRLLRKG(Rosenbluh et al. 2004) RGE RGERPPR (Yu et al. 2017)
H6R6 H6R6 (Sun et al. 2017) RH9 RRHHRRHRR (Alhakamy et al. 2013)
H16 H16 (Iwasaki et al. 2015) RL9 RRLLRRLRR (Alhakamy et al. 2013)
HA2(1-23) GLFGAIAGFIENGWEGMIDGWYG (Esbjörner et al. 2007) RL16 RRLRRLLRRLLRRLRR (Joanne et al. 2009)
HAIYPRH HAIYPRH (Shteinfer-Kuzmine et al. 2017) RT53 RQIKIWFQNRRMKWKKAKLNAEKLKDFKIRLQYFARGLQVYIRQLRLALQGKT (Jagot-Lacoussiere et al. 2016)
hBD3-3 GKCSTRGRKCCRRKK (Lee et al. 2015b) RTP004 RKKRRQRRRG-K15-GRKKRRQRRR) (Lee et al. 2015a)
HBP GKRKKKGKGLGKKRDPCLRKYK (Luo et al. 2016) RV24 RRRRRRRRRGPGVTWTPQAWFQWV (Lo and Wang, 2012)
hLF KCFQWQRNMRKVRGPPVSCIKR (Duchardt et al. 2009) RVG YTIWMPENPRPGTPCDIFTNSRGKRASNG (Kumar et al. 2007)
Hph-1 YARVRRRGPRR (Jung et al. 2011) RVG-9R YTIWMPENPRPGTPCDIFTNSRGKRASNGGGGRRRRRRRRR(Rassu et al. 2017)
HR9 CH5-R9-H5C (Liu et al. 2013a) RVG29 YTIWMPENPRPGTPCDIFTNSRGKRASNGGGGRRRRRRRRR(Villa-Cedillo et al. 2017)
Hst5 DSHAKRHHGYKRKFHEKHHSHRGY (Luque-Ortega et al. 2008) RW9 RRWWRRWRR (Alhakamy et al. 2013)
I1WL5W WKKIWSKIKKLLK (Bi et al. 2014) RW16 RRWRRWWRRWWRRWRR (Jobin et al. 2013)
I4WL5W IKKWWSKIKKLLK (Bi et al. 2014) (RXR)4 (R-Ahx-R)4 (Saleh et al. 2010)
ID No.2 MAAWMRSLFSPLKKLWIRMH (Eudes and Macmillan 2014) (rXr)4 (r-Ahx-r)4 (Vij et al. 2016b)
IMT-P8 RRWRRWNRFNRRRCR (Gautam et al. 2016) S155 VKKKKIKREI-KIAAQRYGRELRRMADEFHV (Haidar et al. 2017)
INF GLFEAIEGFIENGWEGMIDGWYGC (Pichon et al. 1997) S4(13)-PV ALWKTLLKKVLKAPKKKRKV (Mano et al. 2007)
iNGR CRNGRGPDC (Alberici et al. 2013) SAP VRLPPPVRLPPPVRLPPP (Pujals et al. 2006)
isl-1 RVIRVWFQNKRCKDKK (Kilk et al. 2001) SAP(E) VELPPPVELPPPVELPPP (Martin et al. 2011)
JB9 cskc (Basu and Wickstrom, 1997) all-D-SAP (vrlppp)3 (Pujals et al. 2007)
JB434 R9GGLAA-Aib-SGWKH6 (Sangtani et al. 2018) SAPSp-lipo stearyl-GGGGHGAHEHAGHEHAAGEHHAHE (Suzuki et al. 2017)
KAFAK KAFAKLAARLYRKALARQLGVAA (Bartlett et al. 2013) SAR6EW SAR6EW (Im et al. 2017)
KALA WEAKLAKALAKALAKHLAKALAKALKACEA (Wyman et al. 1997) sC18 GLRKRLRKFRNKIKEK (Oren et al. 1999)
Kalata B1 polycyclic CGETCVGGTCNTPGCTCSWPVCTRNGLPV (Daly et al. 1999) (sC18)2 (GLRKRLRKFRNKIKEK)2 (Gronewold et al. 2017)
