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Are Mast Cells MASTers in Cancer?

Gilda Varricchi, Maria Rosaria Galdiero, Stefania Loffredo and 4 more

Frontiers in Immunology · 2017 · Review

doi:10.3389/fimmu.2017.00424

Abstract

Prolonged low-grade inflammation or smoldering inflammation is a hallmark of cancer. Mast cells form a heterogeneous population of immune cells with differences in their ultra-structure, morphology, mediator content, and surface receptors. Mast cells are widely distributed throughout all tissues and are stromal components of the inflammatory microenvironment that modulates tumor initiation and development. Although canonically associated with allergic disorders, mast cells are a major source of pro-tumorigenic (e.g., angiogenic and lymphangiogenic factors) and antitumorigenic molecules (e.g., TNF-α and IL-9), depending on the milieu. In certain neoplasias (e.g., gastric, thyroid and Hodgkin's lymphoma) mast cells play a pro-tumorigenic role, in others (e.g., breast cancer) a protective role, whereas in yet others they are apparently innocent bystanders. These seemingly conflicting results suggest that the role of mast cells and their mediators could be cancer specific. The microlocalization (e.g., peritumoral vs intratumoral) of mast cells is another important aspect in the initiation/progression of solid and hematologic tumors. Increasing evidence in certain experimental models indicates that targeting mast cells and/or their mediators represent a potential therapeutic target in cancer. Thus, mast cells deserve focused consideration also as therapeutic targets in different types of tumors. There are many unanswered questions that should be addressed before we understand whether mast cells are an ally, adversary, or innocent bystanders in human cancers.

Full text

From PubMed Central, U.S. National Library of Medicine, as current on Oct 18, 2026. Paragraph numbers are ours, so an answer can point at one.

Introduction

Mast cells were first identified in human tumors and named by Paul Ehrlich. These cells are present in all classes of vertebrates, and it has been estimated that they have emerged >500 million years ago, long before the development of adaptive immunity. Mast cells are distributed throughout nearly all human tissues and often in close proximity to epithelia, fibroblasts, blood and lymphatic vessels, and nerves.

Human mast cells form a heterogeneous population of cells with differences in their ultrastructure, morphology, mediator content, and surface receptors. Human mast cells derive from CD34+, CD117+ pluripotent hematopoietic stem cells, which arise in the bone marrow. Mast cell progenitors enter the circulation and subsequently complete their maturation in tissues. These cells store and release upon activation a wide spectrum of biologically active mediators that individually have been shown to have potential positive or negative effects on various target cells. Increasing evidence indicates that mast cells act as sentinels of the surrounding environment, with the capacity to rapidly perceive tissue insults and initiate biochemical programs of inflammation or repair.

Mast cells are activated not only by IgE, specific antigens, and superallergens, the main mechanisms which account for their function in allergic disorders, but also by a plethora of immunologic and non-immunologic stimuli. Figure 1 schematically illustrates the constellation of surface receptors expressed by human mast cells.

Mast cells and their mediators have been canonically associated with a detrimental role in allergic diseases, but these cells can induce a protective immune response of the host against noxious substances, viral and microbial pathogens. Interestingly, epidemiological and experimental studies indicate an inverse association between IgE-mediated allergies and cancer, implying tumor-protective effect of IgE.

The initiation and progression of cancer are multistep processes characterized by the accumulation of a variable number of genetic and epigenetic alterations. The immunosurveillance system recognizes and eliminates mutant cells constantly generated. However, immune-resistant cancer cells can slip through this system and proceed to develop tumors.

Normal microenvironment [immune cells, fibroblasts, blood and lymphatic vessels, and interstitial extracellular matrix (ECM)] plays a central role in maintaining tissue homeostasis and is a barrier to tumorigenesis. Incorrect signals (chemokines, cytokines, reactive oxygen species, lipid mediators, etc.) from an aberrant microenvironment alter tissue homeostasis and initiate/promote tumor growth. Thus, the multiple interactions between stromal and tumor cells are crucial for the initial phases of tumor development.

Prolonged low-grade inflammation or smoldering inflammation is a hallmark of cancer. Several cells of the innate and adaptive immune system (macrophages, mast cells, lymphocytes, neutrophils, NK, and NK T cells) are stromal components of the inflammatory microenvironment that can promote the development of experimental and human tumors.

Why are Mast Cells Increased in Tumors?

The presence of mast cells in human tumors, initially reported by Ehrlich, was extended by Eugen Westphal. Tumor-associated mast cells (TAMCs) are present in the microenvironment of several human solid and hematologic tumors.

Peritumoral and/or intratumoral mast cell density is increased in different types of human cancer. Tumor cells produce several chemotactic factors acting on receptors expressed by mast cells. Stem cell factor (SCF), also produced by mast cells, activates the mast cell Kit receptor (CD117), vascular endothelial growth factors (VEGFs) act on VEGFR-1 and VEGFR-2, angiopoietin 1 (Ang1) acts on Tie2 receptor, and CXCL8/IL-8 acts on CXCR1 and CXCR2. Mast cells express CCR2, CXCR2, and CXCR3, which can be important for TAMC localization because their respective ligands, CCL2, CXCL1, and CXCL10, are produced by human tumors. PGE2 and histamine are chemotactic for mature mast cells through the engagement of EP2 receptor and H4R, respectively. LTB4 may be involved in recruitment of mast cell progenitors from the circulation via the activation of BLT1 and BLT2. Finally osteopontin (OP), which is upregulated in human cancer, induces mast cell migration and degranulation.

The Contribution of Mast Cells to Cancer is Tumor Dependent

The increasing heterogeneity of different subsets of immune cells (e.g., macrophages, T helper cells, mast cells, neutrophils, NK, NK T cells, etc.), their plasticity, and their reciprocal interactions have complicated the comprehension of the role of the inflammatory microenvironment in tumor initiation and development.

A large number of studies have tried to identify the contributory functions of TAMCs in tumor growth. In the majority of studies, TAMCs appear functional—either actively promoting or suppressing tumor development and growth—whereas in a few cases they may be simple inert bystanders. In several studies, mast cells appear to play a pro-tumorigenic role in human (Table 1) and experimental tumors (Table 2). Evidence for an antitumorigenic role for mast cells is provided in Table 3. Studies supporting a non-contributing role of mast cells in tumors are outlined in Table 4.

In several solid tumors, such as thyroid, gastric, pancreas, bladder cancers, and Merkel cell carcinoma, mast cells always appear to be pro-tumorigenic. Similarly, in several hematologic tumors, such as different types of Hodgkin’s and non-Hodgkin’s lymphoma, and plasmacytoma, mast cells are associated with poor prognosis. There are certain tumors such as breast cancer in which mast cells always appear to play an antitumorigenic role. The role of mast cells in the pathogenesis of human melanomas is still unclear and appears to depend on both the microlocalization of these cells and the subtypes of tumor.

These apparently conflicting results are intriguing and suggest that the role of mast cells and their mediators in tumors could be cancer specific. Figure 2 schematically illustrates the role of mast cells in different human tumors.

Role of TAMCs in Tumor Angiogenesis and Lymphangiogenesis

Angiogenesis, the formation of new blood vessels, is an essential process for supplying growing malignant tissues with essential nutrients and oxygen. Lymphangiogenesis, the formation of new lymphatic vessels, is important in the development of metastases. Judah Folkman, the father of angiogenesis, suggested that mast cells and macrophages could be attracted by chemotactic molecules produced by tumor cells and could be an important source of proangiogenic factors. Several groups have demonstrated that mast cells produce several proangiogenic (VEGF-A, VEGF-B, and FGF-2) and lymphangiogenic factors (VEGF-C and -D). In addition, we have found that VEGFs are chemotactic for mast cells, indicating that mast cells are a target, in addition to be a source, for VEGFs. Several studies have highlighted the association and/or the correlation in human tumors between increased mast cell density and angiogenesis by evaluating the expression of the proangiogenic isoform VEGF-A.

