From: Subject: IL-23 promotes tumour incidence and growth : Nature Date: Fri, 19 May 2006 11:36:53 +0200 MIME-Version: 1.0 Content-Type: multipart/related; type="text/html"; boundary="----=_NextPart_000_001F_01C67B38.869A4500" X-MimeOLE: Produced By Microsoft MimeOLE V6.00.2900.2869 This is a multi-part message in MIME format. ------=_NextPart_000_001F_01C67B38.869A4500 Content-Type: text/html; charset="iso-8859-1" Content-Transfer-Encoding: quoted-printable Content-Location: http://www.nature.com/nature/journal/vaop/ncurrent/full/nature04808.html IL-23 promotes = tumour incidence and growth : Nature
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Letter

Nature advance online publication 10 May 2006 | = doi:10.1038/nature04808; = Received 22=20 November 2005; Accepted 11 April 2006; Published online 10 May 2006

IL-23 promotes tumour incidence and growth

John L. Langowski1,2,=20 Xueqing Zhang1,2,=20 Lingling Wu1,=20 Jeanine D. Mattson1,=20 Taiying Chen1,=20 Kathy Smith1,=20 Beth Basham1,=20 Terrill McClanahan1,=20 Robert A. Kastelein1=20 and Martin Oft1

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Abstract

Chronic inflammation has long been associated with = increased=20 incidence of malignancy and similarities in the regulatory mechanisms = have been=20 suggested for more than a century1.=20 Infiltration of innate immune cells, elevated activities of matrix=20 metalloproteases and increased angiogenesis and vasculature density are = a few=20 examples of the similarities between chronic and tumour-associated=20 inflammation2.=20 Conversely, the elimination of early malignant lesions by immune = surveillance,=20 which relies on the cytotoxic activity of tumour-infiltrating T cells or = intra-epithelial lymphocytes, is thought to be rate-limiting for the = risk to=20 develop cancer3.=20 Here we show a molecular connection between the rise in = tumour-associated=20 inflammation and a lack of tumour immune surveillance. Expression of the = heterodimeric cytokine interleukin (IL)-23, but not of its close = relative IL-12,=20 is increased in human tumours. Expression of these cytokines = antagonistically=20 regulates local inflammatory responses in the tumour microenvironment = and=20 infiltration of intra-epithelial lymphocytes. Whereas IL-12 promotes=20 infiltration of cytotoxic T cells, IL-23 promotes inflammatory responses = such as=20 upregulation of the matrix metalloprotease MMP9, and increases = angiogenesis but=20 reduces CD8 T-cell infiltration. Genetic deletion or antibody-mediated=20 elimination of IL-23 leads to increased infiltration of cytotoxic T = cells into=20 the transformed tissue, rendering a protective effect against chemically = induced=20 carcinogenesis. Finally, transplanted tumours are growth-restricted in = hosts=20 depleted for IL-23 or in IL-23-receptor-deficient mice. Although many = strategies=20 for immune therapy of cancer attempt to stimulate an immune response = against=20 solid tumours, infiltration of effector cells into the tumour tissue = often=20 appears to be a critical hurdle4,=20 5,=20 6,=20 7.=20 We show that IL-23 is an important molecular link between = tumour-promoting=20 pro-inflammatory processes and the failure of the adaptive immune = surveillance=20 to infiltrate tumours.

IL-12 and IL-23 are members of a small family of = pro-inflammatory=20 heterodimeric cytokines8.=20 Both cytokines share a common p40 subunit that is covalently linked = either to a=20 p35 subunit to form IL-12 or to a p19 subunit to form IL-23 (ref. 9).=20 The receptor for IL-12 is comprised of an IL-12R3Dbeta=201 and IL-12R3Dbeta=202 subunit, whereas the receptor for IL-23 is comprised of the = IL-12R3Dbeta=201 subunit and a novel component termed IL-23R10.=20 Both cytokines are expressed predominantly by activated dendritic cells = and=20 phagocytic cells. Receptors for both cytokines are expressed on T cells, = natural=20 killer cells and natural killer T cells, but low levels of IL-23 = receptor=20 complexes are also found on monocytes, macrophages and dendritic-cell=20 populations10.=20 Despite these similarities, there is increasing evidence that IL-12 and = IL-23=20 drive divergent immunological pathways. Whereas IL-12 leads to = development of=20 'classical' interferon-3Dgamma=20-producing Th1 cells and enhances cytotoxic, antimicrobial = and=20 antitumour responses, IL-23 drives a pathway that leads to the = generation of=20 IL-17-producing CD4+ T cells11,=20 12,=20 13.=20 The induction of IL-23-derived processes leads to recruitment of a range = of=20 inflammatory cells as well as THIL-17 T cells, and has been = shown to=20 be crucial to the pathogenesis of a number of immune-mediated = inflammatory=20 diseases14.

