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. 2020 Feb 25;11:108. doi: 10.3389/fneur.2020.00108

Table 2.

Studies demonstrating neuroprotective and other neuroactive properties of peptides fused to TAT and other cell penetrating peptides.

Peptide name Peptide sequence % Arginine Net charge at pH 7 Neuronal injury model References
TAT-NR2B9c (NA-1) YGRKKRRQRRR-KLSSIESDV 30% +7 Excitotoxicity, stroke, HIE, ICH, AD, epilepsy, pain (5968)
JNKI-1D-TAT, JNKI-1-TAT dqsrpvqpflnlttprkprpp-rrrqrrkkrg-NH2,
GRKKRRQRRR-PP-RPKRPTTLNLFPQVPRSQD-NH2
29% +12 Excitotoxicity, stroke, HIE, ICH, TBI, AD, SCI, SMA, epilepsy, pain (60, 6983)
TAT-JIP-1 GRKKRRQRRR-RPKRPTTLNLF 38% +11 Excitotoxicity, stroke, GCI, PD (8486)
δSV1-1-TAT YGRKKRRQRRR-SFNSYELGSL 28% +7 Stroke (87, 88)
TAT-JBD GRKKRRQRRR-PP-RPKRPTTLNLFPQVPRSQDT 28% +11 HIE, GCI (89, 90)
TAT-NPEG4-(IETDV)2 YGRKKRRQRRR-(Peg)4-(IESDV)2 28% +9 Stroke, pain, epilepsy, cortical spreading depression (9195)
JNK3-N-TAT YGRKKRRQRR-RCSEPTLDVKI 29% +6.9 PD (96, 97)
Src40–49Tat KPASADGHRGY-GRKKRRQRRR 33% +9.1 Pain (98)
TAT-SabKIM1 GFESLSVPSPLDLSGPRVVAPP-RRRQRRKKRG-NH2 22% +8 PD (99)
TAT-CBD3 YGRKKRRQRRR-ARSRLAELRGVPRGL 38% +11 Excitotoxicity, stroke, TBI, pain (100105)
R9-CBD3 RRRRRRRRR-ARSRLAELRGVPRGL 54% +12
TAT-CBD3A6K YGRKKRRQRRR-ARSRLKELRGVPRGL 38% +12
TAT-CRMP-2 YGRKKRRQRR-GVPRGLYDGVCEV 26% +6.9 Excitotoxicity, stroke, OGD (106108)
TAT-NR2Bct YGRKKRRQRRR-KKNRNKLRRQHSY 37% +14.1 Excitotoxicity, stroke (109111)
TAT-NR2Bcts YGRKKRRQRRR-NRRRNSKLQHKKY 35% +14.1 Excitotoxicity (109, 110)
Tat-D2LIL3−29−2 YGRKKRRQRRR-MKSNGSFPVNRRRMD 34% +11 Depression (112)
Penetratin-COG133 (COG112) Ac-RQIKIWFQNRRMKWKK-LRVRLASHLRKLRKRLL-NH2 24% +14.1 TBI, EAE, AD, axonal regeneration, spinal cord demyelination (40, 41, 47, 113115)
TAT-NR2Bct-CTM YGRKKRRQRRR-KKNRNKLRRQHSY-KFERQKILDQRFFE 35% +15.1 Stroke (116)
CN2097 RRRRRRRC-s-s-CKNYKKTEV (cyclic or linear) 41% +9 Excitotoxicity, pain (14, 117)
P42-TAT AASSGVSTPGSAGHDIITEQPRS-GG-YGRKKRRQRRR 19% +7.1 Huntington's disease (118)
TAT-p53DM YGRKKRRQRRR-RVCACPGRDRRT 43% +11 288,289 (14, 109, 119, 120)
TAT-p53DMs YGRKKRRQRRR-CCPGECVRTRRR 43% +11 Excitotoxicity (109)
TAT-CN21 YGRKKRRQRR-KRPPKLGQIGRSKRVVIEDDR 29% +11 Excitotoxicity, stroke, GCI (121123)
PYC36-TAT,
PYC36D-TAT
GRKKRRQRRRGG-LQGRRRQGYQSIKP,
pkisqygqrrrgqlgg-rrrqrrkkrg
35% +12 Excitotoxicity (10)
TAT-GluR6-9c YGRKKRRQRR-RLPGKETMA 32% +8 Excitotoxicity, GCI, stroke, OGD (124126)
TAT-mGluR1 YGRKKRRQRRR-VIKPLTKSYQGSGK 24% +11 Excitotoxicity, HIE, SAH (127129)
