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697 New Insights Into the Mechanisms of Argonaute Protein Competition and Implications for RNAi Gene Therapies Molecular Therapy Volume 23, Supplement 1, May 2015 Copyright © The American Society of G[.]

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Molecular Therapy Volume 23, Supplement 1, May 2015

Copyright © The American Society of Gene & Cell Therapy S277

Gene InhIbItIon

directly to cRGD peptide or to internalizing RGD (iRGD), which

is a variant RGD peptide that triggers permeabilization of tumor

endothelium and internalization by cells through secondary binding

to neuropilin The cRGD- and iRGD-conjugated KRAS Adaptors

were tested for efficacy against subcutaneous MiaPaca-2 xenografts,

and tumor growth was inhibited to equal or greater extent as with

the original cRGD-dendrimer system iRGD may be of particular

benefit for pancreatic adenocarcinomas, which have a densely fibrotic

stroma that impedes drug delivery Enhanced delivery of small- and

large-molecule therapeutics into primary pancreatic adenocarcinomas

in KPC mice has been achieved previously by conjugating or

co-injecting iRGD peptide

We have shown that U1 Adaptors can successfully target human

KRAS both in vitro and in vivo These results support the continued

development of U1 Adaptor technology as a strategy for therapeutic

suppression of KRAS in pancreatic cancer

695 Splice-Correction of X-Linked

Agammaglobulinemia in a Human BAC-Transgenic

Mouse Model Using Oligonucleotides

Burcu Bestas,1 Pedro M.D Moreno,1 Emelie K.M Blomberg,1

Dara K Mohammad,1 Karin E Lundin,1 Robert Månsson,2 Anna

Berglöf,1 Jesper Wengel,3 Edvard C.I Smith.1

1 Laborattory Medicine, Karolinska Institutet, Huddinge/Stockholm,

Sweden; 2 Medicine, Karolinska Institutet, Huddinge/Stockholm,

Sweden; 3 Physics Chemistry and Pharmacy, University of Southern

Denmark, Odense, Denmark.

The inherited immunodeficiency, X-linked agammaglobulinemia

(XLA), is caused by mutations in the BTK gene, and results in

a B-lineage developmental block We have recently assessed the

treatment potential of splice-correcting oligonucleotides (SCOs)

targeting a mutated BTK transcript, which contains a pseudo-exon

(Bestas et al., J Clin Invest 124: 4067, 2014) In order to study

the potential of SCOs, we engineered a novel, Bacterial Artificial

Chromosome (BAC)-transgenic mouse carrying a mutated human

BTK gene, originally found in an XLA family In order to avoid

any influence of mouse endogenous Btk protein, we bred the

BAC-transgenic mice onto a Btk knockout background In this model it

was possible to correct the defect both in pro-B-cells in vitro, and

also in mature B-cells, as demonstrated by the injection of SCOs in

vivo The corrected mRNA gave rise to a functional BTK protein

As a final proof-of-concept we were also able to correct the defect

in primary patient cells In this study we used different nucleotide

chemistries, 2’-O-methyl, locked nucleic acid (LNA) and morpholino

chemistries We have now included also other nucleotide chemistries

in order to make comparative studies This is to our knowledge the

first time that a lymphocyte defect, caused by abnormal splicing, has

been corrected in vivo using a splice-correction approach

696 Pre-Clinical Evaluation of Allele-Specific

Mutant Huntingtin Gene Silencing Antisense

Oligonucleotides

Amber L Southwell,1 Niels H Skotte,1 Nicholas Caron,1 Holly

Kordasiewicz,2 Michael Oestergaard,2 Crystal N Doty,1 Erika B

Villanueva,1 Yuanyun Xie,1 Boguslaw Felczak,1 Susan M Freier,2

Eric E Swayze,2 Punit P Seth,2 C Frank Bennet,2 Michael R

Hayden.1

1 Centre for Molecular Medicine and Therapeutics, University of

British Columbia, Vancouver, BC, Canada; 2 ISIS Pharmaceuticals,

Carlsbad, CA.

Huntington disease (HD) is a dominant, genetic neurodegenerative

disease characterized by progressive loss of voluntary motor

control, psychiatric disturbance, and cognitive decline, for which

there is currently no disease-modifying therapy HD is caused by the expansion of a CAG tract in the huntingtin (HTT) gene The mutant HTT protein (muHTT) acquires toxic functions, and there is significant evidence that muHTT lowering would be therapeutically efficacious However, the wild-type HTT protein (wtHTT) serves vital functions, making allele-specific muHTT lowering strategies potentially safer than non-selective strategies CAG tract expansion is associated with single nucleotide polymorphisms (SNPs) that can be targeted by gene silencing reagents such as antisense oligonucleotides (ASOs) to accomplish allele-specific muHTT lowering We have evaluated ASOs targeted to HD-associated SNPs in acute in vivo studies including screening, distribution, duration of action and dosing, using a humanized mouse model of HD, Hu97/18, that is heterozygous for the targeted SNPs We have identified four well-tolerated lead ASOs that potently and selectively silence muHTT at

a broad range of doses throughout the central nervous system for 36 weeks or more after a single intracerebroventricular injection