(KFF)3K (KFF)3K (Rownicki et al. 2017) SMTP motif, LRLLR (Fuselier and Wimley, 2017)
K-FGF AAVLLPVLLAAP (Lin et al. 1995) SPACE Cyclic ACTGSTQHQCG (Hsu and Mitragotri, 2011)
KH (KH)9 (Chuah et al. 2016) SRCRP2-11 GRVEVLYRGSW (Tuttolomondo et al. 2017)
KLA KLAKLAKKLAKLAK (Huang et al. 2017) STR-KV H3K3V6 (Pan et al. 2016)
KLAK KLALKLALKALKAALKLA (Oehlke et al. 1998) SS-02 Dmt-r-FK (Alta et al. 2017)
KLA-R7 KLAKLAKKLAKLAKGGRRRRRRR (Lemeshko, 2013) SS-20 F-r-FK (Alta et al. 2017)
KP MAPTKRKGSCPGAAPNKKP (Villa-Cedillo et al. 2017) SS-31 r-Dmt-KF (Zhao et al. 2005)
KST peptide STGKANKITITNDKGRLSK (Adachi et al. 2017) SynB1 RGGRLSYSRRRFSTSTGR (Rousselle et al. 2000)
L1−6 PLILLRLLR (Schmidt et al. 2017) T2 LVGVFH (Kumar et al. 2012)
L5a RRWQW (Liu et al. 2016a) Tat(49-57) RKKRRQRRR (Vives et al. 1997a)
L17E IWLTALKFLGKHAAKHEAKQQLSKL (Akishiba et al. 2017) Tat(48-60) GRKKRRQRRRPPQ (Vives et al. 1997b)
lactoferrampin(265- 284) DLIWKLLSKAQEKFGKNKSR (Reyes-Cortes et al. 2017) Tat(44-57) CGISYGRKKRRQRRR (Niesner et al. 2002)
lactoferricin(17-30) FKCRRWQWRMKKLG (Reyes-Cortes et al. 2017) Tat(37-72) CFITKALGISYGRKKRRQRRRPPQGSQT-HQVSLSKQ (Fawell et al. 1994)
lactoferrin(19-40) KCFMWQEMLNKAGVPKLRCARK (Duchardt et al. 2009) Tat analog GRKKRRQR (Nguyen et al. 2008)
LAH1 KKLALALALALHALALALALKKA (Moulay et al. 2017) Tat-LK15 Tat-KLLKLLLKLLLKLLK (Peng et al. 2017a)
LALF(31-52) HYRIKPTFRRLKWKYKGKFW (Yanez et al. 2017) TCTP MIIFRALISHKK (Bae et al. 2016)
LB FKCRRWQWRMKKLGAPSITCVRRAF) (Liu et al. 2013b) TD-1 ACSSSPSKHCG (Chen et al. 2006)
L-CPP LAGRRRRRRRRRK (Liu et al. 2006) TD2.2 SYWYRIVLSRTGRNGRLRVGRERPVLGESP (Heffernan et al. 2012)
LDP-NLS KWRRKLKKLRPKKKRKV (Ponnappan and Chugh, 2017) TH peptide GYLLGHINLHHLAHL-Aib-HHIL (Chen et al. 2017a)
LE10 LELELELELELELELELELE (Antunes et al. 2013) TM2 PKKGSKKAVTKAQKKDGA (Kochurani et al. 2015)
LF chimera FKCRRWQWRMKKLG-K-RSKNKGFKEQAKSLLKWILD(Reyes-Cortes et al. 2017) Transportan GWTLNSAGYLLGKINLKALAALAKKIL, TP (Pooga et al. 1998)
linTT1 AKRGARSTA (Hunt et al. 2017) TP10 AGYLLGKINLKALAALAKKIL (Soomets et al. 2000)
LK LKKLLKLLKKLLKLAG (Kim et al. 2016b) TPk VRRFkWWWkFLRR (Bahnsen et al. 2015)
LL37 LLGDFFRKSKEKIGKEFKRIVQRIKDFLRNLVPRTES (Kim et al. 2016b) Tpl KWCFRVCYRGICYRRCRGK (Jain et al. 2015)
