MINI REVIEW article

Front. Immunol., 29 August 2014

Sec. Microbial Immunology

Volume 5 - 2014 | https://doi.org/10.3389/fimmu.2014.00412

Unraveling the Convoluted Biological Roles of Type I Interferons in Infection and Immunity: A Way Forward for Therapeutics and Vaccine Design

  • DK

    Danushka Kumara Wijesundara 1,2† *

  • YX

    Yang Xi 3

  • CR

    Charani Ranasinghe 2

  • 1. Virology Laboratory, Department of Surgery, Basil Hetzel Institute, University of Adelaide, Adelaide, SA, Australia

  • 2. Molecular Mucosal Vaccine Immunology Group, The John Curtin School of Medical Research, The Australian National University, Canberra, ACT, Australia

  • 3. Lung and Allergy Research Centre, Translational Research Institute, UQ School of Medicine, The University of Queensland, Woolloongabba, QLD, Australia

Abstract

It has been well-established that type I interferons (IFN-Is) have pleiotropic effects and play an early central role in the control of many acute viral infections. However, their pleiotropic effects are not always beneficial to the host and in fact several reports suggest that the induction of IFN-Is exacerbate disease outcomes against some bacterial and chronic viral infections. In this brief review, we probe into this mystery and try to develop answers based on past and recent studies evaluating the roles of IFN-Is in infection and immunity as this is vital for developing effective IFN-Is based therapeutics and vaccines. We also discuss the biological roles of an emerging IFN-I, namely IFN-ε, and discuss its potential use as a mucosal therapeutic and/or vaccine adjuvant. Overall, we anticipate the discussions generated in this review will provide new insights for better exploiting the biological functions of IFN-Is in developing efficacious therapeutics and vaccines in the future.

Introduction

Since the initial discovery of type I interferons (IFN-Is) as anti-viral agents (1), these cytokines have been extensively studied for their anti-microbial and immune regulatory properties. IFN-I family comprises 13 IFN-α subunits, IFN-β, IFN-ω, IFN-ε, IFN-κ, IFN-τ, and IFN-δ (in mice only) (28). All IFN-Is signal through the IFN-α receptor (IFN-AR) complex to induce synthesis and secretion of IFN-inducible genes or effector proteins with anti-viral, pro-apoptotic, and ubiquitination-modifying properties (911). The signaling pathways that IFN-Is utilize to exert various biological effects have been comprehensively reviewed elsewhere and will not be reviewed here [see Ref. (12)]. Numerous cell types produce IFN-Is (e.g., macrophages, myeloid dendritic cells (DCs), fibroblasts, and epithelial cells), but plasmacytoid DCs (pDCs) appear to be the most prolific producers of IFN-Is (13, 14). The production of these cytokines tends to be beneficial to the host particularly against acute viral infections, but there are considerable evidences to suggest that IFN-Is play detrimental roles in autoimmune diseases (15), bacterial and persistent viral infections. Herein, we review how IFN-Is could play beneficial or detrimental roles in pathogen control predominantly with respect to viral infections and discuss how they could be used as therapeutics and vaccine adjuvants. Furthermore, the importance of considering the emerging IFN-ε in immunity and vaccine development will be discussed.

The Benefits and Detriments of IFN-Is in the Control of Pathogens

The importance of IFN-Is in protecting hosts against pathogens has been demonstrated in several contexts. Firstly, IFN-AR deficient mice tend to be more susceptible to infection with viruses (particularly acute viral infections) compared to wild-type mice. Some examples include Henipavirus (16), acute Friend virus (17), encephalitic flavivirus (18), lymphocytic choriomeningitis virus (LCMV) Armstrong (19), Hazara virus (20), Dengue virus (21), Respiratory Syncytial Virus (22), and numerous other viral infections (23). Secondly, systemic exhaustion of IFN-Is following a primary viral infection has been shown to increase the host susceptibility to secondary unrelated viral infections in mice (24). Thirdly, therapeutic administration of IFN-Is can reduce viral loads in individuals infected with chronic viruses and promote cancer regression (see below Section “The Use of IFN-Is as Therapeutics and Adjuvants”). Finally, pathogens can attenuate IFN-I responses to promote immune evasion. For instance, human immunodeficiency virus (HIV)-1 can reduce the capacity of IFN producing cells to produce IFN-Is (2527), induce cytopathic effects on these cells (2832), and/or block IFN-I mediated intracellular signaling events (33) to help establish a chronic phase infection. Similarly, cancer immune evasion and development could also involve attenuation of IFN-I responses. In agreement with this, Critchley-Thorne et al. (34) have shown that various cancer patients have significantly attenuated expression of interferon stimulate genes in lymphocytes compared to healthy controls.

The benefits of IFN-Is in conferring protection against microbes have been mostly demonstrated using acute viral infection models, but several studies suggest that IFN-Is can also assist in the control of bacterial infections. This was first demonstrated in vitro where De la Maza and colleagues (35) showed that IFN-I inhibit Chlamydia trachomatis infectivity of human and mouse cell lines. Several subsequent studies have shown that IFN-I could indeed play important roles for inhibiting various stages of bacterial infections. Some examples include replication of Chlamydophila pneumoniae (36), recruitment of Myobacterium tuberculosis target cells into the lung during early infection (37), and invasion and transmigration of Streptococcus pneumoniae in the lungs (38). However, IFN-Is do not always appear to render beneficial outcomes in anti-bacterial immunity. Several studies have reported that IFN-AR deficient mice are better protected than WT controls following bacterial infections such as Ehrlichia muris (39), Chlamydia muridarum (40), Listeria monocytogenes (41, 42), Myobacterium species (43, 44), and Francisella tularensis (45). Furthermore, induction of IFN-Is following virus infections could make hosts more susceptible to secondary bacterial infections (4648). The mechanisms as to how IFN-Is exacerbate or make hosts more susceptible to bacterial disease may vary depending on the infection. For instance, IFN-I mediated disease exacerbation has been linked to reduction of interleukin (IL)-17 expressing γδ T cells, increased expression of IL-10 or reduction in cell-mediate immune responses following F. tularensis, M. Leprae, or L. monocytogenes, respectively (42, 44, 45).