Given the combined evidence of a causal relationship = between=20 chronic inflammation and cancer as well as the pivotal role IL-23 plays = in=20 autoimmunity, we hypothesized that IL-23-mediated responses may be = important in=20 tumour promotion. To test this, we employed a comprehensive examination = of=20 messenger RNA expression within various human cancer panels. = Surprisingly,=20 IL-23p19 mRNA was found significantly upregulated in the overwhelming = majority=20 of carcinoma samples from various organ types when compared with their = adjacent=20 normal tissue (Fig.=20 1a, b).=20 Significant mRNA upregulation was observed for both subunits of IL-23 = (Fig.=20 1c), but not for the IL-12-specific subunit p35 (Supplementary=20 Fig. S2). The expression of IL-17, a cytokine central to = tumour-promoting=20 angiogenesis15,=20 was also found to be significantly elevated in human tumours, consistent = with=20 activation of IL-23-induced processes (Supplementary=20 Fig. S2 and data not shown). Additionally, human colorectal tumours = present=20 a high degree of correlation between increased IL-23p19 expression and = increased=20 macrophage and neutrophil infiltration activation16=20 (Supplementary=20 Fig. S3). Expression of IL-23p19 protein was localized within human = tumour=20 tissue but not in the surrounding stroma, and observed on infiltrating=20 CD11c+ dendritic cells as a dominant source of the cytokine = (Supplementary=20 Fig. S4a, b). Supporting this, tumour-infiltrating macrophages=20 (CD11b+) and dendritic cells (CD11c+) isolated = from mouse=20 syngeneic tumour models expressed preferentially high amounts of = IL-23p19 mRNA=20 rather than IL-12p35 when compared to those isolated from the spleen (Supplementary=20 Fig. S4c, d).

Figure=20 1: Overexpression of IL-23 but not of IL-12 in human=20 cancer.

a, Quantitative mRNA expression of IL-23p19 in = human colon=20 tumour (red) compared to normal adjacent tissue of the same individual = (green,=20 connected by a line), or tissue from cancer-free individuals (black, = yellow).=20 b, Significant upregulation of IL-23p19 expression in various = human=20 cancers. The table shows number of individual paired cancerous and = normal=20 adjacent samples with average fold increase in expression, number of = samples=20 with five- and ten-fold increase, and P-value. c, = Expression of=20 both IL-23 subunits is increased in human cancer; each y-axis = position=20 signifies expression within an individual sample. Histological = classification of=20 samples as normal, normal adjacent and cancerous (with stage = classification) is=20 given.

High=20 resolution image and legend (106K)

To examine more directly the role of IL-12 and IL-23 in = epithelial=20 tumorigenesis, we employed a strategy of genetic deletion, analysing the = susceptibility of mice deficient in either IL-12 or IL-23=20 (p35-/- or p19-/- mice) or mice = deficient in=20 both cytokines (p40-/- mice) to tumour formation = during=20 chemical carcinogenesis17.=20 Whereas Il12p35-/- mice showed earlier appearance and=20 developed significantly increased numbers of papillomas compared to = C57BL/6=20 control mice (Fig.=20 2a, b),=20 mice deficient in IL-23p19 were resistant to tumour induction. = Surprisingly,=20 p40-/- mice, deficient in both cytokines, were = resistant to=20 tumour induction, emphasizing the importance of IL-23 for increased = tumour=20 incidence. Papillomas generated by this protocol persisted well over=20 100 days after cessation of TPA promotion (see Methods) and often=20 progressed towards malignancy (Fig.=20 2b and data not shown); however, the low overall incidence of benign = tumours=20 in the absence of IL-23 prevents a statistically relevant comparison = with WT=20 mice in this regard.