The VEGF-A gene can be alternatively spliced to form the proangiogenic VEGF-A165 and the antiangiogenic VEGF-A165b. The vast majority of the studies performed so far evaluated only the proangiogenic isoforms, whereas in certain tumors the antiangiogenic VEGF-A165b isoform is dominant. This finding suggests that the majority of results on VEGF-A plasma levels in cancer need to be reinterpreted or require repeating with tools that can differentiate between the two isoforms of VEGF-A. For instance, we have recently demonstrated that human neutrophils, under certain circumstances, can produce both pro- and antiangiogenic isoforms of VEGF-A. The role of different pro- and antiangiogenic isoforms of VEGFs produced by TAMCs in primary cancers and in the formation of metastases needs further investigation.

Human mast cells produce different matrix metalloproteinases (e.g., MMP-9) and proteases (tryptase and chymase), which regulate the digestion of ECM favoring the implantation of cancer cells in an aberrant microenvironment.

Vascular endothelial growth factor-C, released by melanoma cells, TAMs, and TAMCs, likely represents a major lymphangiogenic factor in this tumor. Mast cells can be found in metastatic lymph nodes of cancer patients, and the role of lymphangiogenic factors in the formation of metastasis should be further addressed.

Epithelial-to-mesenchymal transition (EMT) is a mechanism by which tumor cells gain metastatic features and contribute to chemotherapy drug resistance. In addition, in the context of tumors, EMT can generate cells with stem-like properties (e.g., stemness). We have demonstrated that mast cells can induce EMT and stem cell features in human cancer through the production of CXCL8/IL-8.

The Role of Mast Cells Varies According to the Stage of Tumors

A recent study found that low mast cell count in perilesional stroma of deeply invasive melanomas predicted poor survival. By contrast, mast cell density was not correlated with prognosis in superficially invasive melanomas. The latter findings suggest that the role of mast cells in melanoma is dependent also on the stage of the tumor. The role(s) of these cells in human and experimental melanoma requires additional studies.

Pittoni et al. found that in prostate cancer mast cells exert different functions according to tumor stage. Mast cells were pro-tumorigenic in the initial stages of prostate cancer by supplying MMP-9 in the microenvironment, but became dispensable at later stages.

In stage I non-small-cell lung cancer (NSCLC), but not in stage II, peritumoral but not intratumoral mast cell (tryptase+ chymase+) density was an independent favorable prognostic factor.

Vascular endothelial growth factor-B, an angiogenic factor produced by human macrophages and mast cells, could play a role in early colon cancer development at the stage of adenoma formation.

The Role of Mast Cells in Tumors Varies According to Their Microlocalization

The vast majority of initial studies evaluating mast cell density in different cancers did not examine differences between the periphery and the center of tumors. There is increasing evidence that different stages of tumors can be associated with qualitative and quantitative changes in different types of immune cells in the periphery and center of tumors. The pro- or antitumorigenic role of mast cells in different types of melanomas remains controversial. Siiskonen and collaborators found that tryptase+ chymase+ mast cells in perilesional stroma of melanoma play a protective role. In NSCLC, mast cell infiltration of tumor islets confers a survival advantage independently of tumor stage. In another study, it was found that only in stage I NSCLC increased peritumoral mast cells were associated with a better prognosis. In pancreatic ductal adenocarcinoma, mast cell density in the intratumoral border zone, but not the peritumoral or the intratumoral center zone, was associated with a worse prognosis. In prostate cancer, high intratumoral mast cell density was initially associated with good prognosis. Subsequently, it was reported that intratumoral mast cells inhibited tumor growth, whereas peritumoral mast cells stimulated human prostate cancer.

Mast cells are increased in patients with both cutaneous T-cell lymphoma and cutaneous B-cell lymphoma compared with normal skin, particularly at the periphery of the tumors. Interestingly, the density of mast cells in the center of tumors was similar to normal skin. The density of peripheral mast cells correlated with disease progression.

Collectively, these findings suggest that the microlocalization of mast cells is an important aspect in the initiation and progression of several tumors.

Figure 3 schematically illustrates the mechanisms by which mast cells and some of their mediators may play a pro-tumorigenic or an antitumorigenic role.

Which are the Activators of TAMCs in Tumor Microenvironments?

Peritumoral and intratumoral mast cells operate in an inflammatory microenvironment characterized by hypoxia, the accumulation of lactic acid, adenosine, PGE2, IFN-γ, and by low pH. This milieu is likely to influence mast cell recruitment and activation. Mast cells can be recruited by SCF produced by several tumors and by mast cells themselves. Mast cells can be recruited by VEGFs and Ang1 produced by tumor and immune cells through the engagement of VEGFR-1/VEGFR-2 and Tie2, respectively, expressed by human mast cells.

Hypoxia, a feature of tumor microenvironment, activates human mast cells to release IL-6 and VEGF-A. Adenosine, produced by tumor cells and mast cells, is markedly increased and is an immunosuppressive factor in tumor microenvironment. Adenosine potentiates histamine release and the production of angiogenic factors from human mast cells and macrophages. Cyclooxygenase 2, overexpressed in tumors, generates PGE2 which induces angiogenic and lymphangiogenic factors from human mast cells. Several chemokines (CXCL1, CXCL10, and CXCL12) can activate mast cells and enhance mast cell secretion of CXCL8/IL-8. Thus, these chemokines can promote angiogenesis/lymphangiogenesis via the recruitment of mast cells to the edge of solid tumors.

The impact of IgE-mediated activation of mast cells on tumor development and progression has been investigated. Monomeric IgE, in the absence of antigen, induced VEGF-A production from mast cells and increased melanoma growth. Increased expression of immunoglobulin free light chains (FLC) was found within stroma of various human cancers. In a murine B16F10 melanoma model, inhibition of FLC-mediated mast cell activation reduced tumor growth. Alarmins are upregulated in cancers and can activate mast cells. IL-33 is upregulated in squamous cell carcinoma (SCC), and mast cell activation by IL-33 occurs in skin cancers. IL-33 induces the production of GM-CSF, CXCL8/IL-8, and VEGF-A from mast cells. In addition to the high-affinity receptor for IgE (FcεRI), human mast cells express the IgG receptors FcγRIIA and FcγRI. FcγRI is upregulated by IFN-γ which is highly expressed in tumors. In the tumor microenvironment, antitumor IgG immune complexes may activate mast cells. OP, upregulated in human cancer, is produced by mast cells and induces their migration and degranulation. Platelet-activating factor, produced by human mast cells, upregulates CXCR4 on mast cells and promotes their migration to lymph nodes.

In summary, a plethora of immunologic and non-immunologic factors present in tumor microenvironment can activate TAMCs.

Mast Cells as a Potential Therapeutic Target in Cancer

Several therapeutic strategies have been envisioned to limit tumor growth by targeting mast cells and their mediators. Mast cells play a pro-tumorigenic role in human bladder cancer through stimulating estrogen receptor β (ERβ). In a murine model of bladder cancer, these authors showed that a selective ERβ antagonist inhibited mast cell-promoted tumor growth. It has been found that mast cells can promote the proliferation of colon cancer in vivo. Injection of Fcε-PE40 chimeric toxin, which induced mast cell apoptosis, inhibited colon tumor development in vivo.

Pharmacologic inhibition of mast cell degranulation by cromolyn inhibited Myc-induced pancreatic islet tumors, experimental pancreatic and thyroid cancer, and cholangiocarcinoma.

Pittoni and collaborators have demonstrated that pharmacologic inhibition by cromolyn and genetic ablation of mast cells inhibited prostate cancer in mice. However, mast cells protect from a malignant neuroendocrine tumor. It has been shown that mast cells can promote prostate cancer chemotherapy and radiotherapy resistance via modulation of p38/p53/p21. The authors suggested that targeting these signaling pathways may help to suppress chemo- and radiotherapy resistance in prostate cancer. In a mouse model, mast cells enhanced prostate cancer growth via modulation of androgen receptor and increasing MMP-9 expression. The authors suggested that targeting these mast cell-androgen receptor signals may inhibit tumor growth.

The UV wavelengths in sunlight are the prime etiological cause of skin cancers, including basal cell carcinoma and SCC. Exposure to UV affects skin mast cell migration by altering the CXCR4–CXCL12 axis. The pharmacological blockage of the CXCR4–CXCL12 pathway inhibited sunlight-induced skin cancer.