LLIIL LLIIL (Alaybeyoglu et al. 2017) TPP TKDNNLLGRFELSG (Gehrmann et al. 2014)
LMWP VSRRRRRRGGRRRR (Chen et al. 2017d) TT1 CKRGARSTA (Paasonen et al. 2016)
LP-12 HIITDPNMAEYL (Kumar et al. 2015) vAMP 059 INWKKWWQVFYTVV (Dias et al. 2017)
LPAs RCnRCnK (Gupta et al. 2011) vCPP 0769 RRLTLRQLLGLGSRRRRRSR (Dias et al. 2017)
LTV LTVSPWY (Chopra 2012) vCPP 2319 WRRRYRRWRRRRRWRRRPRR (Dias et al. 2017)
lycosin-I RKGWFKAMKSIAKFIAKEKLKEHL (Tan et al. 2017) VDAC(1-26) MAVPPTYADLGKSARDVFTKGYGFGL (Smilansky et al. 2015)
Lyp1 CGNKRTRGC (Fogal et al. 2008) VP22 NAATATRGRSAASRPTQRPRAPARSASRPRRPVQ (Elliott and O'Hare, 1997)
M918 MVTVLFRRLRIRRACGPPRVRV (El-Andaloussi et al. 2007) V peptide TVDNPASTTNKDKLFAVRK (Manosroi et al. 2014)
Maurocalcine GDCLPHLKLCKENKGCCSKKCKRRGTNIEKRCR (Poillot et al. 2010) VT5 DPKGDPKGVTVTVTVTVTGKGDPKPD (Oehlke et al. 1997)
MAP KLALKLALKALKAALKLA (Oehlke et al. 1998) W(RW)4 W(RW)4 (Nasrolahi Shirazi et al. 2013)
MAP12 LKTLTETLKELTKTLTEL (Oehlke et al. 2002) Xentry LCLR (Montrose et al. 2014)
MCoTI-I polycyclic SGSDGGVCPKILQRCRRDSDCPGACICRGNGYCG(Camarero, 2017) X-pep MAARLC (Adachi et al. 2017)
MCoTI-II polycyclic CPKILKKCRRDSDCPGACICRGNGYCGSGSDGGV(Huang et al. 2015) YKA YKALRISRKLAK (Desai et al. 2014)
MFK MFKLRAKIKVRLRAKIKL (Samuels et al. 2017) YTA2 YTAIAWVKAFIRKLRK (Lindgren et al. 2006)
Mgpe9 CRRLRHLRHHYRRRWHRFRC (Vij et al. 2016a) YTA4 IAWVKAFIRKLRKGPLG (Lindgren et al. 2006)
MitP INLKKLAKL(Aib)KKIL (Howl et al. 2018) Z2 FWIGGFIKKLKRSKLA (Chen et al. 2017c)
m(KLA)-iRGD klaklakklakla-K-GG-iRGD (Qifan et al. 2016) Z3 FKIKKFIGGLWRSKLA (Chen et al. 2017c)
MMGP1 MLWSASMRIFASAFSTRGLGTRMLMYCSLPSRCWRK(Pushpanathan et al. 2013) Z12 KRYKNRVASRKCRAKFKQLLQHYREVAAAKSSENDRLRLLLK(Derouazi et al. 2015)
MPER fragment ELDKWASLWNWFDITNWLWYIK (Song et al. 2009) ZXR-1 FKIGGFIKKLWRSKLA (Chen et al. 2017c)


备注:

Ac=acetyl; 

Ahx or X=aminohexanoic acid; 

Aib=aminoisobutyric acid; 

B=β-alanine; 

Bpg=bishomopropargylglycine; 

Dmt=dimethyltyrosine; 

O=ornithine; pT=phosphoryl-Thr; 

pS=phosphoryl-Ser; 

Φ=l-2-naphthylalanine; 

ri=retro-inverso; 

*=staple tethering site


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