Several reports suggest that the detrimental effects of IFN-Is could also support the establishment of persistent viral infections depending on the quantities and duration of IFN-I induction. IFN-Is have been shown to play significant roles in inhibiting various stages (e.g., replication, virus assembly, protein trafficking, and transcription) of HIV-1 life cycle (4953). However, sustained unlike transient production of IFN-Is resulting from chronic stimulation of pDCs has been proposed to facilitate HIV-1 persistence (54). Similarly following clone 13 LCMV infection transient (within 24 h) hyper-induction of IFN-α and -β has been reported to exacerbate virus pathogenesis and promote viral persistence (19). However, in the same study IFN-Is were crucial for the control of acute Armstrong LCMV infection, which was likely due to lower IFN-I induction following Armstrong compared to clone 13 LCMV infection. In chronic simian immunodeficiency virus (SIV) infection studies, disease free phenotypes of sooty mangabeys have been associated with the abolishment of interferon stimulated gene expression during chronic, but not in acute phase infection (55). Overall, it can be speculated that early, transient yet non-excessive induction of IFN-Is (at least α and β species) are important in the control of acute viral infections. On the contrary, chronic and/or hyper-induction of IFN-Is could provide an environment for enhanced persistence and/or pathogenesis of chronic viral infections.

IFN-Is and Regulation of Adaptive Immunity

Apart from their most celebrated role as direct anti-viral agents, IFN-Is have also been increasingly recognized as potent regulators of cellular immune responses. Of particular interest to vaccine development has been the ability of these cytokines to regulate adaptive immune responses and this aspect is discussed here.

Dendritic cells are often crucial for initiating adaptive immune responses and serve as important targets for IFN-Is to regulate adaptive immunity. Exposure of IFN-Is facilitates maturation of DCs via increasing the expression of DC-associated chemokine receptors, co-stimulatory molecules, and major histocompatibility complex class I and class II antigen presentation (5660). Consequently, DCs that mature following IFN-I exposure can effectively prime protective T cell responses (61). A caveat here is that IFN-I responses could operate in a threshold dependent manner where excessive responsiveness is inhibitory to the ability of DCs to prime T cell responses. For instance, following LCMV infection higher induction of IFN-Is has been associated with heightened expression of programed death-ligand 1 (PD-L1) on DCs and PD-L1 interaction with programed death 1 (PD-1) on T cells can inhibit T cell activation (19, 62).

IFN-Is could also act directly on lymphocytes to alter adaptive immune outcomes. Naïve B cells up-regulate the expression of activation markers CD69, CD86, and CD25 following IFN-I exposure in vitro (63), but in vivo IFN-Is only up-regulate CD69 and CD86 expression on naïve B and T cells (64). The consequences of up-regulating these activation markers are not clear, but in vitro studies suggest it could serve to reduce the activation thresholds of naïve B cells unlike T cells (63, 65). Alternatively, CD69 expression resulting from IFN-I exposure can down-regulate sphigosine-1 phosphate receptor-1 on naïve lymphocytes to retain these cells in secondary lymphoid organs (66). This retention mechanism could facilitate a more durable interaction between naïve lymphocytes and DCs for efficient lymphocyte activation to occur. IFN-Is have been reported to represent a distinct third signal for naïve T cell activation to occur and prevent the expansion of regulatory T cells that can inhibit T cell activation (6769). Furthermore, IFN-Is regulate the functions of lymphocytes even after naïve lymphocyte activation or effector/memory differentiation. Some examples of this include IFN-I mediated enhancement in cell division (63, 70), survival (71, 72), interferon-γ secretion (73), cytotoxicity (74), germinal center formation, and antibody isotype switching (75).

Despite the many studies demonstrating that IFN-Is are capable of boosting adaptive immunity; there have also been several studies in bacterial and chronic viral infection settings suggesting that IFN-I signaling leads to IL-10 production (19, 44, 76, 77). IL-10 is thought to be detrimental to the clearance of these pathogens as has been demonstrated with HIV-1 (78). It is likely that IFN-Is up-regulate PD-1 expression (e.g., on regulatory T cells) and PD-L1 (e.g., on DCs) on cells resulting in a milieu where PD-1/PD-L1 interactions occur; this could facilitate IL-10 production and exhaustion of T cell function during chronic viral infections (19, 7680). A caveat here is that IFN-Is in some instances can also inhibit IL-10 production and IL-10 production can occur independently of IFN-I signaling (76, 81). Furthermore, IFN-Is up-regulate pro-apoptotic molecules such as Bak on T cells to induce apoptosis independently of T cell exhaustion (82).

Overall, IFN-Is play pivotal roles in boosting adaptive immunity, but the switch from becoming a booster to an inhibitor of adaptive immunity may reflect on how much apoptosis, PD-1/PD-L1 interactions and IL-10 signaling are induced on immune cells due to IFN-Is.

The Use of IFN-Is as Therapeutics and Adjuvants

The development of efficient methods to purify IFN-I and subsequent high yield purification of IFN-α2 during the late 1970s paved way for the first IFN-I based human clinical trial in 1986 where IFN-α2 was used for treating hairy cell leukemia (83, 84). Since then the therapeutic use of IFN-Is have shown promising outcomes for treatment of several cancers and viral infections. Therapeutic administration of pegylated IFN-α2 have rendered potent anti-viral and immune enhancing effects against hepatitis B virus infection (85, 86). A recent clinical trial has shown that similar outcomes could be achieved even when pegylated IFN-α2 is administered to HIV-infected patients (87). Systemic administration of IFN-α and/or IFN-β has also been reported to reduce viral growth and clinical manifestations of herpes zoster, herpes simplex virus, and cytomegalovirus (CMV) infections (8891). Furthermore, systemic or intralesional administration of IFN-α and/or IFN-β has been shown to induce a regression of skin-associated wart infections following papilloma virus infections (9298). IFN-Is have also been used in synergic regimens where administration of IFN-α2 or -β2 and anti-viral drugs (e.g., ribavirin and faldaprevir) could effectively reduce viral loads of certain hepatitis C virus (HCV) genotypes and is currently the best treatment for HCV-infected patients (99102). A caveat here is that these regimens have also been reported to cause adverse side-effects (103). Apart from treatment of pathogen infections, IFN-Is especially IFN-α2, have also been used for treatment and regression of various cancers (e.g., leukemia, prostrate cancer, and cervical intraepithelial neoplasia) (104106).