Figure=20 2: Tumour resistance with reduced inflammation in = IL-23-deficient=20 animals.

a, b, IL-23-deficient mice are resistant, = whereas=20 IL-12-deficient mice are more susceptible, to cutaneous chemical = carcinogenesis.=20 Papillomas persisted more than 100 days following cessation of = treatment=20 (b, right panel). c, Cutaneous IL-17 and G-CSF mRNA = expression=20 following 140 days of carcinogenesis. d, Increased = granulocyte=20 infiltration in the dermis of WT and Il12p35-/- mice.=20 e=96l, Cutaneous expression of activated MMP9 = (e=96h)=20 and the endothelial marker CD31 (i=96l) following = 140 days of=20 carcinogenesis. We note increased angiogenesis in C57BL/6 and=20 Il12p35-/- mice. Data are mean 3Dplusminus=20 s.d.; scatter plots show incidence with bars = depicting=20 the mean. The significance of results in the figures is represented by:=20 *P < 0.05, **P < 0.01 and ***P < 0.001; = n=20 7 (a, b), n =3D 3 = (c, d)=20 per group.

High=20 resolution image and legend (221K)

To explore the role of IL-23 in tumour susceptibility, = we examined=20 the expression of inflammatory immune cell products in = carcinogen-treated skin.=20 IL-23 is known to modulate the homeostasis of neutrophil infiltration = into=20 tissues by inducing expression of IL-17 and G-CSF18=97two=20 cytokines that are also linked to tumour growth15,=20 19.=20 We found IL-17 highly expressed in the hyperplastic skin of both = wild-type=20 C57BL/6 (WT) and Il12p35-/- mice, but barely = detectable in=20 Il23p19-/- and Il12p40-/- mice. = Similarly,=20 G-CSF levels were decreased in Il23p19-/- mice (Fig.=20 2c). The number of granulocytes (GR1+) and macrophages=20 (CD11b+, F4/80+) in the dermis of=20 Il23p19-/- and Il12p40-/- mice was = also=20 decreased compared to the skin of C57BL/6 and = Il12p35-/- mice=20 (Fig.=20 2d, Supplementary=20 Fig. S5a, b and data not shown). In murine cancer models, tumour = development=20 is dependent on MMP9 activity produced by inflammatory macrophages and = mast=20 cells20.=20 Additionally, MMP9 and IL-17 have been identified as both markers and = mediators=20 of tumour angiogenesis15,=20 21.=20 The expression of activated MMP9 was elevated in carcinogen-treated skin = of=20 tumour-prone WT and Il12p35-/- animals, but = dramatically lower=20 in skins of Il23p19-/- and = Il12p40-/- mice=20 (Fig.=20 2e=96h, Supplementary=20 Fig. S5d, e). Angiogenic markers also followed this pattern, with=20 significantly lower vessel density in IL-23p19-deficient animals (Fig.=20 2i=96l, Supplementary=20 Fig. S5c). Thus, the absence of IL-23 resulted in a significant = reduction in=20 numerous inflammatory factors and cell types essential for tumour = promotion.

