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PLENARY PAPER
From the Department of Dermatology, University of
Pittsburgh, PA.
Interferon Dendritic cells (DCs) are rare sentinel cells that
provide a first defense against invading microbial and viral
pathogens.1,2 Encounter with these antigens induces DC
maturation and allows for the subsequent activation of naive,
antigen-specific T cells and B cells.2,3 DCs can be
divided into distinct subsets based on differential phenotype and
function. In humans, myeloid DCs (or DC1) express "myeloid" markers
CD13, CD33, high levels of granulocyte-macrophage colony-stimulating
factor (GM-CSF) receptors, and accessory molecules. They can induce
T-cell proliferation and TH1-like cytokine profiles.2
Human lymphoid DCs or DC24,5 do not express the CD11c
antigen; instead, they express high levels of interleukin-3R (IL-3R),
CD68, CD4, CD45R, and the inhibitory receptor IL-T3.6 Type
1 interferons are clinically important in infectious diseases and in
the treatment of leukemia and lymphoma. Many different cell types have
the capacity to produce interferon In contrast to human DCs, mouse DC subsets described so far express
CD11c.9,10 Murine myeloid DCs can be isolated based on the
coexpression of CD11c and CD11b and can induce a strong allogeneic
mixed-lymphocyte reaction (MLR).11 The
lymphoid-related DC expresses CD11c and CD8 The origin of plasmacytoid DCs remains controversial. In humans, a
recent study on immunoglobulin (Ig) gene arrangement suggests a
lymphoid origin.16 However, as their original
terms To identify the mouse natural IFN-producing cell, or pDC, total
leukocyte populations were isolated from different tissues and
stimulated with herpes simplex virus (HSV), and culture supernatants were analyzed for IFN- Using multicolor fluorescence-activated cell sorting (FACS), the
phenotype murine pDC could be determined. It was found to exhibit a
phenotype slightly different from that of its human counterpart in that
it expressed CD11c, but they were similar in expressing CD45R and low
levels of CD4 and IL-3R. Short-term culture of murine pDC in media
containing IL-3 and anti-CD40 antibodies induced maturation, resulting
in the evolution of long cellular protrusions and high expression of
major histocompatibility complex class 2 and accessory molecules.
Mice
Reagents
Cytokine mobilization Mice were mobilized with recombinant Flt3L ± GM-CSF or combinations thereof at a concentration of 20 µg/mouse per day in phosphate-buffered saline (PBS) by subcutaneous injections in the neck for 7 consecutive days under an IACUC-approved protocol.Cell preparation Bone marrow cells were obtained by flushing femurs and tibias with PBS using a 23-gauge needle. Red cells were hypotonically lysed, and the residual cells were washed and kept on ice until used. Spleens were isolated, and a single-cell suspension was made by passing the spleen through a nylon cell strainer (Falcon; Becton Dickinson, Franklin Lakes, NJ). After lysis of red cells by ammonium chloride solution, cells were incubated with antibodies to block FcR interactions (FcR-block, CD16/CD32, Pharmingen, at 1 µg/1 × 106 cells at 10 × 106/mL in PBS) for 15 minutes on ice. For isolation of CD11c+ cells using directly conjugated MACS magnetic beads (Miltenyi Biotec), CD11c beads were added according to the manufacturer's instructions. After incubation for 20 minutes at 4°C, cells were washed and passed over a MACS column. Positively selected cells were isolated and suspended in appropriate buffer. Purity was checked routinely by FACS and was found to be greater than 97% (not shown). For analysis by FACS, cells were stained with directly conjugated antibodies.Fluorescence-activated cell sorting Splenocytes were isolated as described above. After blocking of FcR-binding sites, T and B cells were depleted using antibodies to CD3 and surface immunoglobulin, respectively. After incubation, goat anti-rat-coated magnetic beads (MACS) were added, and, after further incubation, cells were passed over a MACS column. The negative fraction was collected, washed, and stained with antibodies to CD3-fluorescein isothiocyanate (FITC), sIgM-FITC, CD11c-phycoerythrin (PE), and CD11b-APC. Cells were sorted on a FACS Star Plus (Becton Dickinson, Mountain View, CA). Gates were set to exclude debris and dead cells. T and B cells were further excluded by gating on the FITC-negative population. CD11c+CD11b+ and CD11c+ CD11b cells were collected. Cell purity usually exceeded
96% (data not shown). Alternatively, total DCs were isolated using
directly conjugated CD11c antibody-coated magnetic beads. After they
were washed, DCs were stained with antibodies to CD8 -PE, B220-APC, and a cocktail of FITC-labeled antibodies (CD11b, sIg, CD3). Gates were
set to exclude monocytes, T cells, and B cells, and DCs were sorted
based on the expression of CD8 and B220. Purity exceeded 96% (data
not shown).