Collectively, these findings indicate that mast cells and their mediators deserve focused consideration as therapeutic targets in different types of cancer.

Outstanding Questions

There is compelling evidence that human mast cells isolated from various anatomical sites respond to different stimuli and release distinct mediators. Peritumoral and intratumoral TAMCs are embedded by a wide spectrum of mediators and in close contact with several stromal cells. It will be important to identify the stimuli that can activate TAMCs in different tumor microenvironments. Similarly, it will be important to identify preformed and de novo synthesized mediators released in situ by TAMCs.

Studies on mast cell biology are routinely conducted at physiological pH and normoxia. By contrast, the metabolic phenotype of tumors is characterized by low pH and areas of either hypoxia or normoxia. Tumor-associated macrophages in normoxic tumor tissues express M1 markers, whereas those in hypoxic tumor tissues preferentially express M2 markers. These findings caution against the over interpretation of results from studies of whole TAMC populations. It will be of fundamental importance to investigate how hypoxic conditions and metabolism activate/modulate the production of pro-inflammatory and angiogenic/lymphangiogenic factors from TAMCs. Proteomic and lipidomic analyses of mast cells will help to characterize the proangiogenic and antitumorigenic profiles of TAMCs from different human tumors.

Analysis of mast cells in draining lymph nodes and in ectopic lymphoid structures of tumors has only recently begun. The role of mast cells in draining lymph nodes, in tertiary lymphoid tissues, and at metastatic sites of different tumors remains to be explored.

IgE has been suggested to play a protective role in tumor growth. Additional studies should investigate the role, if any, of IgE-mediated activation of mast cells in different human tumors.

The pro- or antitumorigenic role(s) of mast cells in different human tumors appears to be generally, but not always, cancer specific. We cannot exclude the possibility that subpopulation of TAMCs could play different, even opposite effects in various types/subtypes of tumors.

There is preliminary evidence that peritumoral mast cells play different roles compared to intratumoral mast cells. Studies in other experimental and human tumors will clarify whether the microlocalization of mast cells can markedly influence their effects.

Within the last years, gene expression profiling has demonstrated that several individual human cancers (e.g., melanoma, gastric, lung, and breast cancers) are heterogeneous with a spectrum of molecular changes. The complex heterogeneity (spatial, temporal, intratumor, intertumor) of the tumor microenvironment adds an additional layer of complexity. An important task will be to correlate the role of TAMCs in different subtypes of human cancers as defined by genetic markers.

There is recent evidence in melanoma, in prostate, and in pancreatic cancer that mast cells can play different roles in early and late phases of tumor initiation and growth. This fascinating hypothesis deserves to be further investigated in order to clarify the functional role of TAMCs in the progression of experimental and human cancers.

Two strains of mast cell-deficient mice with mutations affecting Kit, KitW/w−v and KitW−sh/W−sh, have been extensively used to study the role of mast cells in tumor growth. These mice are profoundly deficient in mast cells and also exhibit several other abnormalities, such as basophil deficiency. Recent evidence suggests that basophils can play a role in human pancreatic cancer. New Kit-independent mast cell-deficient mice have been used to evaluate the role of mast cells in cutaneous lymphoma, malignant pleural effusion, and skin cancerogenesis. Collectively, results obtained with mast cell-deficient mouse models should be interpreted with caution because even new mouse mutants with unperturbed Kit function also showed some hematological abnormalities.

Mast cells are plastic cells: their phenotype depends on their anatomical location and the physiological or pathological context. TAMCs are exposed in a hostile tumor environment to increased levels of lactate, PGE2, adenosine, IFN-γ, and a low pH. This metabolic milieu can profoundly alter mast cell behavior. It has been shown that it is possible to reverse the immunosuppressive and pro-tumoral properties of tumor-associated macrophages. A better knowledge of the pro-tumorigenic profile of TAMCs could help to “re-educate” these cells to play an antitumorigenic role.

Tumor cells evade host immune attack by expressing several checkpoint inhibitors, such as programmed cell death-1 (PD-1) ligands (PD-L1 and PD-L2) which inhibit PD-1+ lymphocytes in tumor microenvironment. Monoclonal antibodies targeting the PD-1/PD-L1 pathway unleash antitumor immunity and have revolutionized the management of a wide spectrum of malignancies. Certain cancer cells (e.g., melanoma) express also PD-1, in addition to PD-L1, providing an additional tumor intrinsic mechanism enhancing the pro-tumorigenic effect of PD-1/PD-L1 axis. Mouse mast cells highly express PD-L1 and, to a lesser extent, PD-L2. An important task will be to investigate the role of PD-L1+ TAMC in tumor microenvironment.

All the above implies that elucidation of the roles of mast cells in different human tumors will demand studies of increasing complexity beyond those assessing merely mast cell density and microlocalization.

Conclusion

In several human and experimental tumors, mast cells and their mediators play a pro-tumorigenic role. However, in other tumors and even in the same tumor, mast cells seem to play a protective role. These apparently controversial results might reflect differences in stage, grade, and subtypes of tumors, different methods to identify mast cells (e.g., tryptase+, chymase+, toluidine blue, CD117+, Giemsa), or different microanatomical compartment (i.e., peritumoral vs intratumoral) analyzed in the various studies. Whatever the mechanisms, there are many unanswered questions that need to be addressed before we understand whether mast cells are an ally, adversary, or innocent bystander in human cancers.

Author Contributions

GV, MG, and SL conceived and designed the review. All the authors contributed intellectually and to the writing of the submitted version of the manuscript.

Conflict of Interest Statement

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

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    Not in our library10.1046/j.1365-2362.2003.01152.x

  42. 42.

    Ribatti D Vacca A Ria R Marzullo A Nico B Filotico R Neovascularisation, expression of fibroblast growth factor-2, and mast cells with tryptase activity increase simultaneously with pathological progression in human malignant melanoma. Eur J Cancer (2003 ) 39:666–74.

    Not in our library10.1016/s0959-8049(02)00150-8

  43. 43.

    Siiskonen H Poukka M Bykachev A Tyynela-Korhonen K Sironen R Pasonen-Seppanen S Low numbers of tryptase+ and chymase+ mast cells associated with reduced survival and advanced tumor stage in melanoma. Melanoma Res (2015 ) 25:479–85.

    Not in our library10.1097/cmr.0000000000000192

  44. 44.

    Takahashi K Mulliken JB Kozakewich HP Rogers RA Folkman J Ezekowitz RA. Cellular markers that distinguish the phases of hemangioma during infancy and childhood. J Clin Invest (1994 ) 93:2357–64.

    Not in our library10.1172/jci117241

  45. 45.

    Toth-Jakatics R Jimi S Takebayashi S Kawamoto N. Cutaneous malignant melanoma: correlation between neovascularization and peritumor accumulation of mast cells overexpressing vascular endothelial growth factor. Hum Pathol (2000 ) 31:955–60.

    Not in our library10.1053/hupa.2000.16658

  46. 46.

    Johnson C Huynh V Hargrove L Kennedy L Graf-Eaton A Owens J Inhibition of mast cell-derived histamine decreases human cholangiocarcinoma growth and differentiation via c-kit/stem cell factor-dependent signaling. Am J Pathol (2016 ) 186:123–33.

    Not in our library10.1016/j.ajpath.2015.09.016

  47. 47.
    Bone marrow angiogenesis and mast cell density increase simultaneously with progression of human multiple myeloma · 1999
    No notice
  48. 48.

    Rabenhorst A Schlaak M Heukamp LC Forster A Theurich S von Bergwelt-Baildon M Mast cells play a protumorigenic role in primary cutaneous lymphoma. Blood (2012 ) 120:2042–54.

    Not in our library10.1182/blood-2012-03-415638

  49. 49.

    Vyzoukaki R Tsirakis G Pappa CA Devetzoglou M Tzardi M Alexandrakis MG. The impact of mast cell density on the progression of bone disease in multiple myeloma patients. Int Arch Allergy Immunol (2015 ) 168:263–8.

    Not in our library10.1159/000443275

  50. 50.