Because the incidence of macroscopic tumours can be = viewed as the=20 result of a disturbed balance between tumour-promoting factors and=20 tumour-cell-eliminating mechanisms, our observations raised the question = of=20 whether endogenous IL-23 would have a direct role in immune surveillance = against=20 tumours. Immune surveillance and anti-tumour responses are vigorously = promoted=20 by IL-12, which induces proliferation and cytotoxic activity of natural = killer=20 cells and CD8+ T cells22.=20 Surprisingly, the carcinogen-induced hyperplastic skin of=20 Il23p19-/- and Il12p40-/- mice were=20 infiltrated with a dramatically increased number of cytotoxic = CD8+ T=20 cells in both the dermis and the epidermis when compared to WT and=20 p35-/- mice (Fig.=20 3a=96e). The expression of markers associated with the activity of = cytotoxic T=20 cells such as Fas ligand, perforin and granzymes were decreased in=20 Il12p35-/- animals but increased in=20 Il23p19-/- mice (Supplementary=20 Fig. S6 and data not shown). Although deficiency of IL-12 or IL-23 = does not=20 alter the development of CD8 cells (ref. 13=20 and data not shown), it may alter the response necessary for tissue=20 infiltration. To test this, we changed the microenvironment of the=20 carcinogen-treated skin of C57BL/6 mice by serial intradermal injection = of IL-12=20 and IL-23. While administration of IL-12 led to increased immune = surveillance,=20 as shown by the influx of greater numbers of CD8+ T cells in = the=20 epidermis, IL-23 injection caused a significant decrease of CD8 T cells = in the=20 hyperplastic skin (Fig.=20 3f, Supplementary=20 Fig. S7a=96c). Cytotoxic activity markers such as perforin, = granzyme, Fas=20 ligand and interferon-3Dgamma=20 increased on IL-12 administration, whereas IL-23 greatly = raised mRNA=20 levels of IL-17 (Supplementary=20 Fig. S7d, e). Systemic treatment with neutralizing antibodies = specific to=20 IL-23p19 led to a marked down-modulation of MMP9 expression and = increased=20 surveillance of CD8+ T cells within the carcinogen-treated = skin (Fig.=20 3g=96h, Supplementary=20 Fig. S8). Taken together, these results provide evidence that IL-23=20 negatively influences both the number and activity of cytotoxic T cells = within=20 transformed tissue.

Figure=20 3: IL-12 stimulates, but IL-23 decreases, immune surveillance by = CD8 T=20 cells.

a=96e, Increased immune surveillance by CD8 = T cells in=20 the epidermis of Il23p19-/- mice (c) and=20 Il12p40-/- mice (d) treated with carcinogen for = 140 days, compared to WT (a) and p35-/-=20 (b). Quantification of infiltrating CD8 T cells for each genotype = is=20 shown in e. f, Alterations in epithelial CD8 infiltration=20 following treatment with exogenous IL-23 or IL-12 compared to = control-treated=20 animals. g, h, Systemic treatment with anti-IL-23p19 = antibody=20 shows marked decrease in MMP9 expression (g), but increase in=20 CD8+ T cells (h), compared to control. Scatter plots = show=20 incidence or density with bars depicting the mean. Significance of = results in=20 figures is represented by: *P < 0.05 and ***P < = 0.001;=20 n =3D 3 per group.

High=20 resolution image and legend (96K)

To further investigate the effects of IL-12 and IL-23 on = immune=20 surveillance, IL-12- and IL-23-cytokine-deficient mice were challenged=20 intradermally with PDV squamous carcinoma cells17.=20 In accordance with the dominant role of IL-12 in tumour immunity, = deficiency of=20 IL-12p35 or IL-12p40 dramatically increased tumour incidence, indicating = the=20 diminished ability of the host to reject the nascent carcinomas (Fig.=20 4a). IL-23p19-deficient mice, however, had a strong resistance to = tumour=20 challenge, similar to that of C57BL/6 control mice. Although previous = reports=20 suggest genetically engineered tumours overexpressing an artificially = linked=20 IL-23p19 and p40 construct are rejected more efficiently23,=20 our results show that at the very least IL-23 deficiency does not impair = the=20 ability of the immune system to eliminate nascent tumours.

Figure=20 4: Functional ablation of IL-23p19 signalling restricts tumour=20 growth.

a, Persistence of immune-sensitive PDV squamous = carcinoma in=20 IL-12-deficient but not WT or p19-/- mice. b, = B16F10=20 melanoma and c, LL/2 lung carcinoma cells exhibit reduced growth = in=20 Il23R-/- mice. d, Tumour growth of = immune-resistant=20 PDV6 squamous carcinoma cells in IL-12- or IL-23-receptor-deficient = mice.=20 e, Growth restriction of PDV6 tumours is CD8-dependent only in=20 Il23R-/- mice. f, Antibody ablation of IL-12p40 = increases both tumour incidence and size compared to anti-IL-23p19 = treatment.=20 g, EP2 breast tumour growth is increased by ablation of IL-12p40 = or=20 reduced by anti-IL-23p19 treatment (***/* for days 18 and 21). Data are = mean=20 3Dplusminus=20 s.d.; scatter plots show incidence with bars = depicting=20 the mean. Significance of results in figures is represented by: = *P <=20 0.05 and ***P < 0.001; NS, not significant; n 5 (a=96e, g), n = 14 (f) mice per group with two tumours = per mouse=20 for all studies.