Studies in pre-clinical models suggest that IFN-Is could also be potent vaccine adjuvants for inducing adaptive immune responses. Some examples include when an influenza vaccine adjuvanted with IFN-α/β administered mucosally induced significantly higher IgG2a and IgA antibody responses and protection compared to non-adjuvanted vaccines (107, 108). Interestingly, the species of IFN-Is used as immune adjuvants could have different immune outcomes in terms of enhancing adaptive immunity. Studies in our laboratory suggest that recombinant pox viral vectors encoding IFN-β compared to those encoding IFN-α4 or IFN-ε significantly enhanced systemic T cell immunity against co-encoded antigens in prime-boost vaccination settings (109). However, Xi et al. (110) using similar prime-boost vaccination settings demonstrated that the use of IFN-ε was much more efficient in inducing T cell immunity in mucosal compartments (e.g., lung and gut) compared to IFN-α4 and IFN-β when used as vaccine adjuvants. Another important consideration here is that the vaccine vectors (i.e., pox viruses) used in our studies are acute attenuated viruses and do not chronically induce IFN-Is as is usually the case with persistent virus infections.

There are several confounding factors that could dictate the use of IFN-I in therapy and as vaccine adjuvants. Firstly, unique biological effects have been reported with different members of the IFN-I family and subtypes of IFN-α. Thus, the choice of IFN-I species (e.g., IFN-α2 or IFN-β) could dictate the success of IFN-I treatment or IFN-I based vaccine formulations. Secondly, members of the IFN-I family have different binding affinities and kinetics to the IFN-AR subunits with current comparative studies suggesting that IFN-β has the highest affinity to IFN-AR and anti-viral capacity (111113). A caveat with these studies is that not all members of the IFN-I family were compared. Thirdly, IFN-Is can cause numerous adverse side-effects and induce autoimmunity (e.g., lupus, thyroiditis, diabetes, dermatitis, Sjogren’s syndrome, and arthritis) especially in patients with a history of autoimmune manifestations (114). The autoimmune outcomes in these settings are thought to be a combination of tolerogenic immune function failures and IFN-I mediated maturation of DCs that present autoantigens to activate autoreactive T cells and B cells that make autoantibodies (115).

Collectively, IFN-Is have shown considerable promise for the treatment of cancers and pathogen infections (e.g., chronic viruses) in some clinical settings. IFN-Is are also promising for use as vaccine adjuvants, but the species of IFN-Is used for this purpose could have a significant bearing on adaptive immunity generated at certain immune compartments. For instance, IFN-β could be used to effectively enhance systemic T cell immune responses, whereas IFN-ε is more promising as an adjuvant to enhance mucosal T cell immunity in the lung and the gut mucosae.

Importance of IFN-ε in Immunity and Vaccine Development

Most studies investigating the roles of IFN-Is have done so mainly analyzing the roles of IFN-α and -β. However, investigating the roles of other IFN-I family members is beneficial for effective therapeutic and vaccine development strategies especially given that higher induction of IFN-α and -β could be detrimental to the host as discussed previously. For this purpose, it is indeed intriguing to evaluate the roles of IFN-ε, which unlike other IFN-Is is constitutively expressed and plays various protective roles in reproductive tissues, gut, lung, and the brain (Table 1). Since our initial studies characterizing the roles of IFN-ε in inducing anti-viral states on cells (109), we have found that this cytokine also possesses potent immune regulatory capacity. Our recent studies indicated that, intranasal immunization of mice with vaccinia virus (VV) encoding murine IFN-ε (VV-HIV-IFN-ε) unlike IFN-α (VV-HIV-IFN-α4) or IFN-β (VV-HIV-IFN-β) could induce rapid clearance of VV in the lung (110). Viral clearance in this instance correlated with several immune outcomes: (i) elevated lung VV-specific CD8+CD107a+IFN-γ+ cell population expressing activation markers CD69/CD103, (ii) enhanced lymphocyte recruitment to lung alveoli with reduced inflammation, and (iii) highly functional CD8+CD4+ double positive T cell subset [CD3highC–C chemokine receptor (CCR)7highCD62Llow] in lung lymph nodes (110). Next when IFN-ε was used in an intranasal/intramuscular heterologous HIV-1 prime-boost vaccination regimen, elevated HIV-specific effector, but not memory CD8+ T cells responses were detected in spleen, genito-rectal nodes, and Peyer’s patches. Furthermore, homing marker α4β7 and CCR9 analysis showed that unlike other IFN-Is, IFN-ε promoted the migration of antigen-specific CD8+ T cells to the gut mucosae (110). These results for the first time established that unlike other IFN-Is, IFN-ε played a unique role at the mucosae. Another recent study has also further substantiated our findings demonstrating that IFN-ε deficient mice were more susceptible to intra-vaginal herpes simplex virus 2 and Chlamydia muridarum infections compared to wild-type mice (117). This suggests that IFN-ε could also be beneficial for the control of certain bacterial infections. A caveat here is that it is unknown whether IFN-ε could cause adverse side-effects in humans as it has not yet been used for treatment or vaccination purposes in humans.

Table 1

SiteFunctionReference
BrainMaintenance of the structure and function(116)
LungPromote clearance of viral infections(110)
Recruitment of unique yet highly anti-viral CD4+CD8+ T cells
GutEnhance expression of CCR9 and α4β7 on anti-viral T cells to promote homing to the gut (i.e., Peyer’s patches)(110)
Reproductive tissuesRegulation of embryonic development Protect male and female reproductive tissues against infections (e.g., herpes and Chlamydia)(117, 118)

Site-specific effects of IFN-ε.

Overall, IFN-ε has great potential to be used as a topical microbicide or a therapeutic to control local lung/gut infections or modulate tissue-specific immunity at sites where pathogens are initially encountered (i.e., mucosal surfaces). Specifically, IFN-ε’s ability to enhance CD8+ T cell homing to the gut [gut is the primary site of HIV virus replication and CD4+ T-cell depletion (119)] and also its ability to control infections at the lung mucosae suggest that administration of pegylated forms of IFN-ε or vaccines encoding IFN-ε could be effective for controlling mucosal pathogens such as HIV-1.

Concluding Remarks

The dual roles of IFN-Is in providing beneficial and detrimental effects to the host in pathogen control is intriguing for developing IFN-I based vaccines and therapies. Lessons learned from acute viral infection models and studies comparing acute versus chronic infection states suggest that transient, but not sustained and/or excessive induction of IFN-Is is likely to confer protective outcomes. IFN-Is have also proven to be promising therapeutic agents against various pathogens and cancers and could also be used as vaccine adjuvants. The caveat here is that the vaccine vector used should ideally not chronically stimulate the production of IFN-Is, which is expected to be detrimental for the generation of robust adaptive immune responses. Our laboratory and others have demonstrated that IFN-ε has great potential to provide protective outcomes against not only mucosal viral infections, but also certain mucosal bacterial infections. Keeping this in mind, more studies need to evaluate the contribution of the different species of IFN-Is not just IFN-α and -β in immunity against infections. These studies are expected to pave way for the development of novel and effective IFN-I based vaccines/therapies against chronic pathogens and cancers.