Cell culture All cell cultures were performed in complete RPMI-1640 (Gibco Life Technologies, Grand Island, NY) supplemented with 5% fetal calf serum (Gibco), 2 mM L-glutamine, penicillin, streptomycin, and 1 mM HEPES buffer (Gibco).ELISA for detection of IFN- 2, 96-well flat-bottomed plates
(Costar, Corning, NY) were coated with 5 µg/mL sheep anti-mouse
IFN-/ (a kind gift from Schering-Plough Research Institute)
overnight at 4°C. After blocking with 3% bovine serum albumin in
PBS-Tween 20 for 1 hour at 37°C, plates were washed, and standard
rmIFN-2 (a kind gift from Schering-Plough) and culture
supernatants were added (standard rmIFN-2 had been
titrated in parallel against a known rmIFN-2 standard
provided by the National Institute of Allergy and Infectious Diseases
reference reagent repository). Plates were then incubated at
4°C overnight. After washing, a rat anti-mouse IFN-2
antibody (4E-A1 [IgG1]; Seikagaku America, Falmouth, MA) was added,
and plates were incubated at room temperature for 2 hours. After
washing, peroxidase-conjugated goat anti-rat immunoglobulin antibody
(Jackson Laboratory) was added, and plates were further incubated for 1 hour. After washing, plates were developed using tetramethylbenzidine
(TMB) substrate (Kirkegaard & Perry, Gaithersburg, MD) and were
stopped by the addition of 1 M sulfuric acid. For determination of
IL-12 (p70), plates were coated with 5 µg/mL rat anti-mouse IL-12
(p70) (Pharmingen) in PBS overnight. After blocking with bovine serum
albumin, supernatants and standard (rmIL-12, a kind gift from the
Genetics Institute, Cambridge, MA) was added. After incubation
overnight at 4°C, a biotinylated rat anti-mouse IL-12 (p40/p70)
antibody was added, and plates were incubated for 2 hours at room
temperature. After washing, extravidin-peroxidase (Sigma) was added and
plates were further incubated for 1 hour. Finally, plates were
developed using TMB substrate as above.
Electron microscopy Transmission electron microscopy was performed on sorted, CD11c+B220+ DCs. Cells were fixed in 2.5% glutaraldehyde in cacodylate buffer and postfixed in 1% OsO4 solution. Cells were dehydrated in a series of alcohol solutions and embedded in epoxide. Sections were examined using a JEOL 1210 microscope (JEOL, Peabody, MA).Staining for intracellular cytokines Cells were cultured for 24 hours together with HSV (10 plaque-forming units [PFU]/cell), and Brefeldin A was added during the last 5 hours of culture. For preparation and staining of cytospins, cells were spun onto slides and fixed for 20 minutes with 2% formaldehyde. Thereafter, cells were permeabilized for 20 minutes using PBS containing 0.55% saponin, 5% fetal calf serum (FCS), and 2 mM HEPES. All subsequent steps were performed in buffer containing saponin. After washing and blocking with 10% goat serum and 5% FCS for 30 minutes, primary rat anti-mouse IFN- (4E-A1; Seikagaku) or
an isotype-matched control antibody was added, and slides were incubated overnight. Rabbit anti-rat immunoglobulin antibodies (Dako
a/s, Glostrup, Denmark) were added, slides were incubated for 30 minutes and washed, and rat APAAP (Dako) was added. Incubation with
rabbit anti-rat immunoglobulin and rat APAAP was repeated. Finally,
slides were developed using Fast Red substrate (Dako).
For intracellular staining of IL-12 (p40), total CD11c+ DCs separated using MACS magnetic beads were cultured for 24 hours together with stimulatory anti-CD40 antibodies (25 µg/mL), SAC (10 µg/mL), or LPS (10µg/mL), and Brefeldin A (10 µg/mL) was added during the last 5 hours of culture. Surface staining of cells was performed using directly FITC-conjugated antibodies to CD11c, CD11b, and B220. After washing, cells were fixed in 2% PFA for 20 minutes and then permeabilized for 30 minutes in PBS containing 0.55% saponin, 5% FCS, and 2 mM HEPES. Directly PE-conjugated rat anti-mouse IL-12 (p40) antibodies (5 µg/mL) were added, and cells were further incubated for 30 minutes. After washing, cells were analyzed by flow cytometry. Measurement of T-cell stimulation For alloreaction, serial dilutions of irradiated DC (2000 cGy) that had been preactivated for 24 hours with stimulatory CD40 antibodies (IgM subclass; Pharmingen), and 2 × 105 total allogeneic CD4+ T cells were cultured for 6 days in round-bottomed, 96-well plates at a final volume of 200 µL. 3H-thymidine (1 µCi/well [0.037 MBq/well]) was added during the last 18 hours of culture. Proliferation was measured in a-scintillation counter.
