MF59 and Pam3CSK4 Boost Adaptive Responses to Influenza Subunit Vaccine through an IFN Type I-Independent Mechanism of Action This information is current as of September 25, 2021. Elena Caproni, Elaine Tritto, Mario Cortese, Alessandro Muzzi, Flaviana Mosca, Elisabetta Monaci, Barbara Baudner, Anja Seubert and Ennio De Gregorio J Immunol 2012; 188:3088-3098; Prepublished online 20 February 2012; Downloaded from doi: 10.4049/jimmunol.1101764 http://www.jimmunol.org/content/188/7/3088 http://www.jimmunol.org/ Supplementary http://www.jimmunol.org/content/suppl/2012/02/21/jimmunol.110176 Material 4.DC1 References This article cites 33 articles, 12 of which you can access for free at: http://www.jimmunol.org/content/188/7/3088.full#ref-list-1

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The Journal of Immunology is published twice each month by The American Association of Immunologists, Inc., 1451 Rockville Pike, Suite 650, Rockville, MD 20852 Copyright © 2012 by The American Association of Immunologists, Inc. All rights reserved. Print ISSN: 0022-1767 Online ISSN: 1550-6606. The Journal of Immunology

MF59 and Pam3CSK4 Boost Adaptive Responses to Influenza Subunit Vaccine through an IFN Type I-Independent Mechanism of Action

Elena Caproni,*,1 Elaine Tritto,†,1 Mario Cortese,* Alessandro Muzzi,* Flaviana Mosca,* Elisabetta Monaci,* Barbara Baudner,* Anja Seubert,* and Ennio De Gregorio*

The innate immune pathways induced by adjuvants required to increase adaptive responses to influenza subunit vaccines are not well characterized. We profiled different TLR-independent (MF59 and alum) and TLR-dependent (CpG, resiquimod, and Pam3CSK4) adjuvants for the ability to increase the immunogenicity to a trivalent influenza seasonal subunit vaccine and to tetanus toxoid (TT) in mouse. Although all adjuvants boosted the Ab responses to TT, only MF59 and Pam3CSK4 were able to enhance hemagglutinin Ab

responses. To identify innate immune correlates of adjuvanticity to influenza subunit vaccine, we investigated the signatures Downloaded from induced by each adjuvant in vitro in splenocytes and in vivo in muscle and lymph nodes using DNA microarrays. We found that flu adjuvanticity correlates with the upregulation of proinflammatory and other genes involved in leukocyte transendothelial migration at the vaccine injection site. Confocal and FACS analysis confirmed that MF59 and Pam3CSK4 were the strongest inducers of blood recruitment in the muscle compared with the other adjuvants tested. Even though it has been proposed that IFN type I is required for adjuvanticity to influenza vaccines, we found that MF59 and Pam3CSK4 were not good inducers of IFN-related innate immunity pathways. By contrast, resiquimod failed to enhance the adaptive response to flu despite a strong activation of the IFN http://www.jimmunol.org/ pathway in muscle and lymph nodes. By blocking IFN type I receptor through a mAb, we confirmed that the adjuvanticity of MF59 and Pam3CSK4 to a trivalent influenza vaccine and to TT is IFN independent. The Journal of Immunology, 2012, 188: 3088–3098.

here are different types of influenza vaccines with protection against virus challenge (7). Alternatively, subunit flu distinct immunogenicity profiles (1–3). Live-attenuated vaccine formulation immunogenicity could be restored by incor- T virus vaccines induce a subclinical infection and there- poration of IFN inducers such as the TLR9 agonist (5). fore are very efficacious (4). Formalin-inactivated whole-virus Clinical experience has shown that the oil-in-water emulsions are vaccines (WV) contain viral RNA, which activates IFN type I extremely efficient in increasing the immunogenicity of flu subunit responses that are required to boost the adaptive immune re- vaccines. The adjuvant MF59 is an oil-in-water emulsion composed by guest on September 25, 2021 sponses (5). Subunit virus vaccines contain hemagglutinin (HA) of small droplets of squalene surrounded by a monolayer of nonionic and neuraminidase Ags, next to some residual structural detergents (8, 9). Different aspects contribute to MF59 adjuvanticity. (M1 and NP), but lack most of the viral RNA, except from some Several studies have shown that MF59 can increase Ag uptake by residual RNA that may be associated with NP (6). These vaccines APCs (10–12) besides inducing the activation of innate immunity are safer compared with WV, but they are also less immunogenic (13). In vitro studies have shown that MF59 induces the secretion of and therefore require an adjuvant when administered to previously in human macrophages, monocytes, and granulocytes unexposed or immunocompromised subjects. It has been proposed (14). We have previously demonstrated that i.m. injection of MF59 that the best strategy to boost subunit flu vaccine formulation is to induces the local activation of innate immune reactions leading to restore IFN type I-associated activities that are lost during the recruitment of blood cells in the muscle (13). Recently, it has been purification process. Accordingly, coadministration of type I rIFN shown that all the recruited cell types can take up both Ag and with a subunit influenza vaccine in mice was shown to increase adjuvant and that MF59 promotes the migration of Ag/adjuvant double-positive cells to the draining lymph nodes (LNs) (15). A MF59-adjuvanted influenza vaccine was developed initially *Research Center, Novartis Vaccine and Diagnostics, 53100 Siena, Italy; and for seasonal vaccination in the elderly to enhance Ab titers and †Preclinical Safety Genomics, Novartis Institute of Biomedical Research, CH-4057 seroconversion and seroprotection rates (9). More recently it was Basel, Switzerland shown that also in young children MF59 enhances seroconver- 1 E.C. and E.T. contributed equally to this study. sion and cross-protection against flu strains mismatched to the Received for publication June 17, 2011. Accepted for publication January 13, 2012. Ags of a seasonal subunit flu vaccine (16). MF59 and a second The microarray data presented in this article have been submitted to the ArrayExpress oil-in-water emulsion, called AS03, have also been successfully database, European Molecular Biology Laboratory-European Bioinformatics (http:// www.ebi.ac.uk/), under accession number E-MTAB-942. used for the development of prepandemic flu vaccines to protect from a potential H5N1 pandemic influenza outbreak (17, 18). The Address correspondence and reprint requests to Dr. Ennio De Gregorio, Novartis Vaccine and Diagnostics, Via Fiorentina 1, 53100 Siena, Italy. E-mail address: clinical data have shown that MF59 increases cross-reactive neu- [email protected] tralizing Ab titers and memory responses to H5 (17, 19). Recently, The online version of this article contains supplemental material. MF59 and AS03 have also been used during the vaccination cam- Abbreviations used in this article: DC, dendritic cell; HA, hemagglutinin; HI, hem- paign in the 2009–2010 H1N1 pandemic outbreak (20). agglutination inhibition; KO, knockout; LN, lymph node; TT, tetanus toxoid; WV, The ability of emulsions to increase the immunogenicity of sub- whole-virus vaccine. unit flu vaccines seems to be quite unique. Clinical trials have shown Copyright Ó 2012 by The American Association of Immunologists, Inc. 0022-1767/12/$16.00 that the administration of MF59-adjuvanted H5N1 induces higher www.jimmunol.org/cgi/doi/10.4049/jimmunol.1101764 The Journal of Immunology 3089 hemagglutination inhibition (HI) and microneutralization titers to measured by endpoint ELISA. MaxiSorp 96-well plates were coated with flu compared with the administration of the same vaccine formulated 2.5 mg/well flu Ag or 0.2 mg/well TT Ag in 100 ml/well PBS. Goat anti- with alum (21). Furthermore, mouse studies have shown that MF59 mouse IgG (Southern Biotechnology Associates), goat anti-mouse IgG1 (Southern Biotech), or goat anti-mouse IgG2a (Southern Biotechnology enhances HI titers to seasonal flu Ags compared with alum and CpG Associates) Abs, all conjugated with alkaline phosphatase, were used at (22). However, the innate immune pathways required for the adju- 1:2000 dilution. The reaction was developed by addition of 100 ml/well vant effect of oil-in-water emulsions to influenza subunit vaccines p-nitrophenyl phosphate disodium salt substrate (Sigma-Aldrich), and ab- and the exact role played by IFN type I are not well characterized. sorbance at 405 nm was measured by Spectramax plate reader. ELISA titers were expressed as the reciprocal dilution that gave an OD higher than In this work, we investigated the innate immune reactions that are the average OD of the blanks plus 3 times the SD. associated with adjuvanticity to influenza subunit vaccines in mouse using a systems biology approach. First, we monitored the ability of Measurement of flu HI inhibition titers MF59, alum, and three TLR-dependent adjuvants, CpG, resiquimod, Sera were pretreated with DENKA receptor-destroying enzyme (Bio- and Pam3CSK4, to increase the immunogenicity of a commercial flu genetics) and then inactivated at 56˚C for 30 min. Two-fold serial seasonal subunit vaccine and to tetanus toxoid (TT). We found that dilutions of 25 ml pretreated sera and positive control sera were treated m despite the fact that all adjuvants were able to increase the immu- with 25 l virus working dilution. Samples were incubated at room temperature; 50 ml turkey RBC suspension was dispensed in each well; nogenicity of TT, only MF59 and to a less extent Pam3CSK4 were and plates were again incubated at room temperature. The test endpoint able to adjuvant the flu vaccine. Then, we analyzed the innate was determined by visual inspection for an agglutination reaction: a red immune signatures induced by each adjuvant in vitro in splenocytes, dot formation indicated a positive reaction (inhibition), whereas a diffuse and in vivo at the injection site (muscle) and in the draining LNs patch of cells a negative reaction (hemagglutination). The titer was de- fined as the highest serum dilution at which hemagglutination was in- using DNA microarrays. Surprisingly, we found that the activation hibited, and the Ab concentration corresponds to the reciprocal value Downloaded from of IFN type I pathway is not required for the adjuvanticity of MF59 of the titer. and Pam3CSK4. In contrast, we found that despite a potent acti- In vitro splenocyte stimulation for microarray analysis vation of IFN type I responses, resiquimod did not trigger any adjuvant effect to the flu vaccine. Ultimately, we identified a gene For microarray experiments on splenocytes, 106 cells/well were treated for signature at the injection site that correlates with adjuvanticity to 6 h with PBS, MF59 (at 1:300 dilution), 40 mg/ml alum, 1 mM CpG, 1% DMSO, 5 mM R848, and 1 mM Pam3CSK4. influenza subunit vaccine. http://www.jimmunol.org/ Muscle, LN, and splenocyte RNA extraction and purification Materials and Methods For muscle RNA extraction, whole muscles were homogenized in 7.5 ml Adjuvants TRIzol (Invitrogen) with an Ultra-Turrax T25 (IKA), and total RNA was MF59 is a Novartis proprietary oil-in-water emulsion consisting of the 5% oil extracted following the manufacturer’s protocol. One hundred micrograms squalene with the two surfactants, 0.5% Tween 80 and 0.5% Span-85, in of RNA from each muscle was purified using the RNeasy RNA purification a sodium citrate buffer (10 mM). The alum used in this study is aluminum columns (Qiagen) following the producer’s protocol. Residual DNA was removed by an additional on-column DNase I digestion step using the hydroxide (Al-OH3). The CpG oligonucleotide sequence used was 59-TCC ATG ACG TTC CTG ACG TT-39 with all phosphothioate backbones (CpG Qiagen RNase-free DNase set (Qiagen). RNA quality was assessed by using the 2100 Bioanalyzer automated microfluidic system in combination