Statements

Acknowledgments

This work was supported by Australian National Health and Medical Research Council project grant award 525431 (Charani Ranasinghe) and ACH2 EOI grants (Charani Ranasinghe).

Conflict of interest

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.

References

  • 1

    IsaacsALindenmannJ. Virus interference. I. The interferon. Proc R Soc Lond B Biol Sci (1957) 147(927):25867.10.1098/rspb.1957.0049

  • 2

    AdolfGR. Human interferon omega – a review. Mult Scler (1995) 1(Suppl 1):S447.

  • 3

    FosterGRFinterNB. Are all type I human interferons equivalent?J Viral Hepat (1998) 5(3):14352.10.1046/j.1365-2893.1998.00103.x

  • 4

    KawamotoSOritaniKAsadaHTakahashiIIshikawaJYoshidaHet alAntiviral activity of limitin against encephalomyocarditis virus, herpes simplex virus, and mouse hepatitis virus: diverse requirements by limitin and alpha interferon for interferon regulatory factor 1. J Virol (2003) 77(17):962231.10.1128/JVI.77.17.9622-9631.2003

  • 5

    MartalJLCheneNMHuynhLPL’HaridonRMReinaudPBGuillomotMWet alIFN-tau: a novel subtype I IFN1. Structural characteristics, non-ubiquitous expression, structure-function relationships, a pregnancy hormonal embryonic signal and cross-species therapeutic potentialities. Biochimie (1998) 80(8–9):75577.10.1016/S0300-9084(99)80029-7

  • 6

    LaFleurDWNardelliBTsarevaTMatherDFengPSemenukMet alInterferon-kappa, a novel type I interferon expressed in human keratinocytes. J Biol Chem (2001) 276(43):3976571.10.1074/jbc.M102502200

  • 7

    OritaniKMedinaKLTomiyamaYIshikawaJOkajimaYOgawaMet alLimitin: an interferon-like cytokine that preferentially influences B-lymphocyte precursors. Nat Med (2000) 6(6):65966.10.1038/76233

  • 8

    PestkaSLangerJAZoonKCSamuelCE. Interferons and their actions. Annu Rev Biochem (1987) 56:72777.10.1146/annurev.bi.56.070187.003455

  • 9

    StarkGRKerrIMWilliamsBRSilvermanRHSchreiberRD. How cells respond to interferons. Annu Rev Biochem (1998) 67:22764.10.1146/annurev.biochem.67.1.227

  • 10

    SenGC. Viruses and interferons. Annu Rev Microbiol (2001) 55:25581.10.1146/annurev.micro.55.1.255

  • 11

    KarpovAV. Endogenous and exogenous interferons in HIV-infection. Eur J Med Res (2001) 6(12):50724.

  • 12

    IvashkivLBDonlinLT. Regulation of type I interferon responses. Nat Rev Immunol (2014) 14(1):3649.10.1038/nri3581

  • 13

    SiegalFPKadowakiNShodellMFitzgerald-BocarslyPAShahKHoSet alThe nature of the principal type 1 interferon-producing cells in human blood. Science (1999) 284(5421):18357.10.1126/science.284.5421.1835

  • 14

    ColonnaMTrinchieriGLiuYJ. Plasmacytoid dendritic cells in immunity. Nat Immunol (2004) 5(12):121926.10.1038/ni1141

  • 15

    HallJCRosenA. Type I interferons: crucial participants in disease amplification in autoimmunity. Nat Rev Rheumatol (2010) 6(1):409.10.1038/nrrheum.2009.237

  • 16

    DhondtKPMathieuCChalonsMReynaudJMVallveARaoulHet alType I interferon signaling protects mice from lethal henipavirus infection. J Infect Dis (2013) 207(1):14251.10.1093/infdis/jis653

  • 17

    GerlachNSchimmerSWeissSKalinkeUDittmerU. Effects of type I interferons on Friend retrovirus infection. J Virol (2006) 80(7):343844.10.1128/JVI.80.7.3438-3444.2006

  • 18

    LobigsMMullbacherAWangYPavyMLeeE. Role of type I and type II interferon responses in recovery from infection with an encephalitic flavivirus. J Gen Virol (2003) 84(Pt 3):56772.10.1099/vir.0.18654-0

  • 19

    TeijaroJRNgCLeeAMSullivanBMSheehanKCWelchMet alPersistent LCMV infection is controlled by blockade of type I interferon signaling. Science (2013) 340(6129):20711.10.1126/science.1235214

  • 20

    DowallSDFindlay-WilsonSRaynerEPearsonGPickersgillJRuleAet alHazara virus infection is lethal for adult type I interferon receptor-knockout mice and may act as a surrogate for infection with the human-pathogenic Crimean-Congo hemorrhagic fever virus. J Gen Virol (2012) 93(Pt 3):5604.10.1099/vir.0.038455-0

  • 21

    ZustRTohYXValdesICernyDHeinrichJHermidaLet alType I IFN signals in macrophages and dendritic cells control dengue virus infection: implications for a new mouse model to test dengue vaccines. J Virol (2014) 88(13):727685.10.1128/JVI.03827-13

  • 22

    GoritzkaMDurantLRPereiraCSalek-ArdakaniSOpenshawPJJohanssonC. Interferon-α/β receptor signaling amplifies early pro-inflammatory 2 cytokine production in the lung during Respiratory Syncytial Virus 3 infection. J Virol (2014) 88(11):612836.10.1128/JVI.00333-14

  • 23

    CarreroJA. Confounding roles for type I interferons during bacterial and viral pathogenesis. Int Immunol (2013) 25(12):6639.10.1093/intimm/dxt050

  • 24

    AlsharifiMRegnerMBlandenRLobigsMLeeEKoskinenAet alExhaustion of type I interferon response following an acute viral infection. J Immunol (2006) 177(5):323541.10.4049/jimmunol.177.5.3235

  • 25

    KamgaIKahiSDeveliogluLLichtnerMMaranonCDeveauCet alType I interferon production is profoundly and transiently impaired in primary HIV-1 infection. J Infect Dis (2005) 192(2):30310.10.1086/430931

