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TABLE 16.1 Examples of Transport of Nucleic Acids by CPP Cargo type Cargo function Target cell/tissue Applied CPP Conjugation method Ref. ss and ds 18- and 36-mers fibroplasts HS68 and NIH-3T3 cells MPG fusion sequence of HIV gp41 + nuclear localization sequence of SV40 T-antigen Ac-GALFLGFLGAAGSTMGA-WSQ- PKSKRKV-cysteamide antisense 26-mer cardiac H9C2 cells MTS-NLS N-terminal part — signal peptide sequence of cayman crocodylus Ig(v) light chain- C-terminal part- simian virus SV40 large tumor antigen (T-ag) NLS MGLGLHLLVLAAALQGAKKKRKV plasmids Plasmid pRL-SV40 Plasmid pcDNA3.1NT-GFP HS-68, NIH-3T3, C2C12 myoblasts, COS-7, CEM-SS λ phage Recombinant λ phage particle COS-1; VA13/2RA; 293; NIH 3T3; HeLa; A431; mouse liver in vivo a Sequence according to the original report. MPG fusion sequence of HIV gp41 + nuclear localization sequence of SV40 T-antigen Ac-GALFLGFLGAAGSTMGA-WSQ- PKSKRKV-cysteamide Tat (43–60) MLGISYGRKKRRQRRRPPQT Noncovalent conjugation 57 S–S 55 Noncovalent conjugation 59 Application of recombinant λ phage displaying Tat peptide on its surface 49 Cell-Penetrating Peptides as Vectors for Delivery of Nucleic Acids 353
354 Cell-Penetrating Peptides: Processes and Applications Recently, Eguchi et al. 49 demonstrated utilization of Tat peptide for gene transfer to various mammalian cell lines (Table 16.1). They constructed recombinant lambda phage particles displaying Tat peptide on their surfaces and carrying mammalian marker genes (GFP and firefly luciferase) as part of their genomes. In animal cells briefly exposed to Tat–phage, the expression of phage marker genes was induced. In contrast, recombinant phage displaying other functional peptides, like the peptide from integrin-binding domain (RGD) or a nuclear localization signal (NLS) similar to controls (wild type phage and naked DNA), could not induce detectable marker gene expression. The expression of marker genes induced by Tat–phage is not affected by temperature or by endosomotropic agents such as monensin and chloroquine (for further reading, see Galloway et al. 50 ). However, expression is partially impaired by inhibitors of caveolae formation such as nystatin 51 and fillipin. 52 The ability of Tat–phage to transport dsDNA was also shown in vivo. Constructs carrying GFP gene were injected intraparenchymally into the mouse liver. Livers were dissected in 2 days and the GFP expression observed by fluorescence microscopy. These data suggest that Tat peptide could be used in gene transfer in vivo. Also, the existence of classical endocytic pathway-independent mechanism for Tatmediated uptake was obviously proved again. 16.4.3 TRANSPORTAN Transportan is a 27-amino acid artificial chimeric peptide developed by the authors. 37 This peptide has been demonstrated to locate through biological membranes and finally localize in the nucleus. The ability to transport antisense PNA molecules in vitro and in vivo was demonstrated in the article published shortly after introducing the peptide. 44 This report compares penetratin and transportan as transmembrane delivery vectors for PNA molecules, demonstrating similar effects with both vectors (see above). A recent publication by Beletskii et al. 53 reported use of transportan to target several PNA oligomers to diploid female ES cells (Table 16.1). The study demonstrates the effect of PNA oligomers on inactivation of X chromosome (Xi) and formation of macrochromatin body (MCB). This complex is formed to silence the X chromosome in female mammals. Transportan coupled with 19-mer PNA complementary to noncoding Xist RNA, a noncoding RNA responsible for Xi silencing, inhibits Xist complexing with Xi. Such inhibition leads to disruption of association between Xi and H2A macrohistone. These results prove the potential of transportan as a cell-penetrating delivery vehicle. 16.4.4 MTS–NLS Certain peptides from protein signal sequences have been shown to penetrate membranes. Membrane translocating sequences (MTS) are used by protein sorting machinery to target proteins into different intracellular compartments. Nuclear proteins are usually tagged with nuclear localization signals (NLSs). MTS–NLS conjugates have shown the ability to enter cells and localize in the nucleus. 54 Peptides containing a hydrophobic N-terminal part (signal peptide sequence of Cayman crocodylus Ig(v) light chain) associated with an NLS (the C-terminal
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CELL- PENETRATING PEPTIDES Processe
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Pharmacology and Toxicology: Basic
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Library of Congress Cataloging-in-P
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to the handbook are prominent resea
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REFERENCES 1. Green, M. and Loewens
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Contributors Mats Andersson Microbi
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Erin T. Pelkey Department of Chemis
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Contents Section I Classes of Cell-
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Chapter 16 Cell-Penetrating Peptide
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The Tat-Derived Cell-Penetrating Pe
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24 Cell-Penetrating Peptides: Proce
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3 Transportans Margus Pooga, Mattia
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Transportans 55 Indeed, galparan is
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Transportans 57 especially the endo
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Transportans 59 In the penetration
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Transportans 61 A 21-mer antisense
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Transportans 63 Commonly, the prote
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Transportans 65 antibiotin antibodi
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Transportans 67 For cross-linking o
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Transportans 69 and still retain ef
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Model Amphipathic Peptides 73 its D
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Model Amphipathic Peptides 75 pmol/
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TABLE 4.1 Internalization of Peptid
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TABLE 4.2 Internalization of Peptid
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Model Amphipathic Peptides 81 relat
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Model Amphipathic Peptides 87 A cat
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Model Amphipathic Peptides 89 At th
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Model Amphipathic Peptides 91 16. H
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Signal Sequence-Based Cell-Penetrat
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116 Cell-Penetrating Peptides: Proc
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Interactions of Cell-Penetrating Pe
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Structure Prediction of CPPs and It
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FIGURE 9.7 (Color Figure 9.7 follow
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FIGURE 9.8 The center of the figure
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Biophysical Studies of Cell-Penetra
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Toxicity and Side Effects of Cell-P
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Kinetics of Uptake of Cell-Penetrat
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Page 350 and 351: Cell-Penetrating Peptide Conjugatio
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Page 370 and 371: Cell-Penetrating Peptides as Vector
Page 380 and 381: CPP 2 Cargo or mRNA CAP Antisense A
Page 384: Cell-Penetrating Peptides as Vector
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Page 418 and 419: Index A Abaecin, 129 Abz radiolabel
Page 420 and 421: Index 399 Diffraction, 168 Disulphi
Page 422 and 423: Index 401 structure prediction, 187
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Index 403 pRB proteins, Tat-E1A bin
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Index 405 lipid perturbation (secon
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