1826). R848 (resiquimod) is an imidazoquinoline resuspended at 100 mM in by guest on September 25, 2021 100% DMSO. Pam3CSK4 (Invivogen) is synthetic triacylated lipoprotein with the RNA 6000 Nano (Agilent) following the producer’s protocol. resuspended at 1 mg/ml in MilliQ water. For LN RNA extraction, the draining inguinal LNs from each mouse were collected in liquid nitrogen and immediately homogenized on ice in lysis Antigens buffer (Qiagen) using the Ultra Turrax T25 (IKA). The tissue lysates were transferred in a QIAshredder column (Qiagen), and RNA was purified using Trivalent flu Ags present in the 2007/2008 seasonal flu vaccine formulation the RNeasy RNA purification columns (Qiagen) following the manufac- used in this study were egg-derived H1N1 (A/Brisbane/59/2007 H1N1-like turer’s protocol, as described before. RNA from splenocytes was extracted virus), H3N2 (A/Brisbane/10/2007 H3N2-like virus), and B (B/Florida/4/ using the Qiagen QIAshredder and RNeasy RNA purification kit (Qiagen) 2006-like virus). TT Ag was prepared and purified in house. All Ags were following manufacturer’s protocol. For all microarray analyses shown in diluted in PBS. this study, three independent experiments were conducted for each group. Mice and immunizations RNA labeling, microarray hybridization, and data analysis Pathogen-free female BALB/c mice 6–8 wk of age were housed and treated For each hybridization, 400 ng RNA was labeled using the QuickAmp according to internal animal ethical committee and European guidelines. labeling kit (Agilent). Muscle or LNs RNAwere labeled with cyanine-3 dye In all experiments, mice were injected i.m. in both quadriceps muscles for one-color microarray analysis. RNA from splenocytes treated with with 50 ml dose per quadriceps with PBS, MF59 (1:1 dilution in PBS), 200 adjuvants were labeled with Cy5 and cohybridized with cyanine-3–labeled mg alum, 20 mg CpG (all in PBS), 1% DMSO in PBS, 25 mg R848, 10 mg RNA extracted from control buffer-treated splenocytes (reference) for two- Pam3CSK4 (both in PBS, 1% DMSO), flu (a mix of H1N1, H3N2, and B color microarray analysis. The efficiency of incorporation of the Cy3 or Ags; 0.1 mg each), and flu plus MF59. For adjuvanticity experiments, mice Cy5 dyes was measured by Nanodrop analysis. A total of 1.65 mg labeled were injected with the aforementioned adjuvants in presence or absence of cRNA was hybridized onto the Agilent 4 3 44k Whole Mouse Genome either 0.1 mg TT Ag or a mix of H1N1, H3N2, and B Ags (0.1 mg each). Microarray chip, detecting .40,000 transcripts, for 17 h at 60˚C following Mice were immunized at days 0 and 21. Sera were collected 2 wk after Agilent protocol. Images were acquired using the Agilent Scanner (Agi- the first and the second immunization for Ab analysis. In Ab-blocking lent) at a resolution of 5 mm in extended dynamic range. experiments, mice were injected i.p. with 50 ml20mg anti–IFN-ab R2 Raw images were preprocessed with the Agilent’s Feature Extraction 9.5 (Space Import-Export; bulk) or isotype control (goat IgG; R&D Systems) Image Analysis Software. Feature intensities and ratios were calculated, 1 d before and 7 d after the first immunization. Frequencies of influenza- and flags for bad quality were assigned to each spot. The BASE 1.2.17 specific T cells in the spleens of four mice/group were measured 2 wk after database/analysis software was used for further data processing. Intensities second immunization. responses in LNs were measured on pools of for each spot were normalized by the global median, and the local back- four mice/group 3 wk after the second immunization. For microarray and ground was subtracted. The average ratio of the experimental replicates confocal microscopy experiments, mice were sacrificed 6 h after treatment was estimated, and aquantile normalization was applied. For multiple and quadriceps muscles and inguinal LNs were collected. For flow cyto- hypothesis testing and type I error rate control, false discovery rate sta- metric analysis of muscle cell composition or inguinal LN cell activation, tistics based on the estimation of the q values for each t test were applied mice were sacrificed 24 h after treatment. (23). Significantly regulated probes were selected by using the following . , Ag-specific Ab measurement cutoff: intensity ratio 4 (log2 ratio $|2|) and p value 0.05 in at least one treatment. Hierarchical clustering was performed with the TMEV 4.3.02 Sera from mice injected with Ag with or without adjuvant twice were software on the log2 transformed dataset applying the Euclidean distance collected 2 wk post-second immunization, and flu- or TT-specific Abs were matrix and the average linkage clustering method. 3090 INNATE IMMUNE SIGNATURES OF VACCINE ADJUVANTS