Isolation of murine plasmacytoid dendritic cells from spleen and bone marrow Mice were mobilized for 7 days with Flt3L, a hematopoietic growth factor known to specifically expand DCs in vivo,9,11 and were killed 1 day after the last cytokine injection. Bone marrow cells were obtained by flushing resected femurs with PBS. Spleens were harvested and dispersed into single-cell suspensions. Total bone marrow and spleen cells were analyzed by flow cytometry and were found to express high numbers of CD11c+CD11b+ (myeloid) and CD11c+CD11b (lymphoid) DCs, respectively
(data not shown). Because a functional characteristic of human
plasmacytoid DCs (pDC2) is their high production of type 1 interferon
after encounter with pathogens, isolated bone marrow and spleen cells
were cultured overnight in the presence or absence of HSV or SAC. Cells
maintained in medium alone were used as control. After 24 hours of
culture, supernatants were harvested and analyzed for their
IFN-2 content by specific ELISA. As shown in Figure
1, bone marrow and, to a lesser degree,
spleen cells from mice that had been mobilized with Flt3L showed
greater production of IFN-2 in response to HSV than
nontreated mice. SAC induced low but reproducible levels of
IFN-2. A significantly greater number of mobilized DCs
could be detected when Flt3L was combined in vivo with GM-CSF, a factor known to support DC generation in vitro and also found to have an
additive effect when coadministered with Flt3L (P.B. et al, unpublished
observation, October 1999). Thus, the combination of
Flt3L+GM-CSF was used throughout the rest of this study.
Bone marrow cells and splenocytes derived from mice mobilized
with Flt3L+GM-CSF were isolated with magnetic beads directly conjugated
with antibodies to CD11c or CD11b. Cells were cultured in medium alone
or together with HSV or SAC, and supernatants were collected after 24 hours. The major IFN- Isolated splenocytes from day 7 Flt3L+GM-CSF-mobilized mice were then
sorted by FACS based on their expression of the
CD11c+CD11b+ (myeloid) and
CD11c+CD11b
Isolation of the major type 1 IFN-producing dendritic cell subset The CD11c+CD11b DC subset contains
a population of CD11c+CD8+ DCs that has been
previously shown to be the major IL-12-producing DC
subset11,19; however, CD11c+CD8+
DCs do not produce significant levels of IFN- after HSV infection. Because this suggested that there was heterogeneity within the CD11c+CD11b subset responsible for the
production of IFN- versus IL-12, DCs were sorted in a manner biased
by the phenotype recently reported for human pDC2.7 Human
pDC2 expresses CD45R, CD4, and the receptor for IL-3,7 but
FACS staining of murine CD11c+CD11b DCs using
antibodies recognizing IL-3R or CD4 showed only weak expression of
these antigens (Table 1). However,
antibodies to B220 (mouse CD45R) could be readily used to subdivide the
DC population further given that flow cytometry of total
CD11c+ DCs revealed a distinct subpopulation of
CD11c+B220+ cells in day 7 Flt3L+GM-CSF-mobilized mice (Figure 3A)
This was also observed to a lesser extent in mice treated with Flt3L
alone (data not shown). This CD11c+B220+
population could be found in bone marrow (BM), peripheral
blood, spleen, and lymph nodes of mobilized mice. Total
CD11c+ cells were isolated by directly conjugated
anti-CD11c magnetic beads and were sorted by flow cytometry using
fluorochrome-labeled antibodies to CD11b-FITC, CD8-PE, and B220-APC.
Anti-CD3- and sIg-FITC-labeled antibodies were used to further
exclude contaminating T and B cells. CD11b+ DCs (myeloid)
were excluded, and gates were set on CD8+- and
B220+- expressing cells, respectively (Figure 3B). FACS
analysis of sorted DCs showed that though both populations expressed
the CD11c antigen, the CD8+ population was typically
CD11cbright and the B220+ population was
CD11cdim (Table 1). B220+CD11c+
cells expressed Thy1.2 and low, but detectable, levels of IL-3R and
CD4. Furthermore, they expressed MHC class 1 (H2-Kd) and
class 2 (I-Ad/b) and accessory and adhesion molecules CD86,
CD11a, and CD54, but no myeloid antigens or CD19. The expression of
Gr-1 varied and was low or undetectable on freshly isolated cells, but
it was slightly up-regulated after overnight culture. The
CD8+CD11c+ DC population expressed high
levels of the MHC class II and CD86 molecules but lacked expression of
the CD4, IL-3R, and Thy1.2 antigens. Table 1 provides a summary of the
phenotype of the distinct murine DC subpopulations either directly
after FACS sorting or after overnight culture in complete
medium.