High=20 resolution image and legend (73K)

To explore the respective importance of the host's = immune response=20 to IL-12 and IL-23 and to control tumour growth, we challenged mice = incapable of=20 responding to IL-23 (Il23R-/- mice) with subcutaneous=20 transplantation of murine tumour cell lines. The growth of B16F10 = melanoma and=20 LL2 lung carcinoma was inhibited in Il23R-/- mice = incapable of=20 responding to IL-23, compared to either WT or mice incapable of = responding to=20 IL-12 (Fig.=20 4b, c=20 and data not shown). We then used an immune-resistant subclone of the = PDV=20 squamous carcinoma (termed PDV6), to investigate the role of IL-12 and = IL-23=20 signalling in the host on tumour growth and CD8 T-cell function. Growth = of=20 squamous carcinomas in Il23R-/- was reduced compared = to that=20 in WT mice, but increased in mice not able to respond to IL-12 (Fig.=20 4d). Although depletion of CD8 T cells in WT mice did not alter PDV6 = tumour=20 growth kinetics, it potentiated tumour growth in = Il23R-/- mice=20 (Fig.=20 4e). These results demonstrate that the absence of IL-23 signalling=20 attenuates tumour growth and suggests an essential role for CD8 T cells = in=20 restricting tumour growth in Il23R-/- mice.

We next asked whether antibody ablation of IL-23 or = combined=20 ablation of IL-23 and IL-12 would have a different effect on the ability = of the=20 host immune system to eliminate early malignant tumour cells. Animals = treated=20 with anti-IL-23p19 showed a decreased risk of PDV tumour formation and = faster=20 elimination of injected tumour cells. However, animals treated with=20 anti-IL-12p40, and thus depleted of both IL-12 and IL-23, had a = significantly=20 increased tumour incidence and developed larger, faster-growing tumours = (Fig.=20 4f and data not shown). Finally, animals bearing tumours formed by a = transformed mammary cell line were treated with IL-23 or IL-12 = neutralizing=20 antibodies. Mice treated with anti-IL-23p19 showed decreased tumour = growth and=20 increased tumour rejection. However, anti-IL-12p40 treatment led to a = marked=20 increase of tumour growth and an increase of metastasis formation (Fig.=20 4g and data not shown). Thus, although elimination of transformed = cells is=20 regulated by IL-12 (ref. 22),=20 the contribution of the pro-inflammatory environment to tumour = propagation can=20 be seen in the result of functional IL-23 ablation.

The importance of inflammatory processes in tumour = progression and=20 initiation is exemplified by the tumour resistance of MMP9- or = COX-2-deficient=20 mice20,=20 24.=20 However, local expression of growth factors and cytokines in tumours may = not=20 only aid tissue remodelling and angiogenesis, but also protect the = tumour from=20 immune-mediated elimination7.=20 Surprisingly, our work shows that IL-23 appears not only to induce = innate=20 inflammatory infiltration, but also to downregulate dramatically the = ability of=20 CD8 T cells to infiltrate tumours (Supplementary=20 Fig. S1). Although the understanding of the biological role of IL-23 = is=20 still evolving, it is known that IL-23 production by dendritic cells and = macrophages can be elicited by bacterial exposure and Toll-like receptor = engagement. The resulting IL-17-dependent immune response constitutes a=20 significant component for rapid neutralization of infectious = organisms9,=20 25,=20 26.=20 However, inappropriate expression of IL-23 also appears to coincide with = numerous autoimmune disorders14,=20 27.=20 IL-23-induced inflammatory processes such as induction of angiogenesis = or=20 macrophage and neutrophil infiltration are valuable defence mechanisms = against=20 bacterial infections. However, this regulatory pathway appears to be not = just=20 inappropriate for eliminating tumours but instead could provide a = protective,=20 tumour-promoting environment for nascent malignancies, suggesting that=20 anti-IL-23p19 therapy may prove efficacious for tumour treatment.