  • 26

    MartinelliECicalaCVan RykDGoodeDJMacleodKArthosJet alHIV-1 gp120 inhibits TLR9-mediated activation and IFN-{alpha} secretion in plasmacytoid dendritic cells. Proc Natl Acad Sci U S A (2007) 104(9):3396401.10.1073/pnas.0611353104

  • 27

    RasaiyaahJTanCPFletcherAJPriceAJBlondeauCHilditchLet alHIV-1 evades innate immune recognition through specific cofactor recruitment. Nature (2013) 503(7476):4025.10.1038/nature12769

  • 28

    Fitzgerald-BocarslyPJacobsES. Plasmacytoid dendritic cells in HIV infection: striking a delicate balance. J Leukoc Biol (2010) 87(4):60920.10.1189/jlb.0909635

  • 29

    SoumelisVScottIGheyasFBouhourDCozonGCotteLet alDepletion of circulating natural type 1 interferon-producing cells in HIV-infected AIDS patients. Blood (2001) 98(4):90612.10.1182/blood.V98.4.906

  • 30

    PacanowskiJKahiSBailletMLebonPDeveauCGoujardCet alReduced blood CD123+ (lymphoid) and CD11c+ (myeloid) dendritic cell numbers in primary HIV-1 infection. Blood (2001) 98(10):301621.10.1182/blood.V98.10.3016

  • 31

    SchmidtBScottIWhitmoreRGFosterHFujimuraSSchmitzJet alLow-level HIV infection of plasmacytoid dendritic cells: onset of cytopathic effects and cell death after PDC maturation. Virology (2004) 329(2):2808.10.1016/j.virol.2004.08.016

  • 32

    BruelTDupuySDemoulinsTRogez-KreuzCDutrieuxJCorneauAet alPlasmacytoid dendritic cell dynamics tune interferon-alfa production in SIV-infected cynomolgus macaques. PLoS Pathog (2014) 10(1):e1003915.10.1371/journal.ppat.1003915

  • 33

    GuhaDAyyavooV. Innate immune evasion strategies by human immunodeficiency virus type 1. ISRN AIDS (2013) 2013:954806.10.1155/2013/954806

  • 34

    Critchley-ThorneRJSimonsDLYanNMiyahiraAKDirbasFMJohnsonDLet alImpaired interferon signaling is a common immune defect in human cancer. Proc Natl Acad Sci U S A (2009) 106(22):90105.10.1073/pnas.0901329106

  • 35

    de la MazaLMPetersonEMGoebelJMFennieCWCzarnieckiCW. Interferon-induced inhibition of Chlamydia trachomatis: dissociation from antiviral and antiproliferative effects. Infect Immun (1985) 47:71922.

  • 36

    BussCOpitzBHockeACLippmannJvan LaakVHippenstielSet alEssential role of mitochondrial antiviral signaling, IFN regulatory factor (IRF)3, and IRF7 in Chlamydophila pneumoniae-mediated IFN-beta response and control of bacterial replication in human endothelial cells. J Immunol (2010) 184:30728.10.4049/jimmunol.0902947

  • 37

    DesvignesLWolfAJErnstJD. Dynamic roles of type I and type II IFNs in early infection with Mycobacterium tuberculosis. J Immunol (2012) 188:620515.10.4049/jimmunol.1200255

  • 38

    LeMessurierKSHackerHChiLTuomanenERedeckeV. Type I interferon protects against pneumococcal invasive disease by inhibiting bacterial transmigration across the lung. PLoS Pathog (2013) 9:e1003727.10.1371/journal.ppat.1003727

  • 39

    ZhangYThaiVMcCabeAJonesMMacNamaraKC. Type I interferons promote severe disease in a mouse model of lethal ehrlichiosis. Infect Immun (2014) 82(4):1698709.10.1128/IAI.01564-13

  • 40

    NagarajanUMPrantnerDSikesJDAndrewsCWJr.GoodwinAMNagarajanSet alType I interferon signaling exacerbates Chlamydia muridarum genital infection in a murine model. Infect Immun (2008) 76(10):46428.10.1128/IAI.00629-08

  • 41

    AuerbuchVBrockstedtDGMeyer-MorseNO’RiordanMPortnoyDA. Mice lacking the type I interferon receptor are resistant to Listeria monocytogenes. J Exp Med (2004) 200(4):52733.10.1084/jem.20040976

  • 42

    ArcherKADurackJPortnoyDA. STING-dependent type I IFN production inhibits cell-mediated immunity to Listeria monocytogenes. PLoS Pathog (2014) 10(1):e1003861.10.1371/journal.ppat.1003861

  • 43

    MancaCTsenovaLFreemanSBarczakAKToveyMMurrayPJet alHypervirulent M. tuberculosis W/Beijing strains upregulate type I IFNs and increase expression of negative regulators of the Jak-Stat pathway. J Interferon Cytokine Res (2005) 25(11):694701.10.1089/jir.2005.25.694

  • 44

    TelesRMGraeberTGKrutzikSRMontoyaDSchenkMLeeDJet alType I interferon suppresses type II interferon-triggered human anti-mycobacterial responses. Science (2013) 339(6126):144853.10.1126/science.1233665

  • 45

    HenryTKirimanjeswaraGSRubyTJonesJWPengKPerretMet alType I IFN signaling constrains IL-17A/F secretion by gammadelta T cells during bacterial infections. J Immunol (2010) 184(7):375567.10.4049/jimmunol.0902065

  • 46

    NavariniAARecherMLangKSGeorgievPMeurySBergthalerAet alIncreased susceptibility to bacterial superinfection as a consequence of innate antiviral responses. Proc Natl Acad Sci U S A (2006) 103(42):155359.10.1073/pnas.0607325103

  • 47

    LiWMoltedoBMoranTM. Type I interferon induction during influenza virus infection increases susceptibility to secondary Streptococcus pneumoniae infection by negative regulation of gammadelta T cells. J Virol (2012) 86(22):1230412.10.1128/JVI.01269-12

  • 48

    TrinchieriG. Type I interferon: friend or foe?J Exp Med (2010) 207(10):205363.10.1084/jem.20101664

  • 49

    BarrSDSmileyJRBushmanFD. The interferon response inhibits HIV particle production by induction of TRIM22. PLoS Pathog (2008) 4(2):e1000007.10.1371/journal.ppat.1000007