    Tripodo C Gri G Piccaluga PP Frossi B Guarnotta C Piconese S Mast cells and Th17 cells contribute to the lymphoma-associated pro-inflammatory microenvironment of angioimmunoblastic T-cell lymphoma. Am J Pathol (2010 ) 177:792–802.

    Not in our library10.2353/ajpath.2010.091286

  51. 51.

    Taskinen M Karjalainen-Lindsberg ML Leppa S. Prognostic influence of tumor-infiltrating mast cells in patients with follicular lymphoma treated with rituximab and CHOP. Blood (2008 ) 111:4664–7.

    Not in our library10.1182/blood-2007-11-125823

  52. 52.

    Franco G Guarnotta C Frossi B Piccaluga PP Boveri E Gulino A Bone marrow stroma CD40 expression correlates with inflammatory mast cell infiltration and disease progression in splenic marginal zone lymphoma. Blood (2014 ) 123:1836–49.

    Not in our library10.1182/blood-2013-04-497271

  53. 53.

    Molin D Edstrom A Glimelius I Glimelius B Nilsson G Sundstrom C Mast cell infiltration correlates with poor prognosis in Hodgkin’s lymphoma. Br J Haematol (2002 ) 119:122–4.

    Not in our library10.1046/j.1365-2141.2002.03768.x

  54. 54.

    Andersen MD Kamper P Nielsen PS Bendix K Riber-Hansen R Steiniche T Tumour-associated mast cells in classical Hodgkin’s lymphoma: correlation with histological subtype, other tumour-infiltrating inflammatory cell subsets and outcome. Eur J Haematol (2016 ) 96:252–9.

    Not in our library10.1111/ejh.12583

  55. 55.

    Englund A Molin D Enblad G Karlen J Glimelius I Ljungman G The role of tumour-infiltrating eosinophils, mast cells and macrophages in classical and nodular lymphocyte predominant Hodgkin lymphoma in children. Eur J Haematol (2016 ) 97:430–8.

    Not in our library10.1111/ejh.12747

  56. 56.

    Marone G Varricchi G Loffredo S Granata F. Mast cells and basophils in inflammatory and tumor angiogenesis and lymphangiogenesis. Eur J Pharmacol (2016 ) 778:146–51.

    Not in our library10.1016/j.ejphar.2015.03.088

  57. 57.

    Yamamoto T Katayama I Nishioka K. Expression of stem cell factor in basal cell carcinoma. Br J Dermatol (1997 ) 137:709–13.

    Not in our library10.1046/j.1365-2133.1997.19402055.x

  58. 58.

    de Paulis A Minopoli G Arbustini E de Crescenzo G Dal Piaz F Pucci P Stem cell factor is localized in, released from, and cleaved by human mast cells. J Immunol (1999 ) 163:2799–808.

    Not in our library

  59. 59.

    Detoraki A Staiano RI Granata F Giannattasio G Prevete N de Paulis A Vascular endothelial growth factors synthesized by human lung mast cells exert angiogenic effects. J Allergy Clin Immunol (2009 ) 123:1142–9, 1149.e1141–1145.

    Not in our library10.1016/j.jaci.2009.01.044

  60. 60.

    Prevete N Staiano RI Granata F Detoraki A Necchi V Ricci V Expression and function of angiopoietins and their tie receptors in human basophils and mast cells. J Biol Regul Homeost Agents (2013 ) 27:827–39.

    Not in our library

  61. 61.

    Visciano C Liotti F Prevete N Cali G Franco R Collina F Mast cells induce epithelial-to-mesenchymal transition and stem cell features in human thyroid cancer cells through an IL-8-Akt-Slug pathway. Oncogene (2015 ) 34:5175–86.

    Not in our library10.1038/onc.2014.441

  62. 62.

    Kay LJ Yeo WW Peachell PT. Prostaglandin E2 activates EP2 receptors to inhibit human lung mast cell degranulation. Br J Pharmacol (2006 ) 147:707–13.

    Not in our library10.1038/sj.bjp.0706664

  63. 63.

    Weller CL Collington SJ Hartnell A Conroy DM Kaise T Barker JE Chemotactic action of prostaglandin E2 on mouse mast cells acting via the PGE2 receptor 3. Proc Natl Acad Sci U S A (2007 ) 104:11712–7.

    Not in our library10.1073/pnas.0701700104

  64. 64.

    Godot V Arock M Garcia G Capel F Flys C Dy M H4 histamine receptor mediates optimal migration of mast cell precursors to CXCL12. J Allergy Clin Immunol (2007 ) 120:827–34.

    Not in our library10.1016/j.jaci.2007.05.046

  65. 65.

    Weller CL Collington SJ Brown JK Miller HR Al-Kashi A Clark P Leukotriene B4, an activation product of mast cells, is a chemoattractant for their progenitors. J Exp Med (2005 ) 201:1961–71.

    Not in our library10.1084/jem.20042407

  66. 66.

    Nagasaka A Matsue H Matsushima H Aoki R Nakamura Y Kambe N Osteopontin is produced by mast cells and affects IgE-mediated degranulation and migration of mast cells. Eur J Immunol (2008 ) 38:489–99.

    Not in our library10.1002/eji.200737057

  67. 67.

    Rao Q Chen Y Yeh CR Ding J Li L Chang C Recruited mast cells in the tumor microenvironment enhance bladder cancer metastasis via modulation of ERbeta/CCL2/CCR2 EMT/MMP9 signals. Oncotarget (2016 ) 7:7842–55.

    Not in our library10.18632/oncotarget.5467

  68. 68.

    Acikalin MF Oner U Topcu I Yasar B Kiper H Colak E. Tumour angiogenesis and mast cell density in the prognostic assessment of colorectal carcinomas. Dig Liver Dis (2005 ) 37:162–9.

    Not in our library10.1016/j.dld.2004.09.028

  69. 69.

    Gulubova M Vlaykova T. Prognostic significance of mast cell number and microvascular density for the survival of patients with primary colorectal cancer. J Gastroenterol Hepatol (2009 ) 24:1265–75.

    Not in our library10.1111/j.1440-1746.2007.05009.x

  70. 70.
    Correlation between serum tryptase, mast cells positive to tryptase and microvascular density in colo-rectal cancer patients: possible biological-clinical significance. · 2014
    Not checked
  71. 71.

    Wang S Li L Shi R Liu X Zhang J Zou Z Mast cell targeted chimeric toxin can be developed as an adjunctive therapy in colon cancer treatment. Toxins (Basel) (2016 ) 8:1–18.

    Not in our library10.3390/toxins8030071

  72. 72.

    Malfettone A Silvestris N Saponaro C Ranieri G Russo A Caruso S High density of tryptase-positive mast cells in human colorectal cancer: a poor prognostic factor related to protease-activated receptor 2 expression. J Cell Mol Med (2013 ) 17:1025–37.

    Not in our library10.1111/jcmm.12073

  73. 73.

    Suzuki S Ichikawa Y Nakagawa K Kumamoto T Mori R Matsuyama R High infiltration of mast cells positive to tryptase predicts worse outcome following resection of colorectal liver metastases. BMC Cancer (2015 ) 15:840.

    Not in our library10.1186/s12885-015-1863-z

  74. 74.

    Fakhrjou A Niroumand-Oscoei SM Somi MH Ghojazadeh M Naghashi S Samankan S. Prognostic value of tumor-infiltrating mast cells in outcome of patients with esophagus squamous cell carcinoma. J Gastrointest Cancer (2014 ) 45:48–53.

    Not in our library10.1007/s12029-013-9550-2

  75. 75.

    Yano H Kinuta M Tateishi H Nakano Y Matsui S Monden T Mast cell infiltration around gastric cancer cells correlates with tumor angiogenesis and metastasis. Gastric Cancer (1999 ) 2:26–32.

    Not in our library10.1007/s101200050017

  76. 76.

    Kondo K Muramatsu M Okamoto Y Jin D Takai S Tanigawa N Expression of chymase-positive cells in gastric cancer and its correlation with the angiogenesis. J Surg Oncol (2006 ) 93:36–42; discussion 42–33.

    Not in our library10.1002/jso.20394

  77. 77.