Methods

Animal models

C57BL/6 mice and=20 Il12p35-/- (Il12atm1Jm), = Il12p40-/-(Il12btm1Jm), = Il12R1-/- (Il12rb1tm1Jm), and=20 IL12R3Dbeta=202-/- (Il12rb2tm1Jm) mice=20 on the C57BL/6 background were obtained from Jackson Laboratory.=20 Il23R-/- (Il23ratm1dnax) = were=20 maintained on a C57BL/6 background;=20 Il23p19-/- (Il23atm1dnax)= were=20 produced by DNAX13=20 and made congenic on the C57BL/6 background (see Supplementary=20 Methods). The two-step skin carcinogenesis model was adapted from = ref. 28:=20 mice were treated with 9,10-dimethyl-1,2-benzanthracene (DMBA;=20 Sigma) in 200 3Dmicro=20l acetone at 100 3Dmicro=20g per mouse once at the age of 2=963 months, then treated = twice-weekly=20 with the tumour promoter 12-O-tetradecanoyl-phorbol acetate (TPA;=20 Fisher) in 200 3Dmicro=20l acetone, 30 3Dmicro=20g per mouse for up to 1 year. Tumours observed arise as = papillomas=20 (keratoacanthomas) but statistically progressed towards carcinomas and=20 metastasized via the lymph drainage. Papilloma counts were conducted = routinely=20 by visual examination. All experiments were repeated at least two times = with=20 similar significance values (P =3D 0.0106 and P < = 0.001,=20 respectively, n 7 for each group in each experiment, with a = total of=20 n 21 per genotype). Mice 8=9610 weeks old were = exposed to=20 carcinogen for five months=97they were either injected subcutaneously = with murine=20 IL-12, IL-23 or PBS at 20 ng per injection site or, for antibody = ablation,=20 systemically injected with 100 3Dmicro=20g of anti-IL-23p19 antibody or isotype control. In both = cases, skin was=20 harvested 72 h later for immunohistochemistry or for TAQMAN gene = expression=20 analysis.

In tumour transplantation studies, transformed cell = lines were=20 mixed 1:1 with growth-factor-reduced Matrigel=20 (Gibco) and injected subcutaneously or intradermally, typically in two = separate=20 sites per animal. Squamous carcinoma cells (PDV29),=20 immune resistant PDV derivative (PDV6;=20 DNAX), melanoma (B16F10;=20 ATCC) and lung (LL2;=20 ATCC) cell lines were injected into C57BL/6 mice, while breast (EP2;=20 DNAX and 4T1;=20 ATCC) and colon (CT26;=20 ATCC) cancer cells were injected into BALB/c mice. For persistence of = PDV=20 transplants, tumour growth was followed for an observation period of=20 2=966 months. Anti-cytokine antibody injections (1 mg per = mouse) were=20 repeated weekly, and expected serum concentrations were confirmed after = one week=20 and at the end of the experiment. For anti-CD8=20 antibody ablation, 0.25 3Dmicro=20g per mouse of antibody (DNAX) was injected subcutaneously = once per=20 week; ablation effectiveness was judged by flow cytometry of treated = spleens at=20 the experimental endpoint. Tumour volume was determined by electronic = calliper=20 measure: (half-length)2 3Dtimes=20 width.