Data deposition were calculated by subtracting the background measured in the corre- sponding negative control for each . The complete set of microarray data have been submitted to the ArrayExpress database, European Molecular Biology Laboratory-European Bioinformat- Quantitative real-time PCR ics (ArrayExpress accession: E-MTAB-942, www.ebi.ac.uk/arrayexpress). Correlation analysis between profiles in the muscle and IgG The expression levels of five genes (Cxcl10, Il6, Ccl2, Tnfa, Mx1) induced titers was calculated as follows. First, the ratios of both gene expression in- by injection of PBS, MF59, alum, and CpG (three mice for each group) tensity levels and IgG titers in each animal induced by each adjuvant were were measured by quantitative RT-PCR using the same RNA samples calculated against the corresponding nonadjuvanted formulations (MF59, previously used for microarray analysis. A total of 400 ng RNA from each alum, and CpG against PBS, R848, and Pam3CSK4 against DMSO, respec- muscle was used for RT-PCR using qScript cDNA SuperMix kit (Quanta tively). Then the ratio values were log2 rescaled, and averages and variances Biosciences), according to the manufacturer’s protocol. Quantitative RT- were calculated. Finally, the linear correlation fit between gene expression and PCR was performed for each cDNA sample in duplicate using Perfecta IgG log2 ratios, weighted on both gene expression and IgG log2 ratio rescaled qPCR FastMix, UNG, Low Rox mix (Quanta Biosciences), and Applied variances, was evaluated. Supplemental Table II shows gene expression values Biosystems TaqMan probes (Cxcl10, IL6, Ccl2, TNFa, Mx1) in Stratagene for the 1410 selected genes together with the correlation data between gene MX3000P System. The thermal cycling program was as follows: an initial expression profiles and influenza vaccine Ab titers for all the genes. 10 min at 45˚C and 3 min at 95˚C, followed by 40 cycles of 15 s at 95˚C and 1 min at 60˚C. The expression level of Gapdh was used as the internal Confocal analysis of muscle cryosections control to calculate the D cycle threshold. OCT-embedded muscle tissue was sectioned transversely at 5 mm, thaw mounted onto Superfrost glass slides, and quickly air dried. Slides were Results fixed in a 4% formaldehyde solution on ice and blocked in 3% BSA, 1% The efficacy of vaccine adjuvants is Ag dependent: MF59 and saponin in PBS. Slides were then stained with goat anti-mouse

Pam3CSK4 are the best adjuvants for a trivalent flu subunit Downloaded from polyclonal IgG (Santa Cruz Biotechnology; 1:50 dilution) and rat anti- mouse CD11b (AbD Serotec; 1:100 dilution), followed by a second-step vaccine incubation with chicken anti-goat AlexaFluor 488 (Molecular Probes; To assess how different classes of adjuvants enhanced Ab titers to 1:200 dilution) and goat anti-rat AlexaFluor 555 (Molecular Probes; 1:200 dilution) secondary Abs. Utrophin, a cytoskeletal playing a role in a coadministered Ag, BALB/c mice were immunized i.m. with TTor anchoring the to the plasma membrane and located in the with seasonal flu Ags (flu) either alone or in combination with the sarcolemma of muscle cells, was used to detect muscle fibers. CD11b/ following adjuvants: the oil-in-water emulsion MF59 and the mineral

Itgam was used as a marker for monocytes, macrophages, and granu- salt alum (both TLR-independent adjuvants), the oligonucleotide http://www.jimmunol.org/ locytes. Nuclei were detected using ToPro3 (Molecular Probes; 1:1000 CpG, the small molecule resiquimod (R848), and the lipopeptide dilution). Slides were mounted with ProLong gold antifade reagent (Invi- trogen), coverslipped, and imaged on a LSM 710 microscope (Zeiss). Pam3CSK4 (agonists of TLR9, TLR7/8, and TLR2, respectively). Two weeks post-second immunization, the Ab titers to both Ags and FACS analysis of LN single-cell suspensions the HI titers to flu were measured. PBS-injected mice were used as Inguinal LNs, excised from mice 24 h after adjuvant injection, were cut on ice; control group for MF59, alum, and CpG, whereas 1% DMSO-treated resuspended in 0.5 ml RPMI 1640 medium (Life Technologies), 16 mlcol- mice were the control group for R848 and Pam3CSK4. MF59 and, to lagenase D (Roche), and 25 ml DNase I (Roche); and incubated at 37˚C for a lesser extent, Pam3CSK4 significantly enhanced total IgG titers to 15 min under gentle agitation. The tissue was completely dissociated with Gentlemax Dissociator (Miltenyi Biotec), followed by an additional 15-min H1N1, H3N2, and B Ags (Fig. 1A). The HI titers reflected what was incubation at 37˚C. Cells were then resuspended in PBS, 5% FCS and observed for total IgG titers: MF59 and Pam3CSK4 were the only by guest on September 25, 2021 counted. A total of 1 3 106 cells/well was plated in duplicate, incubated for adjuvants that induced significantly enhanced HI titers to H3N2 (Fig. 20 min with 20% rabbit serum, and stained with two Ab combinations. For T, 1B). Isotype analysis of Ab titers showed that MF59 increased spe- B, and NK cells, a mix of anti–DX-5 biotin (BD Pharmingen), anti-CD69 cific IgG1 and, to a lesser extent, IgG2a for all influenza Ags (Fig. 1C, PerCP-Cy5.5 (eBioscience), anti-CD3 allophycocyanin-eFluor780 (eBio- science), anti-CD19 A-700 (eBioscience), and Live/Dead Aqua (Invitrogen) 1D). Pam3CSK4 enhanced IgG2a and, to a lesser extent, IgG1 only in was used. For dendritic cells (DCs), anti-CD11c PerCP-Cy5.5, anti-CD86 response to H3N2. R848 and alum were not effective adjuvants for flu allophycocyanin (Invitrogen), anti-MHCII A-700 (eBioscience), and Live/ Ags except for a modest effect of R848 on H3N2 IgG2a response. Dead Aqua (Invitrogen) were used. Anti–DX-5 biotin was detected with CpG did not boost flu total Abs and HI titers, but induced a significant a second-step incubation with streptavidin Pacific Blue (Invitrogen). Cells were resuspended in FACS buffer before acquisition on a FACS LSRII flow increase of IgG2a titers (Fig. 1D). Consistently with the strong effect cytometer (BD Biosciences). of MF59 on IgG1 responses to influenza vaccine, we found that MF59 was the only adjuvant that significantly increased Ag-specific T cell cytokine response IL-4+/IL-13+ CD4 T cells in the spleen (Fig. 1E). To better evaluate To identify influenza-specific T cells, 1.5 3 106 cells/well were prepared the quality of the CD4 response, we extended the analysis to other from spleens or pools of LNs excised from four mice/group, and stimu- . We found that the CD4 response to MF59-adjuvanted lated with 1 mg/ml anti-CD28 with a mix of H1N1, H3N2, and B Ags at vaccine was dominated by IL-4+/IL-13+ cells; however, we could 0.3 mg/ml total concentration. As a positive control, the same number of + + cells was incubated with a mix of anti-CD3 and anti-CD28. As a negative also detect high frequencies of IL-17 and IL-2 CD4 T cells and control, cells were incubated only with anti-CD28. After an overnight CD4 T cells expressing multiple cytokines (Fig. 1F). We found that stimulation, brefeldin A (2.5 mg/ml) was added for additional 4 h to inhibit CpG and Pam3CSK4 decreased the frequencies of Th2 (IL-4+)CD4 cytokine secretion. Cells were then stained with Live/Dead Aqua (Invi- T cells in agreement with the known Th1 immunomodulatory effect trogen), fixed, permeabilized with Cytofix/Cytoperm (BD Biosciences), and then incubated with anti-CD16/CD32 Fc block (BD Biosciences). of TLR agonists (Fig. 1E). Alum and all the TLR agonists tested Spleen-derived T cells were stained with the following mAbs: anti-CD3 failed to enhance influenza-specific CD4 responses (Fig. 1F). The PerCP-Cy5.5 (BD Pharmingen), anti–CD4 V500 (BD Pharmingen), anti– data produced in these experiments in animal model reflect clinical IFN-g PE Green (BD Pharmingen), anti–IL-2 allophycocyanin (BD data showing that MF59 is a very efficient adjuvant for subunit in- Pharmingen), anti-CD8 PE Texas Red (Invitrogen), anti–IL-4 A488 fluenza vaccines, whereas alum is ineffective (21). (eBioscience), anti–IL-13 A488 (eBioscience), anti–TNF-a Alexa700 (BD Pharmingen), anti–IL-17 PE Cy7 (eBioscience), and anti–CD44 V450 (BD Differently to what was observed for flu, all adjuvants signifi- Pharmingen). LN-derived T cells were incubated with the following mAbs: cantly induced TT-specific total IgG titers (Fig. 2A). These data anti-CD3 PerCP-Cy5.5 (BD Pharmingen), anti-CD4 PE-A610 (Invi- show that all compounds tested in this study could act as adju- trogen), anti–IFN-g PE (BD Pharmingen), anti–TNF-a A700 (custom; BD vants, but their efficacy depends on the nature of the coadminis- Pharmingen), anti–IL-2 allophycocyanin (custom; BD Pharmingen), anti- CD44 Horizon V450 (BD Pharmingen), and anti–IL-4 A488 (BD Phar- tered Ag. The analysis of Ab isotypes showed that MF59 and mingen). Cells were then acquired on LRSII (BD Biosciences) and ana- Pam3CSK4 were the best inducers of IgG1 (Fig. 2B), whereas all lyzed using FlowJo software. Frequencies of influenza-specific T cells TLR agonists gave a superior IgG2a response (Fig. 2C). The Journal of Immunology 3091 Downloaded from http://www.jimmunol.org/ by guest on September 25, 2021