    Ammendola M Sacco R Donato G Zuccala V Russo E Luposella M Mast cell positivity to tryptase correlates with metastatic lymph nodes in gastrointestinal cancer patients treated surgically. Oncology (2013 ) 85:111–6.

    Not in our library10.1159/000351145

  78. 78.

    Ammendola M Marech I Sammarco G Zuccala V Luposella M Zizzo N Infiltrating mast cells correlate with angiogenesis in bone metastases from gastric cancer patients. Int J Mol Sci (2015 ) 16:3237–50.

    Not in our library10.3390/ijms16023237

  79. 79.

    Tu JF Pan HY Ying XH Lou J Ji JS Zou H. Mast cells comprise the major of interleukin 17-producing cells and predict a poor prognosis in hepatocellular carcinoma. Medicine (Baltimore) (2016 ) 95:e3220.

    Not in our library10.1097/md.0000000000003220

  80. 80.

    Imada A Shijubo N Kojima H Abe S. Mast cells correlate with angiogenesis and poor outcome in stage I lung adenocarcinoma. Eur Respir J (2000 ) 15:1087–93.

    Not in our library10.1034/j.1399-3003.2000.01517.x

  81. 81.

    Takanami I Takeuchi K Naruke M. Mast cell density is associated with angiogenesis and poor prognosis in pulmonary adenocarcinoma. Cancer (2000 ) 88:2686–92.

    Not in our library10.1002/1097-0142(20000615)88:12<2686::aid-cncr6>3.3.co;2-y

  82. 82.
    Immune and Inflammatory Cell Composition of Human Lung Cancer Stroma. · 2015
    Not checked
  83. 83.

    Holzel M Landsberg J Glodde N Bald T Rogava M Riesenberg S A preclinical model of malignant peripheral nerve sheath tumor-like melanoma is characterized by infiltrating mast cells. Cancer Res (2016 ) 76:251–63.

    Not in our library10.1158/0008-5472.can-15-1090

  84. 84.

    Strouch MJ Cheon EC Salabat MR Krantz SB Gounaris E Melstrom LG Crosstalk between mast cells and pancreatic cancer cells contributes to pancreatic tumor progression. Clin Cancer Res (2010 ) 16:2257–65.

    Not in our library10.1158/1078-0432.ccr-09-1230

  85. 85.

    Chang DZ Ma Y Ji B Wang H Deng D Liu Y Mast cells in tumor microenvironment promotes the in vivo growth of pancreatic ductal adenocarcinoma. Clin Cancer Res (2011 ) 17:7015–23.

    Not in our library10.1158/1078-0432.ccr-11-0607

  86. 86.

    Cai SW Yang SZ Gao J Pan K Chen JY Wang YL Prognostic significance of mast cell count following curative resection for pancreatic ductal adenocarcinoma. Surgery (2011 ) 149:576–84.

    Not in our library10.1016/j.surg.2010.10.009

  87. 87.

    Li L Dang Q Xie H Yang Z He D Liang L Infiltrating mast cells enhance prostate cancer invasion via altering LncRNA-HOTAIR/PRC2-androgen receptor (AR)-MMP9 signals and increased stem/progenitor cell population. Oncotarget (2015 ) 6:14179–90.

    Not in our library10.18632/oncotarget.3651

  88. 88.

    Nonomura N Takayama H Nishimura K Oka D Nakai Y Shiba M Decreased number of mast cells infiltrating into needle biopsy specimens leads to a better prognosis of prostate cancer. Br J Cancer (2007 ) 97:952–6.

    Not in our library10.1038/sj.bjc.6603962

  89. 89.

    Wasiuk A Dalton DK Schpero WL Stan RV Conejo-Garcia JR Noelle RJ. Mast cells impair the development of protective anti-tumor immunity. Cancer Immunol Immunother (2012 ) 61:2273–82.

    Not in our library10.1007/s00262-012-1276-7

  90. 90.

    Wedemeyer J Galli SJ. Decreased susceptibility of mast cell-deficient Kit(W)/Kit(W-v) mice to the development of 1, 2-dimethylhydrazine-induced intestinal tumors. Lab Invest (2005 ) 85:388–96.

    Not in our library10.1038/labinvest.3700232

  91. 91.

    Gounaris E Erdman SE Restaino C Gurish MF Friend DS Gounari F Mast cells are an essential hematopoietic component for polyp development. Proc Natl Acad Sci U S A (2007 ) 104:19977–82.

    Not in our library10.1073/pnas.0704620104

  92. 92.

    Nordlund JJ Askenase PW. The effect of histamine, antihistamines, and a mast cell stabilizer on the growth of cloudman melanoma cells in DBA/2 mice. J Invest Dermatol (1983 ) 81:28–31.

    Not in our library10.1111/1523-1747.ep12538356

  93. 93.

    Jeong HJ Oh HA Nam SY Han NR Kim YS Kim JH The critical role of mast cell-derived hypoxia-inducible factor-1alpha in human and mice melanoma growth. Int J Cancer (2013 ) 132:2492–501.

    Not in our library10.1002/ijc.27937

  94. 94.

    Soucek L Lawlor ER Soto D Shchors K Swigart LB Evan GI. Mast cells are required for angiogenesis and macroscopic expansion of Myc-induced pancreatic islet tumors. Nat Med (2007 ) 13:1211–8.

    Not in our library10.1038/nm1649

  95. 95.

    Soucek L Buggy JJ Kortlever R Adimoolam S Monclus HA Allende MT Modeling pharmacological inhibition of mast cell degranulation as a therapy for insulinoma. Neoplasia (2011 ) 13:1093–100.

    Not in our library10.1593/neo.11980

  96. 96.

    Nakayama T Yao L Tosato G. Mast cell-derived angiopoietin-1 plays a critical role in the growth of plasma cell tumors. J Clin Invest (2004 ) 114:1317–25.

    Not in our library10.1172/jci22089

  97. 97.

    Xie H Li C Dang Q Chang LS Li L. Infiltrating mast cells increase prostate cancer chemotherapy and radiotherapy resistances via modulation of p38/p53/p21 and ATM signals. Oncotarget (2016 ) 7:1341–53.

    Not in our library10.18632/oncotarget.6372

  98. 98.

    Coussens LM Raymond WW Bergers G Laig-Webster M Behrendtsen O Werb Z Inflammatory mast cells up-regulate angiogenesis during squamous epithelial carcinogenesis. Genes Dev (1999 ) 13:1382–97.

    Not in our library10.1101/gad.13.11.1382

  99. 99.

    Sarchio SN Scolyer RA Beaugie C McDonald D Marsh-Wakefield F Halliday GM Pharmacologically antagonizing the CXCR4-CXCL12 chemokine pathway with AMD3100 inhibits sunlight-induced skin cancer. J Invest Dermatol (2014 ) 134:1091–100.

    Not in our library10.1038/jid.2013.424

  100. 100.

    Tournilhac O Santos DD Xu L Kutok J Tai YT Le Gouill S Mast cells in Waldenstrom’s macroglobulinemia support lymphoplasmacytic cell growth through CD154/CD40 signaling. Ann Oncol (2006 ) 17:1275–82.

    Not in our library10.1093/annonc/mdl109

  101. 101.

    Sinnamon MJ Carter KJ Sims LP Lafleur B Fingleton B Matrisian LM. A protective role of mast cells in intestinal tumorigenesis. Carcinogenesis (2008 ) 29:880–6.

    Not in our library10.1093/carcin/bgn040

  102. 102.

    Yang XD Ai W Asfaha S Bhagat G Friedman RA Jin G Histamine deficiency promotes inflammation-associated carcinogenesis through reduced myeloid maturation and accumulation of CD11b+Ly6G+ immature myeloid cells. Nat Med (2011 ) 17:87–95.

    Not in our library10.1038/nm.2278

  103. 103.

    Purwar R Schlapbach C Xiao S Kang HS Elyaman W Jiang X Robust tumor immunity to melanoma mediated by interleukin-9-producing T cells. Nat Med (2012 ) 18:1248–53.

    Not in our library10.1038/nm.2856

  104. 104.

    Siebenhaar F Metz M Maurer M. Mast cells protect from skin tumor development and limit tumor growth during cutaneous de novo carcinogenesis in a kit-dependent mouse model. Exp Dermatol (2014 ) 23:159–64.