  • 50

    SakumaRNoserJAOhmineSIkedaY. Rhesus monkey TRIM5alpha restricts HIV-1 production through rapid degradation of viral Gag polyproteins. Nat Med (2007) 13(5):6315.10.1038/nm1562

  • 51

    SokolskajaELubanJ. Cyclophilin, TRIM5, and innate immunity to HIV-1. Curr Opin Microbiol (2006) 9(4):4048.10.1016/j.mib.2006.06.011

  • 52

    NisoleSStoyeJPSaibA. TRIM family proteins: retroviral restriction and antiviral defence. Nat Rev Microbiol (2005) 3(10):799808.10.1038/nrmicro1248

  • 53

    OkumuraALuGPitha-RoweIPithaPM. Innate antiviral response targets HIV-1 release by the induction of ubiquitin-like protein ISG15. Proc Natl Acad Sci U S A (2006) 103(5):14405.10.1073/pnas.0510518103

  • 54

    BoassoARoyleCMDoumazosSAquinoVNBiasinMPiacentiniLet alOveractivation of plasmacytoid dendritic cells inhibits antiviral T-cell responses: a model for HIV immunopathogenesis. Blood (2011) 118(19):515262.10.1182/blood-2011-03-344218

  • 55

    BosingerSELiQGordonSNKlattNRDuanLXuLet alGlobal genomic analysis reveals rapid control of a robust innate response in SIV-infected sooty mangabeys. J Clin Invest (2009) 119(12):355672.10.1172/JCI40115

  • 56

    CellaMSalioMSakakibaraYLangenHJulkunenILanzavecchiaA. Maturation, activation, and protection of dendritic cells induced by double-stranded RNA. J Exp Med (1999) 189(5):8219.10.1084/jem.189.5.821

  • 57

    ParlatoSSantiniSMLapentaCDi PucchioTLogozziMSpadaMet alExpression of CCR-7, MIP-3beta, and Th-1 chemokines in type I IFN-induced monocyte-derived dendritic cells: importance for the rapid acquisition of potent migratory and functional activities. Blood (2001) 98(10):30229.10.1182/blood.V98.10.3022

  • 58

    MontoyaMSchiavoniGMatteiFGresserIBelardelliFBorrowPet alType I interferons produced by dendritic cells promote their phenotypic and functional activation. Blood (2002) 99(9):326371.10.1182/blood.V99.9.3263

  • 59

    WelshRMBahlKMarshallHDUrbanSL. Type 1 interferons and antiviral CD8 T-cell responses. PLoS Pathog (2012) 8(1):e1002352.10.1371/journal.ppat.1002352

  • 60

    SimmonsDPWearschPACanadayDHMeyersonHJLiuYCWangYet alType I IFN drives a distinctive dendritic cell maturation phenotype that allows continued class II MHC synthesis and antigen processing. J Immunol (2012) 188(7):311626.10.4049/jimmunol.1101313

  • 61

    DiamondMSKinderMMatsushitaHMashayekhiMDunnGPArchambaultJMet alType I interferon is selectively required by dendritic cells for immune rejection of tumors. J Exp Med (2011) 208(10):19892003.10.1084/jem.20101158

  • 62

    CaiGKarniAOliveiraEMWeinerHLHaflerDAFreemanGJ. PD-1 ligands, negative regulators for activation of naive, memory, and recently activated human CD4+ T cells. Cell Immunol (2004) 230(2):8998.10.1016/j.cellimm.2004.09.004

  • 63

    BraunDCaramalhoIDemengeotJ. IFN-alpha/beta enhances BCR-dependent B cell responses. Int Immunol (2002) 14(4):4119.10.1093/intimm/14.4.411

  • 64

    AlsharifiMLobigsMRegnerMLeeEKoskinenAMullbacherA. Type I interferons trigger systemic, partial lymphocyte activation in response to viral infection. J Immunol (2005) 175(7):463540.10.4049/jimmunol.175.7.4635

  • 65

    WijesundaraDKKumarSAlsharifiMMullbacherARegnerM. Antigen-specific activation thresholds of CD8+ T cells are independent of IFN-I-mediated partial lymphocyte activation. Int Immunol (2010) 22(9):75767.10.1093/intimm/dxq064

  • 66

    ShiowLRRosenDBBrdickovaNXuYAnJLanierLLet alCD69 acts downstream of interferon-alpha/beta to inhibit S1P1 and lymphocyte egress from lymphoid organs. Nature (2006) 440(7083):5404.10.1038/nature04606

  • 67

    CurtsingerJMSchmidtCSMondinoALinsDCKedlRMJenkinsMKet alInflammatory cytokines provide a third signal for activation of naive CD4+ and CD8+ T cells. J Immunol (1999) 162(6):325662.

  • 68

    CurtsingerJMGernerMYLinsDCMescherMF. Signal 3 availability limits the CD8 T cell response to a solid tumor. J Immunol (2007) 178(11):675260.10.4049/jimmunol.178.11.6752

  • 69

    SrivastavaSKochMAPepperMCampbellDJ. Type I interferons directly inhibit regulatory T cells to allow optimal antiviral T cell responses during acute LCMV infection. J Exp Med (2014) 211(5):96174.10.1084/jem.20131556

  • 70

    Starbeck-MillerGRXueHHHartyJT. IL-12 and type I interferon prolong the division of activated CD8 T cells by maintaining high-affinity IL-2 signaling in vivo. J Exp Med (2014) 211(1):10520.10.1084/jem.20130901

  • 71

    MarrackPKapplerJMitchellT. Type I interferons keep activated T cells alive. J Exp Med (1999) 189(3):52130.10.1084/jem.189.3.521

  • 72

    KolumamGAThomasSThompsonLJSprentJMurali-KrishnaK. Type I interferons act directly on CD8 T cells to allow clonal expansion and memory formation in response to viral infection. J Exp Med (2005) 202(5):63750.10.1084/jem.20050821

  • 73

    BrinkmannVGeigerTAlkanSHeusserCH. Interferon alpha increases the frequency of interferon gamma-producing human CD4+ T cells. J Exp Med (1993) 178(5):165563.10.1084/jem.178.5.1655

  • 74

    KohlmeierJECookenhamTRobertsADMillerSCWoodlandDL. Type I interferons regulate cytolytic activity of memory CD8(+) T cells in the lung airways during respiratory virus challenge. Immunity (2010) 33(1):96105.10.1016/j.immuni.2010.06.016