FIGURE 1. Humoral and cellular responses to influenza subunit vaccine formulated with different adjuvants. BALB/c mice (8 mice/group) were treated with seasonal trivalent influenza Ags (flu) in combination with MF59, alum, CpG, Pam3CSK4, or R848, or in their respective control buffer (PBS or 1% DMSO). Ab titers or HI titers were measured 2 wk post-second immunization. The data are representative of two independent experiments. (A) Total IgG titers to flu Ags B Florida, H1N1 Brisbane, and H3N2 Brisbane. (B) HI titers to H3N2 Ag: the black circles represent values from single mice, and the red bars represent the geometric mean titers over eight mice. (B) H1N1- and H3N2-specific IgG1 titers (C) and IgG2a titers (D) were measured 2 wk post- second immunization. (E and F) Analysis of CD4 T cell responses to trivalent influenza Ags in combination with the various Ags in the spleen 3 wk post- second immunization. (E) The graph shows the average frequencies of IL-4+/IL-13+ CD4 T cells from four mice/group as a percentage of total CD4 T cells. (F) Frequencies of influenza-specific CD4 T cells expressing single (IFN-g, IL-4, IL-2, IL-17, or TNF-a) or different combinations of two (double) or three (triple) cytokines. For Abs and T cell responses, unpaired two-tailed Student t test was performed comparing each group of mice treated with adjuvanted vaccines with the corresponding nonadjuvanted control group. *p # 0.05.

MF59 is the strongest activator of cytokine genes in the muscle experiments by whole genome microarray analysis. In a previous and induces early recruitment of CD11b+ cells at the injection kinetic study, we followed the profile between 3 h and site 4 d induced by several vaccine adjuvants after i.m. injection. We To identify early innate immune gene signatures that correlate with found that most of the selected early innate immunity genes were flu adjuvanticity, we analyzed the transcriptional profiles of mouse significantly regulated in the 6-h time point; therefore, we selected splenocytes (in vitro), mouse muscles, and draining LNs (in vivo) the 6-h time point for all the microarray experiments performed in after treatment with the same adjuvants tested in the previous this study (13). Significantly regulated genes were selected with an 3092 INNATE IMMUNE SIGNATURES OF VACCINE ADJUVANTS

cytokine-encoding genes modulated in the dataset. As previously described, MF59 induced a large number of genes in- volved in cell recruitment processes (13). Moreover, MF59 was the strongest inducer of Il1-related genes (Il1a, Il1b, Il1rn, Il1f9) and was the only adjuvant that upregulated Th2 cytokines (Il5 and Il4) and Il9 (Supplemental Fig. 1A). Surprisingly, the TLR7 agonist R848 activated only a very small number of cytokine genes: Cxcl9, Cxcl10, Cxcl11, Cxcl13, and Tnfsf10/Trail, all of them known to respond to IFN type I signaling (Supplemental Fig. 1A). However, differently from the muscle, R848 was the strongest modulator of gene expression in the draining LNs, modulating 34 of the 36 se- lected cytokine genes (Fig. 3A). R848 cytokine response in LNs was characterized by a strong activation of IFN type I and II genes (Ifna2, Ifna4, Ifna12, Ifnb1, and Ifng) and Th1 cytokine genes (Il12b). Pam3CSK4 induced the expression of 12 cytokine genes (mainly chemokines and IL-1 family cytokines) in the LNs, whereas all other adjuvants did not have any activity on cytokine expression or on any other pathway examined.

An unbiased analysis of the gene expression dataset highlighted Downloaded from that at 6 h MF59 also induced in the muscle genes involved in leukocyte transendothelial migration (KEGG mmu04670) and cytokine binding (Fig. 3B, Supplemental Fig. 1B). CpG and Pam3CSK4 also regulated some of these genes, like the adhesion molecule genes Itgam, Icam1, and Vcam1. These data suggest that

MF59 could lead to stronger cell recruitment from the bloodstream http://www.jimmunol.org/ to the injection site compared with the other adjuvants tested in these experiments. Among the genes encoding chemoattractants or adhesion molecules, CD11b/Itgam (marker of myeloid lineage) was selected as a biomarker to investigate local cell recruitment events 6 h after i.m. adjuvant injection by confocal immunofluo- rescence analysis of muscle cryosections (Fig. 2C). Muscle fibers were visualized by using an Ab specific for utrophin, a ubiquitous protein located at the neuromuscular synapse and myotendinous junctions that plays a role in anchoring the cytoskeleton to the by guest on September 25, 2021 plasma membrane in muscle cells. Nuclei were visualized by ToPro 3, a cyanine stain that allows DNA detection. At 6 h MF59 and, to a lower level, Pam3CSK4 could draw afflux of CD11b+ cells at the injection site. The muscle infiltrate was con- firmed and further characterized by flow cytometry of muscle single-cell suspensions at 24 h after injection. MF59 induced a strong recruitment of neutrophils, eosinophils, and monocytes, following treatment, whereas Pam3CSK4 induced mainly neutro- phil influx (Fig. 3D). The observed recruitment of CD11b+ cells at the injection site correlates very well with the adjuvanticity of the FIGURE 2. Ab responses to TT vaccine formulated with different ad- different adjuvants to flu Ags (Fig. 1). Indeed, correlation analysis juvant. BALB/c mice (8 mice/group) were treated with TT Ag in combi- between the gene expression profiles, induced by the different nation with MF59, alum, CpG, Pam3CSK4, or R848, or in their respective control buffer (PBS or 1% DMSO). Total Ab titers (A), IgG1 (B), and adjuvants and the flu IgG Ab titers elicited in the adjuvanticity IgG2a (C) were measured 2 wk post-second immunization. Unpaired two- experiments, highlighted P (Selp), a cell adhesion molecule tailed Student t test was performed comparing mice treated with adju- that mediates the interaction of activated endothelial cells or pla- vanted vaccines versus nonadjuvanted control group. *p # 0.05. telets with leukocytes by mediating rapid rolling of leukocytes over vascular surfaces during the initial steps in inflammation, as average log2 ratio $2 compared with control groups and a p value one of the few strongly positive correlating genes (Supplemental #0.05 calculated on the three replicates per treatment group. Func- Table I). Selp, one of the genes belonging to the leukocyte trans- tional analysis of the adjuvant-regulated genes showed that in vitro endothelial migration cluster, was strongly modulated by MF59 in mouse splenocytes only the three TLR agonists were able to and Pam3CSK4 in the muscle (Fig. 3B), further supporting the activate transcription of cytokine-encoding genes (Fig. 3A, Sup- idea that enhanced cell recruitment at the injection might result in plemental Fig. 1A). As expected, all TLR agonists upregulated improved adjuvanticity to subunit influenza vaccine. proinflammatory (Tnf, Il6) and Th1 cytokine (Il12 and Ifng) genes To dissect the contribution of either adjuvant or flu Ags to (Supplemental Fig. 1A). On the contrary, the TLR-independent changes to the muscle or splenocyte transcriptomes, we evaluated adjuvants MF59 and alum did not modulate the expression of cy- in a separate experiment the expression profiles of the three flu Ags tokine genes or any other pathway (Supplemental Fig. 1A, data not alone, flu Ags in combination with the adjuvant (MF59 + flu), and shown), suggesting that MF59 and alum poorly activate immune adjuvant alone (MF59). We found that the flu Ags did not impact cells in vitro. In contrast, in vivo, MF59 was the strongest modulator the basal transcriptional program of splenocytes in vitro (data not of transcriptional events at the injection site, activating 36 of the 43 shown) or of the mouse muscle (Fig. 4). Moreover, the combi- The Journal of Immunology 3093 Downloaded from http://www.jimmunol.org/