    Not in our library10.1111/exd.12328

  105. 105.

    Hedstrom G Berglund M Molin D Fischer M Nilsson G Thunberg U Mast cell infiltration is a favourable prognostic factor in diffuse large B-cell lymphoma. Br J Haematol (2007 ) 138:68–71.

    Not in our library10.1111/j.1365-2141.2007.06612.x

  106. 106.

    Dabiri S Huntsman D Makretsov N Cheang M Gilks B Bajdik C The presence of stromal mast cells identifies a subset of invasive breast cancers with a favorable prognosis. Mod Pathol (2004 ) 17:690–5.

    Not in our library10.1038/modpathol.3800094

  107. 107.

    Amini RM Aaltonen K Nevanlinna H Carvalho R Salonen L Heikkila P Mast cells and eosinophils in invasive breast carcinoma. BMC Cancer (2007 ) 7:165.

    Not in our library10.1186/1471-2407-7-165

  108. 108.

    della Rovere F Granata A Familiari D D’Arrigo G Mondello B Basile G. Mast cells in invasive ductal breast cancer: different behavior in high and minimum hormone-receptive cancers. Anticancer Res (2007 ) 27:2465–71.

    Not in our library

  109. 109.

    Rajput AB Turbin DA Cheang MC Voduc DK Leung S Gelmon KA Stromal mast cells in invasive breast cancer are a marker of favourable prognosis: a study of 4,444 cases. Breast Cancer Res Treat (2008 ) 107:249–57.

    Not in our library10.1007/s10549-007-9546-3

  110. 110.

    Tan SY Fan Y Luo HS Shen ZX Guo Y Zhao LJ. Prognostic significance of cell infiltrations of immunosurveillance in colorectal cancer. World J Gastroenterol (2005 ) 11:1210–4.

    Not in our library10.3748/wjg.v11.i8.1210

  111. 111.

    Carlini MJ Dalurzo MC Lastiri JM Smith DE Vasallo BC Puricelli LI Mast cell phenotypes and microvessels in non-small cell lung cancer and its prognostic significance. Hum Pathol (2010 ) 41:697–705.

    Not in our library10.1016/j.humpath.2009.04.029

  112. 112.

    Ali G Boldrini L Lucchi M Mussi A Corsi V Fontanini G. Tryptase mast cells in malignant pleural mesothelioma as an independent favorable prognostic factor. J Thorac Oncol (2009 ) 4:348–54.

    Not in our library10.1097/jto.0b013e3181989ddb

  113. 113.

    Welsh TJ Green RH Richardson D Waller DA O’Byrne KJ Bradding P. Macrophage and mast-cell invasion of tumor cell islets confers a marked survival advantage in non-small-cell lung cancer. J Clin Oncol (2005 ) 23:8959–67.

    Not in our library10.1200/jco.2005.01.4910

  114. 114.

    Shikotra A Ohri CM Green RH Waller DA Bradding P Mast cell phenotype, TNFalpha expression and degranulation status in non-small cell lung cancer. Sci Rep (2016 ) 6:38352

    Not in our library10.1038/srep38352

  115. 115.

    Chan JK Magistris A Loizzi V Lin F Rutgers J Osann K Mast cell density, angiogenesis, blood clotting, and prognosis in women with advanced ovarian cancer. Gynecol Oncol (2005 ) 99:20–5.

    Not in our library10.1016/j.ygyno.2005.05.042

  116. 116.

    Fleischmann A Schlomm T Kollermann J Sekulic N Huland H Mirlacher M Immunological microenvironment in prostate cancer: high mast cell densities are associated with favorable tumor characteristics and good prognosis. Prostate (2009 ) 69:976–81.

    Not in our library10.1002/pros.20948

  117. 117.

    Chichlowski M Westwood GS Abraham SN Hale LP. Role of mast cells in inflammatory bowel disease and inflammation-associated colorectal neoplasia in IL-10-deficient mice. PLoS One (2010 ) 5:e12220.

    Not in our library10.1371/journal.pone.0012220

  118. 118.

    Antsiferova M Martin C Huber M Feyerabend TB Forster A Hartmann K Mast cells are dispensable for normal and activin-promoted wound healing and skin carcinogenesis. J Immunol (2013 ) 191:6147–55.

    Not in our library10.4049/jimmunol.1301350

  119. 119.

    Xia Q Wu XJ Zhou Q Jing Z Hou JH Pan ZZ No relationship between the distribution of mast cells and the survival of stage IIIB colon cancer patients. J Transl Med (2011 ) 9:88.

    Not in our library10.1186/1479-5876-9-88

  120. 120.

    Dundar E Oner U Peker BC Metintas M Isiksoy S Ak G. The significance and relationship between mast cells and tumour angiogenesis in non-small cell lung carcinoma. J Int Med Res (2008 ) 36:88–95.

    Not in our library10.1177/147323000803600112

  121. 121.

    Tuna B Yorukoglu K Unlu M Mungan MU Kirkali Z. Association of mast cells with microvessel density in renal cell carcinomas. Eur Urol (2006 ) 50:530–4.

    Not in our library10.1016/j.eururo.2005.12.040

  122. 122.

    Ammendola M Sacco R Sammarco G Donato G Zuccala V Romano R Mast cells positive to tryptase and c-kit receptor expressing cells correlates with angiogenesis in gastric cancer patients surgically treated. Gastroenterol Res Pract (2013 ) 2013:703163.

    Not in our library10.1155/2013/703163

  123. 123.

    Esposito I Menicagli M Funel N Bergmann F Boggi U Mosca F Inflammatory cells contribute to the generation of an angiogenic phenotype in pancreatic ductal adenocarcinoma. J Clin Pathol (2004 ) 57:630–6.

    Not in our library10.1136/jcp.2003.014498

  124. 124.

    Marone G Granata F, editors. Angiogenesis, Lymphangiogenesis and Clinical Implications, Chem Immunol Allergy. (Vol. 99). Basel: Karger (2014 ).

    Not in our library10.1159/000352074

  125. 125.

    Folkman J Shing Y Angiogenesis. J Biol Chem (1992 ) 267:10931–4.

    Not in our library

  126. 126.

    Grutzkau A Kruger-Krasagakes S Baumeister H Schwarz C Kogel H Welker P Synthesis, storage, and release of vascular endothelial growth factor/vascular permeability factor (VEGF/VPF) by human mast cells: implications for the biological significance of VEGF206. Mol Biol Cell (1998 ) 9:875–84.

    Not in our library10.1091/mbc.9.4.875

  127. 127.

    Abdel-Majid RM Marshall JS. Prostaglandin E2 induces degranulation-independent production of vascular endothelial growth factor by human mast cells. J Immunol (2004 ) 172:1227–36.

    Not in our library10.4049/jimmunol.172.2.1227

  128. 128.

    Theoharides TC Zhang B Kempuraj D Tagen M Vasiadi M Angelidou A IL-33 augments substance P-induced VEGF secretion from human mast cells and is increased in psoriatic skin. Proc Natl Acad Sci U S A (2010 ) 107:4448–53.

    Not in our library10.1073/pnas.1000803107

  129. 129.

    Sismanopoulos N Delivanis DA Alysandratos KD Angelidou A Vasiadi M Therianou A IL-9 induces VEGF secretion from human mast cells and IL-9/IL-9 receptor genes are overexpressed in atopic dermatitis. PLoS One (2012 ) 7:e33271.

    Not in our library10.1371/journal.pone.0033271

  130. 130.

    Boesiger J Tsai M Maurer M Yamaguchi M Brown LF Claffey KP Mast cells can secrete vascular permeability factor/ vascular endothelial cell growth factor and exhibit enhanced release after immunoglobulin E-dependent upregulation of fc epsilon receptor I expression. J Exp Med (1998 ) 188:1135–45.

    Not in our library10.1084/jem.188.6.1135

  131. 131.

    Detoraki A Granata F Staibano S Rossi FW Marone G Genovese A. Angiogenesis and lymphangiogenesis in bronchial asthma. Allergy (2010 ) 65:946–58.

    Not in our library10.1111/j.1398-9995.2010.02372.x

  132. 132.