We next conducted a pre-clinical therapeutic efficacy trial of lead ASOs and evaluated them for effect on the HD-like phenotypes

of Hu97/18 mice Treated mice underwent longitudinal behavioral and biochemical assessment followed by terminal neuropathology Thus far we have determined that pre-symptomatic allele-specific muHTT silencing prevents onset of behavioral HD-like phenotypes Evaluation of neuropathology and post-symptomatic intervention is ongoing Contingent on findings from these studies and using delivery and dosing information gained from ongoing CNS ASO clinical trials,

a primary SNP-targeted ASO drug could be fairly rapidly translated for human applications

697 New Insights Into the Mechanisms of Argonaute Protein Competition and Implications for RNAi Gene Therapies

Mario Lederle,1 Kathrin Tegeler,1 Daniela Cerny,1 Patrick Zessin,1 Dirk Grimm.1

1 Department of Infectious Diseases/Virology, Heidelberg University Hospital, Cluster of Excellence CellNetworks, Heidelberg, Germany.

RNA interference (RNAi) has become an established tool for basic gene annotation and a promising option for therapeutic concepts that require specific inhibition of dysregulated or exogenous genes However, we and others accumulated strong evidence that over-expression of short hairpin RNAs (shRNAs, an expressable and widely used form of RNAi trigger) can cause cytotoxicity leading up

to organ damage and lethality in shRNA-treated animals A meticulous dissection of the underlying mechanisms is critical as it will ultimately guide the design of safer RNAi therapies and thus foster their clinical translation One informative finding was that co-expression of Ago2 (Slicer, as it cleaves targeted mRNA), a key component of the RNA-induced silencing complex RISC, increases shRNA efficiency in cells and animals, implying that Ago2 is a rate-limiting factor whose saturation may be involved in cytotoxicity Moreover, over-expression

of the three other mammalian Ago proteins - Ago1, Ago3 or Ago4 (all slicing-incompetent) - dampens RNAi potency, underscoring that a delicate balance of all four Ago variants is key to efficient and safe gene silencing Here, we investigated three hypotheses that could explain how relative Ago1-4 levels may affect the functionality of shRNA-loaded RISC and hence the strength of target knockdown: over-abundance of non-Slicer Ago proteins could (i) dysregulate Ago2 expression (transcription and/or translation), (ii) re-localize intra-cellular Ago2 away from sites of RNAi activity, or (iii) quantitatively sequester shRNAs into slicing-incompetent RISC Using codon-optimized Ago variants that are distinguishable by real-time PCR from the endogenous counterparts, as well as Western blotting,

we could eliminate model (i) Likewise, wide-field and confocal

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Molecular Therapy Volume 23, Supplement 1, May 2015 Copyright © The American Society of Gene & Cell Therapy