Freshly isolated and sorted spleen
CD11c+B220+ or
CD11c+CD8
The key markers used to isolate these 3 different mouse DC subsets are
detailed in Figure 5. To obtain myeloid
DCs, PE-conjugated antibodies to CD11b were used and
CD8
Mouse pDC develops into dendritic cells after culture and acquires the capacity to stimulate T cells Visual inspection of cytospins of the CD11c+B220+ DC subset revealed round cells lacking the characteristic dendritic morphology (Figure 6A). Human pDC2 differentiates into typical DC after culture in IL-3 and anti-CD40 antibodies.4 The culture of CD11c+B220+ DCs in media containing IL-3 and anti-CD40 for 2 days similarly revealed "classical" DCs with long protrusions and high MHC class II expression (Figure 6B). Interferon- could be detected by intracellular staining in
CD11c+B220+ DCs after culture with HSV
overnight (Figure 6C-D). IFN--producing cells were clearly
discernible by their strong cytoplasmic staining. Transmission electron
microscopy on freshly isolated, CD11c+B220+ DC
confirmed a well-developed endoplasmic reticulum and multiple mitochondria (Figure 6E). Of note, CD11c+B220+
DCs did not proliferate significantly in culture, regardless of the
combination of stimulating agents evaluated (ie, IL-3, SAC, CD40, HSV;
data not shown).
T-cell costimulation A defining characteristic for most DC subsets is T-cell stimulatory capacity. CD11c+B220+ DCs were examined for their capacity to stimulate T-cell proliferation. Isolated DCs were cultured overnight with stimulatory anti-CD40 antibodies (25 µg/mL). After washing, DCs were irradiated (2000 cGy), and serial dilutions of these APCs were cultured with total allogeneic CD4+ T cells (2 × 105/well). T-cell cocultures with CD11c+CD11b+ and CD11c+CD8+ DCs were generated in parallel.
As shown in Figure 6F, CD40-activated pDC induced T-cell proliferation
to levels similar to those observed for
CD11c+CD8+ lymphoid DCs. Human cultured pDC
has also been shown to induce a more potent T-cell stimulation than
freshly isolated.4 As expected, myeloid DCs induced
comparably strong T-cell stimulation.
In summary, the mouse plasmacytoid DCs exhibited a
CD11cdimB220+Thy1.2+ phenotype but
did not express the CD8
The current study describes the identification of the murine
natural type 1 IFN-producing cell20,21 or plasmacytoid
DC2,7 with the capacity to produce IFN- All murine DC subsets described to date express the CD11c
antigen, whereas those for human DCs do not. Murine pDC also express the CD45R (B220) antigen but at lower levels of CD4 and IL-3R than its
human counterpart. In addition, murine pDC expressed Thy1.2, a marker
expressed on T cells, but did not express the CD8 The existence of the natural IFN-producing cell has been known for some time.24 Although its hematopoietic origin is still a matter of debate, a recent study identified certain transcription factors that regulate T- and B-cell development and pDC2 development, but not DC1, suggesting a lymphoid origin for pDC2.16 Murine pDC express markers associated with lymphoid lineage cells but lack expression of myeloid markers (ie, CD11b) and express low levels of Gr-1 when isolated from BM but not from spleen. However, a low expression of Gr-1 could be found on splenic pDC after overnight culture. A BM-derived cell with the phenotype CD4lowGr-1low that also express low amounts of B220 and Thy-1 was isolated previously25 and found to give rise to both B- and T-lineage cells and to myeloid cells on adoptive transfer. Whether these are the same cells as the plasmacytoid DC described here merits further study. Murine pDC also do not express CD19, a marker that can be found coexpressed with CD11c on a small subset of cells in BM26; this may define an additional subset of DCs. Flt3L with or without GM-CSF was found to induce and enhance the
frequency of the CD11c+B220+DC subset that was
correlated with the mobilization of the IFN- Type 1 interferons produced by pDC would be anticipated to have
multiple effects on the immune system, including the up-regulation of
MHC class I on all cell types and functional activation of macrophages
and natural killer cells.20,21 IFN- Most recent data suggest the involvement of pDC in allergic asthma28 and in the autoimmune condition systemic lupus erythematosus.29 Targeting pDC in murine models for these diseases may provide important insights in their pathogenesis.