    Varricchi G Granata F Loffredo S Genovese A Marone G. Angiogenesis and lymphangiogenesis in inflammatory skin disorders. J Am Acad Dermatol (2015 ) 73:144–53.

    Not in our library10.1016/j.jaad.2015.03.041

  133. 133.

    Varey AH Rennel ES Qiu Y Bevan HS Perrin RM Raffy S VEGF 165 b, an antiangiogenic VEGF-A isoform, binds and inhibits bevacizumab treatment in experimental colorectal carcinoma: balance of pro- and antiangiogenic VEGF-A isoforms has implications for therapy. Br J Cancer (2008 ) 98:1366–79.

    Not in our library10.1038/sj.bjc.6604308

  134. 134.

    Bates DO Mavrou A Qiu Y Carter JG Hamdollah-Zadeh M Barratt S Detection of VEGF-A(xxx)b isoforms in human tissues. PLoS One (2013 ) 8:e68399.

    Not in our library10.1371/journal.pone.0068399

  135. 135.

    Harper SJ Bates DO. VEGF-A splicing: the key to anti-angiogenic therapeutics? Nat Rev Cancer (2008 ) 8:880–7.

    Not in our library10.1038/nrc2505

  136. 136.

    Loffredo S Borriello F Iannone R Ferrara AL Galdiero MR Gigantino V Group V secreted phospholipase A2 induces the release of pro-angiogenic and anti-angiogenic factors by human neutrophils. Front Immunol (2017 ). (in press).

    Not in our library10.3389/fimmu.2017.00443

  137. 137.

    Baram D Vaday GG Salamon P Drucker I Hershkoviz R Mekori YA. Human mast cells release metalloproteinase-9 on contact with activated T cells: juxtacrine regulation by TNF-alpha. J Immunol (2001 ) 167:4008–16.

    Not in our library10.4049/jimmunol.167.7.4008

  138. 138.

    Rinderknecht M Detmar M. Tumor lymphangiogenesis and melanoma metastasis. J Cell Physiol (2008 ) 216:347–54.

    Not in our library10.1002/jcp.21494

  139. 139.

    Granata F Frattini A Loffredo S Staiano RI Petraroli A Ribatti D Production of vascular endothelial growth factors from human lung macrophages induced by group IIA and group X secreted phospholipases A2. J Immunol (2010 ) 184:5232–41.

    Not in our library10.4049/jimmunol.0902501

  140. 140.

    Bowers HM Jr Mahapatro RC Kennedy JW. Numbers of mast cells in the axillary lymph nodes of breast cancer patients. Cancer (1979 ) 43:568–73.

    Not in our library10.1002/1097-0142(197902)43:2<568::aid-cncr2820430225>3.0.co;2-#

  141. 141.

    Zheng X Carstens JL Kim J Scheible M Kaye J Sugimoto H Epithelial-to-mesenchymal transition is dispensable for metastasis but induces chemoresistance in pancreatic cancer. Nature (2015 ) 527:525–30.

    Not in our library10.1038/nature16064

  142. 142.

    Fischer KR Durrans A Lee S Sheng J Li F Wong ST Epithelial-to-mesenchymal transition is not required for lung metastasis but contributes to chemoresistance. Nature (2015 ) 527:472–6.

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  143. 143.

    Scheel C Weinberg RA. Cancer stem cells and epithelial-mesenchymal transition: concepts and molecular links. Semin Cancer Biol (2012 ) 22:396–403.

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    Pittoni P Colombo MP. The dark side of mast cell-targeted therapy in prostate cancer. Cancer Res (2012 ) 72:831–5.

    Not in our library10.1158/0008-5472.can-11-3110

  145. 145.

    Bry M Kivela R Leppanen VM Alitalo K. Vascular endothelial growth factor-B in physiology and disease. Physiol Rev (2014 ) 94:779–94.

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  146. 146.

    Eruslanov EB Bhojnagarwala PS Quatromoni JG Stephen TL Ranganathan A Deshpande C Tumor-associated neutrophils stimulate T cell responses in early-stage human lung cancer. J Clin Invest (2014 ) 124:5466–80.

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  147. 147.

    Zaretsky JM Garcia-Diaz A Shin DS Escuin-Ordinas H Hugo W Hu-Lieskovan S Mutations associated with acquired resistance to PD-1 blockade in melanoma. N Engl J Med (2016 ) 375:819–29.

    Not in our library10.1056/nejmoa1604958

  148. 148.

    Varricchi G Galdiero MR Marone G Granata F Borriello F Marone G. Controversial role of mast cells in skin cancers. Exp Dermatol (2017 ) 26:11–7.

    Not in our library10.1111/exd.13107

  149. 149.

    Hirschhaeuser F Sattler UG Mueller-Klieser W. Lactate: a metabolic key player in cancer. Cancer Res (2011 ) 71:6921–5.

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  150. 150.

    Gottfried E Kreutz M Mackensen A. Tumor metabolism as modulator of immune response and tumor progression. Semin Cancer Biol (2012 ) 22:335–41.

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  151. 151.

    Marone G Findlay SR Lichtenstein LM. Adenosine receptor on human basophils: modulation of histamine release. J Immunol (1979 ) 123:1473–7.

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    Gulliksson M Carvalho RF Ulleras E Nilsson G. Mast cell survival and mediator secretion in response to hypoxia. PLoS One (2010 ) 5:e12360.

    Not in our library10.1371/journal.pone.0012360

  153. 153.

    Walczak-Drzewiecka A Ratajewski M Wagner W Dastych J. HIF-1alpha is up-regulated in activated mast cells by a process that involves calcineurin and NFAT. J Immunol (2008 ) 181:1665–72.

    Not in our library10.4049/jimmunol.181.3.1665

  154. 154.

    Marquardt DL Gruber HE Wasserman SI. Adenosine release from stimulated mast cells. Proc Natl Acad Sci U S A (1984 ) 81:6192–6.

    Not in our library10.1073/pnas.81.19.6192

  155. 155.

    Peachell PT Columbo M Kagey-Sobotka A Lichtenstein LM Marone G. Adenosine potentiates mediator release from human lung mast cells. Am Rev Respir Dis (1988 ) 138:1143–51.

    Not in our library10.1164/ajrccm/138.5.1143

  156. 156.

    Visciano C Prevete N Liotti F Marone G. Tumor-associated mast cells in thyroid cancer. Int J Endocrinol (2015 ) 2015:705169.

    Not in our library10.1155/2015/705169

  157. 157.

    Lin TJ Issekutz TB Marshall JS. SDF-1 induces IL-8 production and transendothelial migration of human cord blood-derived mast cells. Int Arch Allergy Immunol (2001 ) 124:142–5.

    Not in our library10.1159/000053693

  158. 158.

    Jensen-Jarolim E Achatz G Turner MC Karagiannis S Legrand F Capron M AllergoOncology: the role of IgE-mediated allergy in cancer. Allergy (2008 ) 63:1255–66.

    Not in our library10.1111/j.1398-9995.2008.01768.x

  159. 159.

    Chan JK Roth J Oppenheim JJ Tracey KJ Vogl T Feldmann M Alarmins: awaiting a clinical response. J Clin Invest (2012 ) 122:2711–9.

    Not in our library10.1172/jci62423

  160. 160.

    Subramanian H Gupta K Guo Q Price R Ali H. Mas-related gene X2 (MrgX2) is a novel G protein-coupled receptor for the antimicrobial peptide LL-37 in human mast cells: resistance to receptor phosphorylation, desensitization, and internalization. J Biol Chem (2011 ) 286:44739–49.

    Not in our library10.1074/jbc.m111.277152

  161. 161.

    Byrne SN Beaugie C O’Sullivan C Leighton S Halliday GM. The immune-modulating cytokine and endogenous Alarmin interleukin-33 is upregulated in skin exposed to inflammatory UVB radiation. Am J Pathol (2011 ) 179:211–22.

    Not in our library10.1016/j.ajpath.2011.03.010

  162. 162.

    de Vries VC Pino-Lagos K Nowak EC Bennett KA Oliva C Noelle RJ. Mast cells condition dendritic cells to mediate allograft tolerance. Immunity (2011 ) 35:550–61.