  • 75

    Le BonAThompsonCKamphuisEDurandVRossmannCKalinkeUet alCutting edge: enhancement of antibody responses through direct stimulation of B and T cells by type I IFN. J Immunol (2006) 176(4):20748.10.4049/jimmunol.176.8.4682

  • 76

    ZhangLYuanSChengGGuoB. Type I IFN promotes IL-10 production from T cells to suppress Th17 cells and Th17-associated autoimmune inflammation. PLoS One (2011) 6(12):e28432.10.1371/journal.pone.0028432

  • 77

    StewartCAMethenyHIidaNSmithLHansonMSteinhagenFet alInterferon-dependent IL-10 production by Tregs limits tumor Th17 inflammation. J Clin Invest (2013) 123(11):485974.10.1172/JCI65180

  • 78

    SaidEADupuyFPTrautmannLZhangYShiYEl-FarMet alProgrammed death-1-induced interleukin-10 production by monocytes impairs CD4+ T cell activation during HIV infection. Nat Med (2010) 16(4):4529.10.1038/nm.2106

  • 79

    BarberDLWherryEJMasopustDZhuBAllisonJPSharpeAHet alRestoring function in exhausted CD8 T cells during chronic viral infection. Nature (2006) 439(7077):6827.10.1038/nature04444

  • 80

    TerawakiSChikumaSShibayamaSHayashiTYoshidaTOkazakiTet alIFN-alpha directly promotes programmed cell death-1 transcription and limits the duration of T cell-mediated immunity. J Immunol (2011) 186(5):27729.10.4049/jimmunol.1003208

  • 81

    LinLHouJMaFWangPLiuXLiNet alType I IFN inhibits innate IL-10 production in macrophages through histone deacetylase 11 by downregulating microRNA-145. J Immunol (2013) 191(7):3896904.10.4049/jimmunol.1203450

  • 82

    FraiettaJAMuellerYMYangGBoesteanuACGraciasDTDoDHet alType I interferon upregulates Bak and contributes to T cell loss during human immunodeficiency virus (HIV) infection. PLoS Pathog (2013) 9(10):e1003658.10.1371/journal.ppat.1003658

  • 83

    RubinsteinMRubinsteinSFamillettiPCGrossMSMillerRSWaldmanAAet alHuman leukocyte interferon purified to homogeneity. Science (1978) 202:128990.10.1126/science.725605

  • 84

    PestkaS. The interferons: 50 years after their discovery, there is much more to learn. J Biol Chem (2007) 282:2004751.10.1074/jbc.R700004200

  • 85

    LauGKPiratvisuthTLuoKXMarcellinPThongsawatSCooksleyGet alPeginterferon Alfa-2a, lamivudine, and the combination for HBeAg-positive chronic hepatitis B. N Engl J Med (2005) 352(26):268295.10.1056/NEJMoa043470

  • 86

    MarcellinPLauGKBoninoFFarciPHadziyannisSJinRet alPeginterferon alfa-2a alone, lamivudine alone, and the two in combination in patients with HBeAg-negative chronic hepatitis B. N Engl J Med (2004) 351(12):120617.10.1056/NEJMoa040431

  • 87

    AzzoniLFoulkesASPapasavvasEMexasAMLynnKMMounzerKet alPegylated Interferon alfa-2a monotherapy results in suppression of HIV type 1 replication and decreased cell-associated HIV DNA integration. J Infect Dis (2013) 207(2):21322.10.1093/infdis/jis663

  • 88

    JonesBRCosterDJFalconMGCantellK. Topical therapy of ulcerative herpetic keratitis with human interferon. Lancet (1976) 2(7977):128.10.1016/S0140-6736(76)92850-6

  • 89

    LuiSFAliAAGrundyJEFernandoONGriffithsPDSwenyP. Double-blind, placebo-controlled trial of human lymphoblastoid interferon prophylaxis of cytomegalovirus infection in renal transplant recipients. Nephrol Dial Transplant (1992) 7(12):12307.

  • 90

    MeriganTCRandKHPollardRBAbdallahPSJordanGWFriedRP. Human leukocyte interferon for the treatment of herpes zoster in patients with cancer. N Engl J Med (1978) 298(18):9817.10.1056/NEJM197805042981801

  • 91

    PazinGJArmstrongJALamMTTarrGCJannettaPJHoM. Prevention of reactivated herpes simplex infection by human leukocyte interferon after operation on the trigeminal root. N Engl J Med (1979) 301(5):22530.10.1056/NEJM197908023010501

  • 92

    DeunasLAlcantudVAlvarezFArteagaJBenitezABopuzaMet alUse of interferon-alpha in laryngeal papillomatosis: eight years of the Cuban national programme. J Laryngol Otol (1997) 111(2):13440.10.1017/S0022215100136667

  • 93

    EronLJJudsonFTuckerSPrawerSMillsJMurphyKet alInterferon therapy for condylomata acuminata. N Engl J Med (1986) 315(17):105964.10.1056/NEJM198610233151704

  • 94

    HaglundSLundquistPGCantellKStranderH. Interferon therapy in juvenile laryngeal papillomatosis. Arch Otolaryngol (1981) 107(6):32732.10.1001/archotol.1981.00790420001001

  • 95

    HealyGBGelberRDTrowbridgeALGrundfastKMRubenRJPriceKN. Treatment of recurrent respiratory papillomatosis with human leukocyte interferon. Results of a multicenter randomized clinical trial. N Engl J Med (1988) 319(7):4017.10.1056/NEJM198808183190704

  • 96

    LaceMJAnsonJRKlingelhutzAJHaradaHTaniguchiTBosslerADet alInterferon-beta treatment increases human papillomavirus early gene transcription and viral plasmid genome replication by activating interferon regulatory factor (IRF)-1. Carcinogenesis (2009) 30(8):133644.10.1093/carcin/bgp150

  • 97

    PazinGJHoMHaverkosHWArmstrongJABreinigMCWechslerHLet alEffects of interferon-alpha on human warts. J Interferon Res (1982) 2(2):23543.10.1089/jir.1982.2.235

  • 98

    WeckPKBuddinDAWhisnantJK. Interferons in the treatment of genital human papillomavirus infections. Am J Med (1988) 85(2A):15964.