FIGURE 3. MF59 is the strongest modulator of cytokine genes in the muscle and induces early recruitment of CD11b+ cells. (A) Number of significantly regulated cytokine genes in splenocytes (in vitro), skeletal muscle, and inguinal LNs 6 h after treatment with the adjuvants tested in this study. (B) Ex- pression profile of significantly expressed genes belonging to the leukocyte transendothelial migration pathway (KEGG mmu04670) in the muscle 6 h after treatment with the various adjuvants. Some genes, like Icam1, appear more than once in the cluster because they are detected by multiple unrelated probes in the Agilent 44k Whole-Mouse Genome Array. Genes were selected for fold change $|4| and p # 0.05 in at least one condition (see Materials and by guest on September 25, 2021 Methods). The data are shown in log2 scale ranging from 23, green, downregulated to +3, red, upregulated. (C) Confocal analysis of muscle cryosections taken 6 h after treatment showing utrophin in green, CD11b+ cells in purple, and a nuclei in blue. Scale bar, 40 mm. Results are representative of three independent experiments. (D) FACS analysis of cell composition in the muscle 24 h after treatment with various adjuvants. Myeloid lineage cell (CD11b+) neutrophils were defined as Ly6G+, monocytes as Ly6Chigh, eosinophils as Ly6Glow/side scatterhigh, macrophages as F4/80+, and DCs as CD11c+/MHC class IIhigh. The events are shown as average counts over three mice with the relative SD. nation of Ag with the adjuvant did not alter the transcriptional (CD11c+/CD11b+) expressing CD86 and high levels of MHC class II profile of the adjuvant alone. molecules (Fig. 5G). In summary, R848 was able to strongly induce the IFN pathway and cytokine responses in vitro, to activate the IFN R848 strongly modulates IFN-related genes in the muscle and pathway at the injection site, and to strongly activate IFN and cy- draining LNs tokine pathways in LNs, leading to T, NK, and B cell activation and Analysis of genes encoding for components of the IFN signaling to DC recruitment. Interestingly, even if the broad immunostimu- pathway or for IFN-responsive genes revealed that both R848 latory activity of R848 correlated with TT adjuvanticity, it was not and CpG strongly induced these genes either in vitro in mouse sufficient to promote flu adjuvanticity. By contrast, MF59, which is splenocytes or in vivo in muscle (Supplemental Fig. 2A), in agree- the best adjuvant for a flu subunit vaccine among the adjuvants tested ment with the available data on TLR7 and TLR9 signaling pathways in this work, was unable to activate either splenocytes or LNs in vivo, (24, 25). Interestingly, in the draining LNs, R848 was the only whereas it worked efficiently locally at the injection site. regulator of IFN-related genes (61 of the 62 significantly modulated To validate the microarray data, we have analyzed the expression genes in this cluster), whereas none of the other adjuvants tested profile of five genes (Cxcl10, Il6, Ccl2, Tnfa, and Mx1) in muscle had an impact on transcription of these genes (Fig. 5A). In addition injected with PBS, MF59, alum, and CpG by quantitative RT- to cytokine and IFN pathways, R848 was also a selective inducer of PCR. The data obtained with microarray and RT-PCR using the cell activation marker genes such as CD69, CD86, and CD40 (Fig. same RNA samples were very similar for all the genes tested 5B). These data together with the cytokine data shown in Fig. 2 and (Supplemental Fig. 2B). Supplemental Fig. 1A suggest that R848 could induce a broad ac- tivation of the immune cells in the LNs. CD69 and CD86 were used MF59 and Pam3CSK4 adjuvanticity to flu and TT is IFN type I as biomarkers in multicolor flow cytometric stainings of LN cells independent 24 h after i.m. injection of the different adjuvants in the absence of Type I IFN was shown to be a powerful adjuvant for a subunit flu coadministered Ag. In agreement with the microarray data, R848 vaccine formulation (6). Similarly to MF59, rIFN type I coad- was the only compound able to induce an increased expression of ministered to a subunit flu vaccine increased HI titers and pro- CD69 on T, B, and NK cells (Fig. 5C–E, respectively) and of CD86 tection against intranasal influenza challenge in the mouse model. in B cells (Fig. 5F). Moreover, R848 increased the number of DCs The effect was shown to depend on IFN type I receptor (IFN-RI), 3094 INNATE IMMUNE SIGNATURES OF VACCINE ADJUVANTS

adjuvanted vaccine was not affected by anti-IFN type I receptor Ab treatment even after 2 wk from the second dose (Fig. 6E–G). Similar data were obtained for IgG1 and IgG2a isotypes (data not shown). Surprisingly, the Ab response to H3N2 and B Ags adjuvanted with Pam3CSK4 was enhanced by IFNAR2 Ab, suggesting that IFN type I may have an inhibitory effect on the adjuvanticity of TLR2 agonists after the booster dose. In parallel to the Ab response, we assessed the Ag-specific T cell response in cells derived from the draining LNs by intracellular cytokine staining 3 wk after the second immunization. The presence of the anti-IFNAR2 Abs did not reduce the total number of influenza-specific CD4 T cells induced by MF59 or Pam3CSK4 (Fig. 6H). In the group treated with anti-IFNAR2 Ab immunized with flu plus Pam3CSK4, we detected an increase of IL-4+ CD4 T cells and a decrease of IFN-g+ CD4 T cells, suggesting that IFN type I is involved in modulating Th1/Th2 balance in re- sponse to TLR2 agonist.

Discussion

In the current work, we compared MF59 with alum and with the Downloaded from TLR-dependent adjuvants, CpG, R848, and Pam3CSK4 for their ability to boost adaptive responses to flu subunit vaccine and to TT. Although all the adjuvants were able to induce good Ab titers to TT, only MF59 and, to a lower extent, Pam3CSK4 significantly in- creased the humoral response to flu Ags. We cannot explain why HA