    Not in our library10.1016/j.immuni.2011.09.012

  163. 163.

    Rivellese F Suurmond J Habets K Dorjee AL Ramamoorthi N Townsend MJ Ability of interleukin-33- and immune complex-triggered activation of human mast cells to down-regulate monocyte-mediated immune responses. Arthritis Rheumatol (2015 ) 67:2343–53.

    Not in our library10.1002/art.39192

  164. 164.

    Okayama Y Hagaman DD Metcalfe DD. A comparison of mediators released or generated by IFN-gamma-treated human mast cells following aggregation of Fc gamma RI or Fc epsilon RI. J Immunol (2001 ) 166:4705–12.

    Not in our library10.4049/jimmunol.166.7.4705

  165. 165.

    Daeron M. Innate myeloid cells under the control of adaptive immunity: the example of mast cells and basophils. Curr Opin Immunol (2016 ) 38:101–8.

    Not in our library10.1016/j.coi.2015.12.004

  166. 166.

    Oldford SA Marshall JS. Mast cells as targets for immunotherapy of solid tumors. Mol Immunol (2015 ) 63:113–24.

    Not in our library10.1016/j.molimm.2014.02.020

  167. 167.

    Shijubo N Kojima H Nagata M Ohchi T Suzuki A Abe S Tumor angiogenesis of non-small cell lung cancer. Microsc Res Tech (2003 ) 60:186–98.

    Not in our library10.1002/jemt.10257

  168. 168.

    Triggiani M Hubbard WC Chilton FH. Synthesis of 1-acyl-2-acetyl-sn-glycero-3-phosphocholine by an enriched preparation of the human lung mast cell. J Immunol (1990 ) 144:4773–80.

    Not in our library

  169. 169.

    Damiani E Puebla-Osorio N Gorbea E Ullrich SE. Platelet-activating factor induces epigenetic modifications in human mast cells. J Invest Dermatol (2015 ) 135:3034–40.

    Not in our library10.1038/jid.2015.336

  170. 170.

    Chacon-Salinas R Chen L Chavez-Blanco AD Limon-Flores AY Ma Y Ullrich SE. An essential role for platelet-activating factor in activating mast cell migration following ultraviolet irradiation. J Leukoc Biol (2014 ) 95:139–48.

    Not in our library10.1189/jlb.0811409

  171. 171.

    de Paulis A Marino I Ciccarelli A de Crescenzo G Concardi M Verga L Human synovial mast cells. I. Ultrastructural in situ and in vitro immunologic characterization. Arthritis Rheum (1996 ) 39:1222–33.

    Not in our library10.1002/art.1780390723

  172. 172.

    Movahedi K Laoui D Gysemans C Baeten M Stange G Van den Bossche J Different tumor microenvironments contain functionally distinct subsets of macrophages derived from Ly6C(high) monocytes. Cancer Res (2010 ) 70:5728–39.

    Not in our library10.1158/0008-5472.can-09-4672

  173. 173.

    Motakis E Guhl S Ishizu Y Itoh M Kawaji H de Hoon M Redefinition of the human mast cell transcriptome by deep-CAGE sequencing. Blood (2014 ) 123:e58–67.

    Not in our library10.1182/blood-2013-02-483792

  174. 174.

    Lundstrom SL Saluja R Adner M Haeggstrom JZ Nilsson G Wheelock CE. Lipid mediator metabolic profiling demonstrates differences in eicosanoid patterns in two phenotypically distinct mast cell populations. J Lipid Res (2013 ) 54:116–26.

    Not in our library10.1194/jlr.m030171

  175. 175.

    Sotiriou C Pusztai L Gene-expression signatures in breast cancer. N Engl J Med (2009 ) 360:790–800.

    Not in our library10.1056/nejmra0801289

  176. 176.

    Popper HH Ryska A Timar J Olszewski W. Molecular testing in lung cancer in the era of precision medicine. Transl Lung Cancer Res (2014 ) 3:291–300.

    Not in our library10.3978/j.issn.2218-6751.2014.10.01

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    Kim SY Kim SN Hahn HJ Lee YW Choe YB Ahn KJ. Metaanalysis of BRAF mutations and clinicopathologic characteristics in primary melanoma. J Am Acad Dermatol (2015 ) 72:1036–46.e1032.

    Not in our library10.1016/j.jaad.2015.02.1113

  178. 178.

    Riquelme I Saavedra K Espinoza JA Weber H Garcia P Nervi B Molecular classification of gastric cancer: towards a pathway-driven targeted therapy. Oncotarget (2015 ) 6:24750–79.

    Not in our library10.18632/oncotarget.4990

  179. 179.

    Coussens LM Zitvogel L Palucka AK. Neutralizing tumor-promoting chronic inflammation: a magic bullet? Science (2013 ) 339:286–91.

    Not in our library10.1126/science.1232227

  180. 180.

    Bedard PL Hansen AR Ratain MJ Siu LL. Tumour heterogeneity in the clinic. Nature (2013 ) 501:355–64.

    Not in our library10.1038/nature12627

  181. 181.

    Mancardi DA Jonsson F Iannascoli B Khun H Van Rooijen N Huerre M Cutting edge: the murine high-affinity IgG receptor FcgammaRIV is sufficient for autoantibody-induced arthritis. J Immunol (2011 ) 186:1899–903.

    Not in our library10.4049/jimmunol.1003642

  182. 182.

    Feyerabend TB Weiser A Tietz A Stassen M Harris N Kopf M Cre-mediated cell ablation contests mast cell contribution in models of antibody- and T cell-mediated autoimmunity. Immunity (2011 ) 35:832–44.

    Not in our library10.1016/j.immuni.2011.09.015

  183. 183.

    De Monte L Wormann S Brunetto E Heltai S Magliacane G Reni M Basophil recruitment into tumor-draining lymph nodes correlates with Th2 inflammation and reduced survival in pancreatic cancer patients. Cancer Res (2016 ) 76:1792–803.

    Not in our library10.1158/0008-5472.can-15-1801-t

  184. 184.

    Rodewald HR Feyerabend TB. Widespread immunological functions of mast cells: fact or fiction? Immunity (2012 ) 37:13–24.

    Not in our library10.1016/j.immuni.2012.07.007

  185. 185.

    Reber LL Marichal T Galli SJ. New models for analyzing mast cell functions in vivo. Trends Immunol (2012 ) 33:613–25.

    Not in our library10.1016/j.it.2012.09.008

  186. 186.

    Duluc D Corvaisier M Blanchard S Catala L Descamps P Gamelin E Interferon-gamma reverses the immunosuppressive and protumoral properties and prevents the generation of human tumor-associated macrophages. Int J Cancer (2009 ) 125:367–73.

    Not in our library10.1002/ijc.24401

  187. 187.

    Hagemann T Lawrence T McNeish I Charles KA Kulbe H Thompson RG “Re-educating” tumor-associated macrophages by targeting NF-kappaB. J Exp Med (2008 ) 205:1261–8.

    Not in our library10.1084/jem.20080108

  188. 188.

    Sharma P Allison JP. Immune checkpoint targeting in cancer therapy: toward combination strategies with curative potential. Cell (2015 ) 161:205–14.

    Not in our library10.1016/j.cell.2015.03.030

  189. 189.

    Tan S Chen D Liu K He M Song H Shi Y Crystal clear: visualizing the intervention mechanism of the PD-1/PD-L1 interaction by two cancer therapeutic monoclonal antibodies. Protein Cell (2016 ) 7:866–77.

    Not in our library10.1007/s13238-016-0337-7

  190. 190.

    Kleffel S Posch C Barthel SR Mueller H Schlapbach C Guenova E Melanoma cell-intrinsic PD-1 receptor functions promote tumor growth. Cell (2015 ) 162:1242–56.

    Not in our library10.1016/j.cell.2015.08.052

  191. 191.

    Nakae S Suto H Iikura M Kakurai M Sedgwick JD Tsai M Mast cells enhance T cell activation: importance of mast cell costimulatory molecules and secreted TNF. J Immunol (2006 ) 176:2238–48.

    Not in our library10.4049/jimmunol.176.4.2238

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