Antibodies, cytokines, immunohistochemistry, = zymogram

Recombinant murine=20 IL-12 was obtained from R&D, while rmIL-23=20 was produced at DNAX as previously described9.=20 For the ablation of IL-12, monoclonal antibody C17.8 (ref. 30)=20 was used. Mouse anti-murine=20 IL-23p19 antibodies were developed at DNAX. Antibodies were injected = subcutaneously in equal PBS volume at 1 mg per mouse per week. = Expected=20 serum concentrations of C17.8 and anti-IL-23p19 were confirmed by = immunoassays=20 (ELISAs) specific for each antibody. Mouse skin biopsies were harvested = from=20 macroscopically tumour-free areas of carcinogen-exposed skin, = snap-frozen, fixed=20 and subjected to immunohistochemistry and counterstained with = haematoxylin.=20 Antibodies used include CD3,=20 CD8, CD11b, CD11c, CD31 (BD Biosciences), active MMP9, perforin and = F4/80=20 (R&D). Density of staining was determined by analysing multiple 20 = fields with ImagePro=20 Plus software (MediaCybernetics). Gelatin zymogram=20 gels (Invitrogen) were performed following the supplier's=20 instructions.

Gene expression analysis

For TAQMAN analysis, total RNA was isolated by = standard=20 methodologies and reverse transcribed. Expression analysis for = marker-specific=20 mRNA was measured using real-time quantitative PCR (ABI 5700) with SYBR=20 Green PCR Mastermix (Applied Biosystems). Ubiquitin levels were = measured in=20 a separate reaction and used to normalize the data by the 3DDelta=20 - 3DDelta=20Ct method, using the mean cycle threshold=20 (Ct) value for ubiquitin and the gene of interest for = each=20 sample; the expression=20 1.8Ct(ubiquitin)-Ct(gene of = interest)=20 3Dtimes=20 104 was used to obtain the = normalized values=20 (see Supplementary=20 Methods). For murine samples, punch biopsies were taken from at = least three=20 independent, macroscopically and histologically tumour-free areas for = each=20 condition.

Tumour-infiltrating immune cell isolation

Subcutaneous 4T1 and CT26 tumour masses were = dissected and=20 digested with 2 mg ml-1 collagenase and=20 0.5 mg ml-1 DNase=20 (Sigma). Single-cell suspensions were created and live cells collected = by=20 centrifugation over a Histopaque=20 (Sigma) gradient. Similar isolations from naive spleen were performed = for=20 comparison. Collected cells were positively selected for CD11b or CD11c = by magnetic=20 selection (Miltenyi Biotec) according to the manufacturer's=20 specifications.

Statistical analyses

The following statistical analyses were used. For = data in Fig.=20 1 and Supplementary=20 Fig. S1, we used the Mann=96Whitney test (non-paired = Wilcox)=97nonparametric,=20 group comparison. In Figs=20 2, 3,=20 4a=96c=20 and Supplementary=20 Figs S3 and S4, we used the Student's t-test. In Fig.=20 4a=96c, e=96g,=20 we used the Kruskal=96Wallis test followed by Dunn's multiple comparison = test=20 (non-parametric), or analysis of variance (ANOVA) (parametric) followed = by=20 Tukey's multiple comparison test. In Fig.=20 4d, we used Wilcoxon matched pairs test. All tests (other than in Supplementary=20 Fig. S2) were performed using Prism=20 4 (GraphPad Software). Linear regression analysis (Supplementary=20 Fig. S2) was performed with Spotfire DecisionSite=20 software.

Human tumour panels, immunohistochemistry

Paired human tumour and normal adjacent tissues = were=20 obtained from patients undergoing therapeutic or routine surgery, = respectively=20 (see Supplementary=20 Methods). All sample diagnoses and staging were confirmed internally = by a=20 pathologist. Immunohistochemistry was performed on frozen sections with = murine=20 anti-human IL-23p19 (DNAX) with biotinylated=20 horse anti-mouse IgG (Vector Laboratories).

Acknowledgments

We thank C. Strader for her comments on the manuscript. = Author=20 Contributions J.L.L. and X.Z. contributed equally to this work.

Competing interests statement:

The authors declared no competing interests.

Supplementary=20 information accompanies this paper.

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  1. Schering-Plough BioPharma (formerly DNAX Research, Inc.), = 901=20 California Avenue, Palo Alto, California 94304, USA
  2. *These authors contributed equally to this work

Correspondence to: Martin Oft1=20 Correspondence and requests for materials should be addressed to M.O.=20 (Email: martin.oft@spcorp.com).

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RESEARCH
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