S278

Gene InhIbItIon

microscopy analyses revealed no evidence for Ago2 re-localization

upon Ago1/3/4 over-expression and vice versa We therefore focused

on hypothesis (iii) and fused the Ago1-4 cDNAs with GFP or RFP

tags that enabled their specific immunoprecipitation, alone or in

combination, using the TRAP system Indeed, we found that Slicer

and non-Slicers compete for shRNA loading and mRNA knockdown

in a dose-dependent manner, readily explaining all prior observations

of RNAi enhancement by Ago2 over-expression, or shRNA inhibition

by Ago1/3/4, respectively Importantly, the combination of GFP/

RFP-Ago fusion constructs with the co-precipitation protocols newly

established during this work should be widely useful to address other

vital questions in the RNAi field, such as the selective processing of

different cellular small RNAs by the four Ago proteins Moreover,

our notion that over-expression of non-Slicers has no adverse effect

on endogenous Ago2 levels or localization paves the way for novel

strategies to fine-tune therapeutic RNAi in a safe and potent manner

As a prototype, we present a new shRNA/Ago co-expression system

whereby Ago1/2 levels can be regulated independently through

exogenous macrolides, permitting a tight control of target knockdown

efficiencies and providing an original avenue to reduce toxicity in

future RNAi gene therapy applications

698 Eliminating TLR9+ Prostate Cancer Stem

Cells In Vivo Using NF-κB/RELA- or

STAT3-Targeting CpG-siRNA Conjugates

Dayson F Moreira,1 Qifang Zhang,1 Dewan M Hossain,1

Sergey Nechaev,1 Haiqing Li,1 Claudia M Kowolik,1 Massimo

D’Apuzzo,3 Stephen Forman,1 Jeremy Jones,1 Sumanta K Pal,1

Marcin Kortylewski1.1

1 Beckman Research Institute at City of Hope, Duarte, CA.

Treatment of hormone-refractory and frequently reoccurring

prostate cancers is a major clinical challenge Here, we demonstrate

that Toll-like Receptor 9 (TLR9) is commonly upregulated in

late-stage prostate cancers and provides a potential therapeutic target As

a sensor of immunogenic cell death, TLR9 bridges intra-prostatic

inflammation to cancer stem cell phenotype Our limited dilution/

serial transplantation experiments in vivo demonstrate that TLR9

is essential for prostate cancer cells’ potential to propagate and

self-renew In addition, low expression or silencing of TLR9 limits

the clonogenic potential and mesenchymal stem cell-like properties

of prostate cancer cells Genome-wide transcriptional analysis

of cancer cells isolated from xenotransplanted prostate tumors

revealed a unique TLR9-dependent gene expression signature

Further analysis of the TLR9 downstream signaling indicated that

tumorigenic transcription factors NF-κB/RELA and STAT3 cooperate

to orchestrate expression of stem cell-related genes Both RELA and

STAT3 bound and co-regulated promoters of NKX3.1 and KLF4

prostate cancer stem cell-related genes We further demonstrated the

feasibility of targeting prostate cancer-propagating potential in vivo

by TLR9-targeted siRNA delivery using CpG-siRNA conjugates

Local administration of CpG-RELAsiRNA or CpG-STAT3siRNA

but not control conjugates, inhibited tumor growth and cancer cell

clonogenic potential in two xenotransplanted prostate cancer models

Selective elimination of tumor-propagating cells using TLR9-targeted

blockade of NF-κB/RELA and STAT3 signaling has potential for

clinical translation to benefit patients with late-stage prostate cancers

699 shRNA Sense Strand Neutralization Reduces Off-Targeting, Ameliorates Toxicity and Enhances Efficacy of RNAi-Based HBV Gene Therapy

Thomas Michler,1 Stefanie Grosse,2 Stefan Mockenhaupt,2 Ulrike Protzer,1 Dirk Grimm.2

1 Virology, Technical University, Munich, Germany; 2 Virology, University of Heidelberg, Heidelberg, Germany.

A decade after first reports of successful inhibition of hepatitis B virus (HBV) replication in vivo using shRNAs, only a single RNAi therapeutic is under phase II efficacy investigation in Hepatitis

B patients This slow clinical translation is puzzling, as current standard-of-care requires life-long and daily drug treatment which is often accompanied by side effects and occurrence of resistant HBV mutants Also, while preventing the generation of infectious HBV virions, reverse-transcriptase inhibitors fail to suppress HBV surface and e-antigen, which enable viral persistence by modulating the host immune system and contribute to HBV-associated hepatocellular carcinoma These problems may be solved using RNAi as anti-HBV therapeutic since (i) long-term suppression can be achieved by a single dose of an shRNA-encoding vector, and (ii) all four HBV transcripts share a common 3 end, allowing concurrent inhibition

of HBV pre-genomic RNA and all viral proteins with a single RNAi molecule, and potentially restoring antiviral immunity and preventing carcinogenesis Still, clinical translation remains hampered

by safety concerns as shRNA over-expression in the mouse liver causes elevated liver transaminases, jaundice and weight loss One explanation is that ectopic RNAi triggers overload the endogenous miRNA pathway, perturbing miRNA biogenesis and/or activity, and causing cytotoxicity Evidence for this saturation model is that (i) shRNAs circumvent Drosha processing, a gatekeeper for miRNA biogenesis, (ii) embedding shRNAs in a recombinant miRNA context yields lower and safer levels of mature RNAi triggers, and (iii) overexpression of Argonaute-2 (Ago2) ameliorates toxicity and enhances RNAi efficacy Moreover, RNAi triggers can perturb cell physiology through unwanted inhibition of off-target genes with partial complementarities to one of the two strands of the RNAi molecule To alleviate such unintended gene silencing, we have developed a novel bi-cistronic AAV vector that expresses, in addition to the shRNA, a second RNA hairpin called tough decoy or TuD In cell culture, the TuD effectively sequestered and inactivated shRNA sense strands, and thereby improved RNAi specificity These remarkable features translated well into an HBV-transgenic mouse model, where sense strand off-target activity was likewise prevented,

as validated by transcriptome analysis of liver RNA, coinciding with ameliorated toxicity Enhanced in vivo safety was also noted for two alternative AAV vectors, either co-expressing the shRNA and Ago2,

or embedding the shRNA in a miR-122 backbone Notably, the new shRNA/TuD vector outperformed the other expression strategies regarding efficacy with stable HBV reduction of up to 98% over

3 months We speculate that its enhanced antiviral efficacy results from increased loading of the desired antisense strand into RISC in absence of the sense strand, possibly also further amending toxicity

by attenuating RISC saturation The simple TuD design and the versatility of our new AAV vector pave the way for adaptation of our strategy to other applications and should facilitate clinical translation

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