and TT have different requirements for adjuvanticity based on the http://www.jimmunol.org/ existing data. One possibility is that HA and TT interact differen- tially with thevarious adjuvants used in this study. Indeed, it has been shown that Ag-adjuvant codelivery can have a significant impact on adaptive immune responses (26). In addition, the ability of HA to bind sialic acid, a saccharide present in most human cell types, may affect its biodistribution after i.m. injection and its ability to reach FIGURE 4. Flu Ags do not alter the expression profile induced by MF59 the draining LNs. This may favor the adjuvants that exert a strong in the muscle. Expression profiles of mouse muscle 6 h after treatment with local response, such as MF59 and Pam3CSK4, and not the adjuvants the three flu Ags in PBS (flu), flu Ags in combination with MF59 (MF59 + that activate innate immunity directly in the LNs, such as R848. by guest on September 25, 2021 flu), and adjuvant alone (MF59). (A) Overview of significantly regulated The analysis of Ab subclasses in response to TT and influenza B genes; ( ) subset of genes encoding for cytokines hierarchically clustered, showed that TLR agonists induced a Th1-biased response. MF59 2 as described in Supplemental Fig. 1. The log2 scale ranges from 6, green, induced a mixed response, even though the IgG1 to IgG2 ratio was downregulated genes to +6, red, upregulated genes. shifted toward IgG1, confirming that MF59 is an inducer of Th2- biased responses in BALB/c (22, 27), a mouse strain genetically as the adjuvant effect of IFN type I on influenza subunit vaccine was skewed toward Th2 responses (28). abolished in IFN-RI knockout (KO) animals (7). Our microarray In agreement with Ab responses to influenza Ags, we found that analysis highlighted that the tested adjuvants induced differentially MF59 was a strong inducer of IL-4+/IL-13+ CD4 T cells known to the IFN-related genes. CpG and R848 were the most potent activators exert B helper function for production of IgG1 Abs. Despite the of IFN-dependent genes, whereas very few IFN genes were activated ability of Pam3CSK4 to enhance the Ab response to influenza by MF59, alum, or Pam3CSK4 (Supplemental Fig. 2). These data vaccine, we could not measure a parallel increase of flu-specific would suggest that the adjuvanticity of MF59, Pam3CSK4, and alum CD4 T cells. We suggest that Pam3CSK4 may have a modest is largely IFN independent, whereas the adjuvanticity of R848 and effect on CD4 T cells that we cannot measure with the sensitivity CpG depends on IFN. To verify whether a functional type I IFN of our current T cell assay. pathway is required for the mechanism of action of these adjuvants, Interestingly, the data obtained in our mouse model are con- mice treated with anti-IFN type I receptor Ab (anti-IFNAR2) and sistent with clinical data showing that MF59 can boost the re- then immunized with Ag plus adjuvant were compared with immu- sponses to flu vaccines better than alum in humans (21). With the nized mice treated with the control isotype Ab. All the adjuvants exception of alum, we have investigated the effect of the various were tested in combination with TT (Fig. 6A), whereas only MF59 adjuvants by simply mixing them with flu and TT. Therefore, we and Pam3CSK4, the two adjuvants able to enhance flu immunoge- cannot exclude that R848 and CpG would boost flu subunit vac- nicity, were tested in combination with flu Ags (Fig. 6B–H). Two cine responses if physically linked to the HA-Ag through direct weeks post-second immunization, sera were analyzed for Ag-specific conjugation or through a delivery system. Indeed, several pub- Ab titers. Mice treated with anti-IFNAR2, MF59, and Pam3CSK4 lications have shown that direct conjugation of TLR2, 7, and 9 were still able to induce strong TT-specific total IgG Abs; alum and agonists to the Ag increases their adjuvanticity (26, 29, 30). CpG adjuvanticity was partially inhibited, whereas Ab titers dropped In a first attempt to determine whether differential adaptive re- considerably when R848 was used as adjuvant (Fig. 6A). Mice sponses to flu correlated to distinct innate immune reactions in- treated with anti-IFNAR2 and immunized with trivalent flu vaccine duced by the different adjuvants, we analyzed the transcriptional together with MF59 or Pam3CSK4 produced total IgG titers to the profiles induced by each adjuvant in mouse splenocytes. three flu Ags similarly to isotype Ab-treated control mice after pri- Consistently with previous observations, TLR-dependent adju- mary immunization (Fig. 6B–D). Moreover, the response to MF59- vants induced strong transcriptional changes (31). Indeed, TLR The Journal of Immunology 3095 Downloaded from http://www.jimmunol.org/ by guest on September 25, 2021

FIGURE 5. R848 is the strongest modulator of IFN pathway in muscle and LNs and induces LN cell activation. (A) Number of significantly regulated IFN-related genes in splenocytes, skeletal muscle, and inguinal LNs 6 h after treatment with various adjuvants. (B) Microarray expression of selected cell surface-expressed genes in the draining LNs 6 h after treatment with the various adjuvants. (C–E) Multicolor flow cytometric analysis of CD69 on T cells (C), B cells (D), and NK cells (E) isolated from inguinal LNs 1 d after i.m. injection with the different adjuvants. (F) Multicolor flow cytometric analysis of CD86 on B cells isolated from the draining LNs 1 d after i.m. injection with the different adjuvants. (C–F) Data are shown as mean fluorescence intensity (MFI) of either CD69 or CD86 on the relative subsets. (G) Multicolor flow cytometric analysis of MHC class II expression on DCs isolated from the draining LNs 1 d after i.m. injection with the different adjuvants. The data represent the number of DCs expressing CD86 and high level of MHC class II molecules after each treatment. SD bars were calculated over at least three independent experiments. agonists rare known to directly activate splenocytes by binding to infiltration of CD11b+ cells into the muscle 6 h after injection of TLRs. By contrast, MF59 and alum were not able to modulate blood MF59. CD11b+ cell recruitment at 6 h was observed also after cell transcription in vitro, even though it has been shown that MF59 Pam3CSK4 injection. Further characterization of muscle single- and alum induced secretion of chemokines in monocytes (14), and cell suspensions by flow cytometry of the muscle 1 d after injec- that alum activated the NOD-like receptor protein 3 inflammasome tion revealed that the cell populations recruited by MF59 and complex (32). We hypothesize that the activation of chemokine Pam3CSK4 differed greatly. MF59 induced significant recruitment secretion by blood cells in vitro by alum and MF59 is driven by post- of granulocytes, monocytes, and DCs, whereas Pam3CSK4 could transcriptional events. More work needs to be done to investigate only induce neutrophil infiltration, which can explain some of the whether alum and MF59 stimulate mRNA translation or secretion of differences in the quality of the immune responses elicited by these preformed chemokines. Next, we analyzed the transcriptional pro- two adjuvants. Overall, these results are in agreement with results files induced by each vaccine adjuvant in the muscle and in draining published by Mosca et al. (13) and by Calabro et al. (15) on the LNs. Even if MF59 did not activate innate immunity in vitro, it was effect of MF59 on muscle transcriptome and cell recruitment. The the strongest inducer of transcriptional changes at the injection site. striking correlation between the early cell recruitment events in the In particular, MF59 was the most potent inducer of genes coding for muscle and the ability of the vaccine adjuvants to boost adaptive chemokines, chemokine receptors, and transendothelial migration response to HA suggests that leukocyte migration to the injection factors. In agreement with microarray data, we detected a significant site promotes adjuvanticity to flu subunit vaccines. Of course, these 3096 INNATE IMMUNE SIGNATURES OF VACCINE ADJUVANTS Downloaded from http://www.jimmunol.org/ by guest on September 25, 2021

FIGURE 6. MF59 and Pam3CSK4 adjuvanticity to flu and TT is IFN independent. BALB/c mice (6 mice/group) were injected i.p. with anti–IFN-ab receptor 2 Ab (anti-IFNAR2) or control (iso), 1 d before and 7 d after the first immunization and immunized with TT or seasonal trivalent flu Ags either alone or in combination with various adjuvants. (A) Two weeks post-second immunization, sera were analyzed for total IgG titers to TT. Two weeks post- first immunization, sera were analyzed for total IgG titers for H1N1 (B), H3N2 (C), and B Florida (D). Two weeks post-second immunization, IgG titer sera were analyzed for total IgG titers to H1N1 (E), H3N2 (F), and B Florida (G). The values are shown as mean titer over six mice. Unpaired two-tailed Student t test was performed comparing mice treated with IFNAR2 versus mice treated with control Ab, *p # 0.05, **p # 0.01. (H) CD4+ T cell responses to trivalent influenza Ags were measured 3 wk post-second immunization from a pool of four draining LN/group. The histograms show the percentage of CD44+CD4 T cells positive for any cytokine. -, Unstimulated T cells; 3 mg, T cells stimulated with 0.3 mg/ml flu Ags. The percentage of CD4 T cells positive for each cytokines following stimulation with flu Ags is shown in the graph below each group. events are also linked to local reactogenicity of the vaccine. Inter- the muscle. Probably the blood cell types present in the muscle estingly, R848 was able to strongly activate both IFN and cytokine respond differently to TLR7 activation compared with mouse blood pathways in splenocytes, but induced only IFN-responsive genes in cells in vitro. The Journal of Immunology 3097