  • 99

    McHutchisonJGGordonSCSchiffERShiffmanMLLeeWMRustgiVKet alInterferon alfa-2b alone or in combination with ribavirin as initial treatment for chronic hepatitis C. Hepatitis Interventional Therapy Group. N Engl J Med (1998) 339(21):148592.10.1056/NEJM199811193392101

  • 100

    PoynardTMarcellinPLeeSSNiederauCMinukGSIdeoGet alRandomised trial of interferon alpha2b plus ribavirin for 48 weeks or for 24 weeks versus interferon alpha2b plus placebo for 48 weeks for treatment of chronic infection with hepatitis C virus. International Hepatitis Interventional Therapy Group (IHIT). Lancet (1998) 352(9138):142632.10.1016/S0140-6736(98)07124-4

  • 101

    BakerDE. Pegylated interferon plus ribavirin for the treatment of chronic hepatitis C. Rev Gastroenterol Disord (2003) 3(2):93109.

  • 102

    SulkowskiMSAsselahTLalezariJFerenciPFainboimHLeggettBet alFaldaprevir combined with pegylated interferon alfa-2a and ribavirin in treatment-naive patients with chronic genotype 1 HCV: SILEN-C1 trial. Hepatology (2013) 57(6):214354.10.1002/hep.26276

  • 103

    TakagiHHoshinoTNaganumaAKoitabashiEUeharaSSakamotoNet alDrug induced hypersensitivity syndrome by triple therapy of peginterferon alpha2b, ribavirin and telaprevir in patient with double positive for HBV and HCV. Hepatogastroenterology (2013) 60(127):155760.

  • 104

    BonifaziFde VivoARostiGGuilhotFGuilhotJTrabacchiEet alChronic myeloid leukemia and interferon-alpha: a study of complete cytogenetic responders. Blood (2001) 98(10):307481.10.1182/blood.V98.10.3074

  • 105

    LiYFWangQZZhangTTLiLWangJPDingGFet alLow dose of interferon-alpha improves the clinical outcomes of docetaxel in patients with castration-resistant prostate cancer: a pilot study. Oncol Lett (2014) 7(1):12530.10.3892/ol.2013.1653

  • 106

    MachadoFAAbdallaDRMontesLEtchebehereRMMichelinMAMurtaEF. An evaluation of immune system cell infiltrate in the cervical stroma of patients with grade III cervical intraepithelial neoplasia after treatment with intralesional alpha-2B interferon. Eur J Gynaecol Oncol (2014) 35(1):205.

  • 107

    BracciLCaniniIVendittiMSpadaMPuzelliSDonatelliIet alType I IFN as a vaccine adjuvant for both systemic and mucosal vaccination against influenza virus. Vaccine (2006) 24(Suppl 2):S256.10.1016/j.vaccine.2005.01.121

  • 108

    ProiettiEBracciLPuzelliSDi PucchioTSestiliPDe VincenziEet alType I IFN as a natural adjuvant for a protective immune response: lessons from the influenza vaccine model. J Immunol (2002) 169(1):37583.10.4049/jimmunol.169.1.375

  • 109

    DaySLRamshawIARamsayAJRanasingheC. Differential effects of the type I interferons alpha4, beta, and epsilon on antiviral activity and vaccine efficacy. J Immunol (2008) 180(11):715866.10.4049/jimmunol.180.11.7158

  • 110

    XiYDaySLJacksonRJRanasingheC. Role of novel type I interferon epsilon in viral infection and mucosal immunity. Mucosal Immunol (2012) 5(6):61022.10.1038/mi.2012.35

  • 111

    RoismanLCPiehlerJTrossetJYScheragaHASchreiberG. Structure of the interferon-receptor complex determined by distance constraints from double-mutant cycles and flexible docking. Proc Natl Acad Sci U S A (2001) 98:132316.10.1073/pnas.221290398

  • 112

    LamkenPLataSGavutisMPiehlerJ. Ligand-induced assembling of the type I interferon receptor on supported lipid bilayers. J Mol Biol (2004) 341:30318.10.1016/j.jmb.2004.05.059

  • 113

    JaksEGavutisMUzeGMartalJPiehlerJ. Differential receptor subunit affinities of type I interferons govern differential signal activation. J Mol Biol (2007) 366:52539.10.1016/j.jmb.2006.11.053

  • 114

    BanchereauJPascualV. Type I interferon in systemic lupus erythematosus and other autoimmune diseases. Immunity (2006) 25:38392.10.1016/j.immuni.2006.08.010

  • 115

    RonnblomL. The type I interferon system in the etiopathogenesis of autoimmune diseases. Ups J Med Sci (2011) 116:22737.10.3109/03009734.2011.624649

  • 116

    PengGLeiKJJinWGreenwell-WildTWahlSM. Induction of APOBEC3 family proteins, a defensive maneuver underlying interferon-induced anti-HIV-1 activity. J Exp Med (2006) 203(1):416.10.1084/jem.20051512

  • 117

    FungKYManganNECummingHHorvatJCMayallJRStifterSAet alInterferon-epsilon protects the female reproductive tract from viral and bacterial infection. Science (2013) 339(6123):108892.10.1126/science.1233321

  • 118

    HermantPFranciusCClotmanFMichielsT. IFN-epsilon is constitutively expressed by cells of the reproductive tract and is inefficiently secreted by fibroblasts and cell lines. PLoS One (2013) 8(8):e71320.10.1371/journal.pone.0071320

  • 119

    VeazeyRSDeMariaMChalifouxLVShvetzDEPauleyDRKnightHLet alGastrointestinal tract as a major site of CD4+ T cell depletion and viral replication in SIV infection. Science (1998) 280(5362):42731.10.1126/science.280.5362.427

Summary

Keywords

type I interferons, human immunodeficiency virus, IFN-ε, vaccine adjuvants, interferon immunity

Citation

Wijesundara DK, Xi Y and Ranasinghe C (2014) Unraveling the Convoluted Biological Roles of Type I Interferons in Infection and Immunity: A Way Forward for Therapeutics and Vaccine Design. Front. Immunol. 5:412. doi: 10.3389/fimmu.2014.00412

Received

30 May 2014

Accepted

13 August 2014

Published

29 August 2014

Volume

5 - 2014

Edited by

Christine Anne Biron, Brown University, USA

Reviewed by

Janis J. Weis, University of Utah, USA; Jieliang Li, Temple University, USA

Copyright

*Correspondence: Danushka Kumara Wijesundara, Virology Laboratory, Department of Surgery, Basil Hetzel Institute, 37a Woodville Road, Woodville, SA 5011, Australia e-mail:

Danushka Kumara Wijesundara and Yang Xi have contributed equally to this work.

This article was submitted to Microbial Immunology, a section of the journal Frontiers in Immunology.

Disclaimer

All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher.

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