TAT-K13 YGRKKRRQRR-KEIVSRNKRRYQED 33% +9 Stroke (130)
TAT-Indip YGRKKRRQRRR-GEPHKFKREW 33% +9.1 Excitotoxicity, ALS (109, 131)
TAT-Indip-K/R YGRKKRRQRRR-GEPHRFRREW 43% +9.1 Excitotoxicity (109)
TAT-GESV,
D-TAT-GESV
RRRQRRKKRG-YAGQWGESV,
rrrqrrkkrg-yagqwgesv
32% +7 Excitotoxicity, HIE, pain (132134)
TAT-NEP1-40 YGRKKRRQRRR-RIYKGVIQAIQKSDEGHPFRAYLESEV AISEELVQKYSNS 16% +7.1 Stroke, OGD (135, 136)
TAT-NBD YGRKKRRQRRR-TALDWSLWQTE 27% +6 HIE (137)
TAT-ψεHSP90 YGRKKRRQRRR-PKDNEER 39% +8 Stroke, OGD (138)
TAT-Bec YGRKKRRQRRR-GG-TNVFNATFEIWHDGEFGT 19% +6.1 SCI (139)
TAT-gp91ds GRKKRRQRRR-CSTRIRRQL-NH2 47% +12 SCI, TBI, SAH (140142)
TAT-ISP GRKKRRQRRR-CDMAEHMERLKANDSLKLSQEYESI-NH2 20% +6 SCI (143)
Tat-Cav3.2-III-IV YGRKKRRQRRR-EARRREEKRLRRLERRRRKAQ 50% +16 Pain (144)
TAT-μCL YGRKKRRQRRR-PPQPDALKSRTLR 33% +10 Retinal degeneration (145)
ST2-104 RRRRRRRRR-ARSRLAELRGVPRGL 54% +12 Pain (146)
TAT-STEP YGRKKRRQRRR -GLQERRGSNVSLTLDM 30% +8 Excitotoxicity, stroke, OGD (147)
TAT-K YGRKKRRQRRR-PP-LNRTPSTVTLNNNT 26% +9 Excitotoxicity (148)
TAT-P110 YGRKKRRQRRR-GG-DLLPRGT 35% +9 Stroke, Huntington's disease (149, 150)
TAT-C6 GRKKRRQRRR-CRRGGSLKAAPGAGTRR 37% +14 Stroke (151)
Analog 4 and 5 Y-βP-WFGG-RRRRR, YaWFGG-RRRRR 45% +5 Pain (152)
Aβ1-6A2VTAT(D) grkkrrqrrr-gggg-dvefrh 35% +8.1 AD (153)
DEETGE-CAL-TAT RKKRRQRRR-PLFAER-LDEETGEFLP-NH2 28% +5 GCI (154)
TAT-T406 RKKRRQRR-IAYSSSETPNRHDML 29% +7.1 Pain (155)
TAT-21-40 RKKRRQRRR-RIPLSKREGIKWQRPRFTRQ 38% +14 Excitotoxicity, stroke, OGD (156)
TAT-C1aB YGRKKRRQRRR-HLSPNKWKW 30% +10.1 Excitotoxicity, stroke (157)
TAT-2ASCV YGRKKRRQRRR-TVNEKVSC 31% +8 Pain (158)
TAT-NTS YGRKKRRQRRR-RSFPHLRRVF-NH2 43% +12.1 Stroke, OGD (159)
TAT-CBD3M5L YGRKKRRQRR-ARSRMA 44% +9 Pain (160)
TDP-r8 YrFG-rrrrrrrr-G 69% +9 Pain (161)
TAT-Pro-ADAM10 YGRKKRRQRR-PKLPPPKPLPGTLKRRRPPQP 27% +14 Huntington's disease (162)

At the N-terminus, Ac indicates acetyl and at the C-terminus NH2 indicates amide. Lower case single letter code indicates D-isoform of the amino acid. Aib, 2-Aminoisobutyric acid or 2-methylalanine; s-s, disulfide bond; AD, Alzheimer's disease; ALS, amyotrophic lateral sclerosis; EAE, Experimental autoimmune encephalomyelitis; GCI, global cerebral ischaemia; HIE, hypoxia-ischaemia encephalopathy; ICH, intracerebral hemorrhage; OGD, oxygen glucose deprivation; PD, Parkinson's disease; SAH, subarachnoid hemorrhage; SCI, spinal cord injury; SMA, spinal muscular atrophy; stroke, ischaemic stroke; TBI, traumatic brain injury.