The substantial differences between splenocytes and muscle Acknowledgments transcriptional profiles in response to MF59, alum, and R848 show We thank M. Tortoli and the Animal Care Facility at Novartis Vaccines that in vitro assays are not predictive of adjuvant activity in vivo. Siena for all mice experiments; C. Malzone for help on the analysis of In contrast to the injection site, MF59 had no impact on the IgG titers; D. Casini for support with the ELISA experiments; and S. Nuti, transcriptional activity of the draining LNs, whereas the TLR7 S. Tavarini, and C. Sammicheli for help on the FACS experiments. agonist R848 was the strongest modulator of LN transcription. Both the cytokine/chemokine and the IFN-related gene clusters were Disclosures strongly induced by R848 in the draining LNs. Moreover, flow All authors, with the exception of E.T., are employees of Novartis Vaccines cytometric analysis of LN single-cell suspensions showed that and Diagnostics. E.T. is employed by Novartis Institute of Biomedical Re- R848 could activate T, B, and DCs. Because the adjuvants were search, in the same Novartis group. administered without Ags, the observed activation of both APCs and adaptive immune cells in LNs by R848 is Ag independent, References and may not increase specific adaptive responses when R848 is 1. Fiore, A. E., D. K. Shay, K. Broder, J. K. Iskander, T. M. Uyeki, G. Mootrey, coadministered with HA. In addition, R848 may even trigger the J. S. Bresee, and N. S. Cox; Centers for Disease Control and Prevention (CDC); activation of autoreactive B and T cell clones. In a recently pub- Advisory Committee on Immunization Practices (ACIP). 2008. Prevention and control of influenza: recommendations of the Advisory Committee on Immu- lished article, Koyama et al. (5) demonstrated that WV influenza nization Practices (ACIP), 2008. MMWR Recomm. Rep. 57(RR-7): 1–60. vaccines require TLR7-mediated IFN type I response for immu- 2. Gross, P. A., F. A. Ennis, P. F. Gaerlan, L. J. Denson, C. R. Denning, and nogenicity. Moreover, rIFN-a, when administered at the time of D. Schiffman. 1977. A controlled double-blind comparison of reactogenicity, immunogenicity, and protective efficacy of whole-virus and split-product influ- vaccination, was shown to act as a powerful adjuvant for a flu enza vaccines in children. J. Infect. Dis. 136: 623–632. Downloaded from subunit vaccine inducing protection against influenza virus chal- 3. Gross, P. A., and F. A. Ennis. 1977. Influenza vaccine: split-product versus lenge (7). Microarray and adjuvanticity data observed in our ex- whole-virus types: how do they differ. N. Engl. J. Med. 296: 567–568. 4. Rhorer, J., C. S. Ambrose, S. Dickinson, H. Hamilton, N. A. Oleka, F. J. Malinoski, perimental settings would, however, suggest that IFN type I and J. Wittes. 2009. Efficacy of live attenuated influenza vaccine in children: activation can be dispensable in making a good adjuvant to flu a meta-analysis of nine randomized clinical trials. Vaccine 27: 1101–1110. 5. Koyama, S., T.Aoshi, T.Tanimoto,Y.Kumagai,K.Kobiyama, T.Tougan, K.Sakurai, vaccine. Indeed, MF59 and Pam3CSK4 are not good inducers of C. Coban, T. Horii, S. Akira, and K. J. Ishii. 2010. Plasmacytoid dendritic cells de-

IFN-related genes, but are efficient adjuvants for the influenza lineate immunogenicity of influenza vaccine subtypes. Sci. Transl. Med. 2: 25ra24. http://www.jimmunol.org/ vaccine. In addition, despite the ability of CpG to induce a strong 6. Co, M. D., L. Orphin, J. Cruz, P. Pazoles, K. M. Green, J. Potts, A. M. Leporati, J. A. Babon, J. E. Evans, F. A. Ennis, and M. Terajima. 2009. In vitro evidence IFN response in the muscle and of R848 to trigger the IFN that commercial influenza vaccines are not similar in their ability to activate responses in muscle and LNs, none of these adjuvants were able to human T cell responses. Vaccine 27: 319–327. boost adaptive responses to subunit flu vaccination. Analysis of 7. Proietti, E., L. Bracci, S. Puzelli, T. Di Pucchio, P. Sestili, E. De Vincenzi, M. Venditti, I. Capone, I. Seif, E. De Maeyer, et al. 2002. Type I IFN as a natural adaptive immune responses in mice in which the IFN type I sig- adjuvant for a protective immune response: lessons from the influenza vaccine naling was inhibited by treatment with an IFN type I receptor- model. J. Immunol. 169: 375–383. blocking Ab further confirmed this hypothesis. The adjuvanticity 8. O’Hagan, D. T., A. Wack, and A. Podda. 2007. MF59 is a safe and potent vaccine adjuvant for flu vaccines in humans: what did we learn during its de- of R848 to TT was almost completely abrogated in the absence of velopment? Clin. Pharmacol. Ther. 82: 740–744. IFN type I signaling. These data are consistent with a recent study 9. Podda, A. 2001. The adjuvanted influenza vaccines with novel adjuvants: ex- by guest on September 25, 2021 perience with the MF59-adjuvanted vaccine. Vaccine 19: 2673–2680. showing that IFN type I is required for R848 adjuvanticity to OVA 10. Dupuis, M., T. J. Murphy, D. Higgins, M. Ugozzoli, G. van Nest, G. Ott, and (33). 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Our data on Ab titers and cellular responses R. Rappuoli, and E. De Gregorio. 2008. Molecular and cellular signatures of after the influenza vaccine boost confirmed that MF59 is inde- human vaccine adjuvants. Proc. Natl. Acad. Sci. USA 105: 10501–10506. 14. Seubert, A., E. Monaci, M. Pizza, D. T. O’Hagan, and A. Wack. 2008. The pendent of IFN type I, and they suggest that IFN type I may even adjuvants aluminum hydroxide and MF59 induce monocyte and granulocyte play a regulatory role in response to Pam3CSK4 by modulating chemoattractants and enhance monocyte differentiation toward dendritic cells. J. Immunol. 180: 5402–5412. the Th1/Th2 balance. 15. Calabro, S., M. Tortoli, B. C. Baudner, A. Pacitto, M. Cortese, D. T. O’Hagan, By contrast to our data, a previous study has shown that Ab E. De Gregorio, A. Seubert, and A. Wack. 2011. Vaccine adjuvants alum and responses to MF59-adjuvanted flu are partially affected in an IFN- MF59 induce rapid recruitment of neutrophils and monocytes that participate in antigen transport to draining lymph nodes. Vaccine 29: 1812–1823. RI KO background (7). These differences may be explained by the 16. Vesikari, T., M. Pellegrini, A. Karvonen, N. Groth, A. Borkowski, D. T. O’Hagan, fact that immune cells that develop in the absence of IFN-RI may and A. Podda. 2009. Enhanced immunogenicity of seasonal influenza vaccines in have an intrinsic defect to respond to vaccination that is not de- young children using MF59 adjuvant. Pediatr. Infect. Dis. J. 28: 563–571. 17. Galli, G., D. Medini, E. Borgogni, L. Zedda, M. Bardelli, C. Malzone, S. Nuti, pendent on the adjuvant used. Indeed, in the previous study by S. Tavarini, C. Sammicheli, A. K. Hilbert, et al. 2009. Adjuvanted H5N1 vaccine Proietti et al. 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Galli, G., K. Hancock, K. Hoschler, J. DeVos, M. Praus, M. Bardelli, C. Malzone, F. Castellino, C. Gentile, T. McNally, et al. 2009. Fast rise of juvant Pam3CSK4 strongly correlates with the ability of these broadly cross-reactive antibodies after boosting long-lived human memory + adjuvants to enhance Ab responses to flu Ags. Within this CD11b B cells primed by an MF59 adjuvanted prepandemic vaccine. Proc. Natl. Acad. population, a higher frequency of professional APCs recruited by Sci. USA 106: 7962–7967. 20. Clark, T. W., M. Pareek, K. Hoschler, H. Dillon, K. G. Nicholson, N. Groth, and MF59 compared with Pam3CSK4 correlated to an even more en- I. Stephenson. 2009. Trial of 2009 influenza A (H1N1) monovalent MF59- hanced immune response to flu vaccine. adjuvanted vaccine. N. Engl. J. Med. 361: 2424–2435. 3098 INNATE IMMUNE SIGNATURES OF VACCINE ADJUVANTS

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Supplementary figure 2: A. Cluster analysis of IFN-related genes. Genes encoding IFN- related genes (manually curated using several databases) were selected and clustered as described in Suppl. Fig 1. The log2 scale ranges from -3, green, down-regulated genes to +3, red, up-regulated genes. B. RT-qPCR validation of microarray data. The expression profiles of 5 genes (Cxcl10, IL6, Ccl2, TNFa and Mx1) in the muscle in response to MF59, alum or CpG treatments were measured by RT-qPCR (left panels) or microarray (right panels) from the sample RNA samples. The data is displayed as fold-change relative to the expression measured in PBS-injected muscles.

Supplementary table I: Correlation between gene expression profiles and influenza vaccine antibody titers. The table shows the log2 fold change induced by each adjuvant, the R2 correlation coefficient and related p-value calculated for the correlation based on Student's t-statistics for those genes having a p-value < 0.01 for all three flu vaccine antigens.

Supplementary table II: Significantly regulated genes in the muscle. The table shows gene expression values (the average log2 fold changes over at least 3 animals, p-value and q-value) for the 1410 selected genes together with the correlation data (R2 correlation coefficient and Student's t-test p-value) between gene expression profiles and influenza vaccine antibody titers for all the genes.

Supplementary Table I

Log2(mean expression value) correlation (R2) correlation p-values Agilent Gene Reporter Id Accession Symbol Gene Name MF59 Alum CpG R848 Pam3 B H1N1 H3N2 B H1N1 H3N2 A_52_P57843 NM_011347 Selp selectin, platelet 2.73 0.73 0.62 -0.21 2.16 0.835 0.873 0.834 0.0054 0.0032 0.0055 Rho family GTPase A_52_P383137 BC048531 Rnd1 1 2.18 0.89 0.78 -0.02 1.83 0.864 0.881 0.841 0.0036 0.0028 0.0050 RAB11 family interacting protein 1 A_52_P68033 NM_001080813 Rab11fip1 (class I) 2.97 -0.04 1.00 1.00 2.02 0.930 0.842 0.896 0.0009 0.0050 0.0021 RAB11 family interacting protein 1 A_52_P595668 NM_001080813 Rab11fip1 (class I) 2.33 1.00 1.00 1.00 1.75 0.872 0.795 0.858 0.0032 0.0085 0.0040 A_52_P665240 NM_001003668 Krt83 83 4.21 1.00 1.00 1.00 1.00 0.921 0.843 0.887 0.0012 0.0049 0.0025 D630042 RIKEN cDNA A_52_P62489 AK054244 P16Rik D630042P16 gene -2.39 0.08 -0.13 1.00 -1.14 0.908 0.844 0.876 0.0016 0.0048 0.0030 A_51_P376590 BB161836 4.02 1.00 1.00 1.00 1.00 0.851 0.856 0.822 0.0044 0.0041 0.0064 killer cell lectin-like receptor, subfamily A_52_P34058 NM_010648 Klra3 A, member 3 4.48 1.00 1.00 1.00 1.00 0.857 0.833 0.820 0.0040 0.0055 0.0065 % 96$#%#0!%)7 (&%(0( $# $4#%" BD "1# & CAC #@ A GE(AFCBEG2EhT!Y)6A16A40(FF2GG2TFH)Y3(@69R! `rh`hggpit a$X!b !(Ch hWqr hWpg iWqv iWqs hWpv `rh`hgguri 3(@69RP6G5F2DH2A02F6@69(E6GRisT@2@)2E0 0(@is3 hWrr gWqt iWsq hWqi hWht F2E6A2aBE0RFG26A2bC2CG61(F26A56)6GBET09(12!T `rh`hghhqs @2@)2Ei 2EC6A)i pWtg gWih gWtv gWgg hWii `rh`hghuru 16F0FT9(E425B@B9B4ha$EBFBC569(b $94h YhWhr YiWgr gWvq gWqi gWpt `rh`hghviv 920G6AT4(9(0GBF2)6A16A4TFB9H)92p 4(9Fp iWiq gWqg YgWhv YgWgu YgWqp `rh`hgituv 0B@C92@2AG0B@CBA2AGhTDFH)0B@CBA2AGT305(6A 3hD0 YiWvs gWhq YgWgh gWpi gWgs `rh`hgphpp gWgg rWti gWgg gWgg gWgg `rh`hgpiii FB9HG20(EE62E3(@69RpvaS6A0GE(AFCBEG2EbT@2@)2Eq 90pv(q iWqr gWrt gWpt YgWvi gWii `rh`hgrhtu &Y)BQ0BAG(6A6A442A2q BQq YgWsp gWsv YgWgt pWpv YgWhh `rh`hgrpug 90$ighgggr;hr42A2 ighgggr;hr68 qWup qWvg gWgg gWgg gWgg `rh`hgruut 9R@C5B0RG2GE(AF@2@)E(A2(1(CGBEh (Qh pWvv gWgg gWgg gWgg gWgg `rh`hgtpgi 0(I2B96AhT0(I2B9(2CEBG26A 3(Ih YiWgi YiWit gWgs YgWsu YgWhs `rh`hguhgu 92H06A2E605E2C2(G0BAG(6A6A4ir EE0ir hWru gWrv hWru gWru iWgs `rh`hgvqqv 92H8BGE62A2!qE202CGBEh G)qEh iWri hWii YgWrr YhWsh YgWsq `rh`hgvrgu 52C(G6G6FI6EHF0299H9(EE202CGBEi ;(I0Ei pWpr gWgu YhWgu YgWvu YgWpu `rh`hhhhsq 5B3(@69R>!(F2h A1h pWuu hWtq hWtr gWht pWgt `rh`hhhqvi AH092(E3(0GBET6AG2E92H86ApTE24H9(G21 369p gWut iWph gWpu iWpg hWhq `rh`hhiqgr C9(F@6AB42A(0G6I(GBETHEB86A(F2E202CGBE 9(HE qWgt hWpu gWug gWqp gWsq FB9HG20(EE62E3(@69Rta0(G6BA60(@6AB(061GE(AFCBEG2ET `rh`hhitpq RwFRFG2@bT@2@)2Eu 90t(u YhWgt YhWhq YgWis pWgu YgWgr `rh`hhivss 05B92FG2EB9irY5R1EBQR9(F2 35ir5 qWtt hWuv hWps gWph hWpu `rh`hhqsgs 3Y(FFB06(G21CEBG26Ah 08(Ch YhWqh YiWgr gWth gWpt gWps `rh`hhqshs )(F6092H06A2S6CC2EGE(AF0E6CG6BA3(0GBET!0Y9682 !(G3 iWgp gWpi iWgh YgWpt iWph `rh`hhrppr GE6C(EG6G2@BG63Y0BAG(6A6A4sp !E6@sp YgWgp gWri gWit iWtp gWiq `rh`hhsggt 56FGBA212(02GR9(F2i ;1(0i YhWtt YiWsq gWiv gWqg gWqi `rh`hhthsr sppgqgv$ig68 qWsi qWqt gWgg gWgg gWgg `rh`hhtvri 52(GF5B08CEBG26Ah ;FC(h( gWug iWvr gWvt YhWhs YhWit `rh`hhuspt S6A036A42E333;GRC20BAG(6A6A4hp '0p5hp YhWis YiWgt YgWgv YgWhu gWpi CEBG26AC5BFC5(G(F2ia3BE@2E9RibTE24H9(GBER `rh`higgvt FH)HA6GasrbT)2G(6FB3BE@ CCiEh) gWgg qWts pWvh rWhh qWsr `rh`higiig F64A(9Y6A1H021CEB9632E(G6BAY(FFB06(G21h9682p 6C(h9p gWgs YgWuv gWrg YiWit YgWpp `rh`higupg @(GE6Q@2G(99BC2CG61(F2hg @Chg gWgg rWgi gWgg gWgg gWgg `rh`hihuvh  YE29(G213p)BGH96AH@FH)FGE(G2i (0i iWsp hWhq hWir YgWut hWgv

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