Dr W. J. Storkus is greatly acknowledged for support, help, and fruitful discussions. I thank Dr L. D. Falo for support, Mr. R. Lakomy for expert help with FACS sorting, and Drs S. Watkins and A. Bursick at the Center for Biological Imaging for help with electron microscopy.
Submitted March 9, 2001; accepted July 26, 2001.
The publication costs of this article were defrayed in part by page charge payment. Therefore, and solely to indicate this fact, this article is hereby marked "advertisement" in accordance with 18 U.S.C. section 1734.
Reprints: Pia Björck, Department of Dermatology, University of Pittsburgh, BST W 1546, 200 Lothrop St, Pittsburgh, PA 15213; e-mail: bjorckp{at}msx.upmc.edu.
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 A. Hosmalin and P. Lebon Type I interferon production in HIV-infected patients J. Leukoc. Biol., November 1, 2006; 80(5): 984 - 993. [Abstract] [Full Text] [PDF]
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U. Yrlid, V. Cerovic, S. Milling, C. D. Jenkins, J. Zhang, P. R. Crocker, L. S. Klavinskis, and G. G. MacPherson Plasmacytoid Dendritic Cells Do Not Migrate in Intestinal or Hepatic Lymph J. Immunol., November 1, 2006; 177(9): 6115 - 6121. [Abstract] [Full Text] [PDF]
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A. L. Blasius, E. Giurisato, M. Cella, R. D. Schreiber, A. S. Shaw, and M. Colonna Bone Marrow Stromal Cell Antigen 2 Is a Specific Marker of Type I IFN-Producing Cells in the Naive Mouse, but a Promiscuous Cell Surface Antigen following IFN Stimulation. J. Immunol., September 1, 2006; 177(5): 3260 - 3265. [Abstract] [Full Text] [PDF]
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B. C. Harman, J. P. Miller, N. Nikbakht, R. Gerstein, and D. Allman Mouse plasmacytoid dendritic cells derive exclusively from estrogen-resistant myeloid progenitors Blood, August 1, 2006; 108(3): 878 - 885. [Abstract] [Full Text] [PDF]
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 S. Delhaye, S. Paul, G. Blakqori, M. Minet, F. Weber, P. Staeheli, and T. Michiels Neurons produce type I interferon during viral encephalitis PNAS, May 16, 2006; 103(20): 7835 - 7840. [Abstract] [Full Text] [PDF]
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 A. Berhanu, J. Huang, S. M. Alber, S. C. Watkins, and W. J. Storkus Combinational FLt3 Ligand and Granulocyte Macrophage Colony-Stimulating Factor Treatment Promotes Enhanced Tumor Infiltration by Dendritic Cells and Antitumor CD8+ T-Cell Cross-priming but Is Ineffective as a Therapy. Cancer Res., May 1, 2006; 66(9): 4895 - 4903. [Abstract] [Full Text] [PDF]
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 A. Roghanian, S. E. Williams, T. A. Sheldrake, T. I. Brown, K. Oberheim, Z. Xing, S. E. M. Howie, and J.-M. Sallenave The Antimicrobial/Elastase Inhibitor Elafin Regulates Lung Dendritic Cells and Adaptive Immunity Am. J. Respir. Cell Mol. Biol., May 1, 2006; 34(5): 634 - 642. [Abstract] [Full Text] [PDF]
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U. Yrlid, S. W. F. Milling, J. L. Miller, S. Cartland, C. D. Jenkins, and G. G. MacPherson Regulation of Intestinal Dendritic Cell Migration and Activation by Plasmacytoid Dendritic Cells, TNF-{alpha} and Type 1 IFNs after Feeding a TLR7/8 Ligand J. Immunol., May 1, 2006; 176(9): 5205 - 5212. [Abstract] [Full Text] [PDF]
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H. Wang, J. Arp, X. Huang, W. Liu, S. Ramcharran, J. Jiang, B. Garcia, N. Kanai, W. Min, P. J. O'Connell, et al. Distinct Subsets of Dendritic Cells Regulate the Pattern of Acute Xenograft Rejection and Susceptibility to Cyclosporine Therapy J. Immunol., March 15, 2006; 176(6): 3525 - 3535. [Abstract] [Full Text] [PDF]
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T. Ito, H. Kanzler, O. Duramad, W. Cao, and Y.-J. Liu Specialization, kinetics, and repertoire of type 1 interferon responses by human plasmacytoid predendritic cells Blood, March 15, 2006; 107(6): 2423 - 2431. [Abstract] [Full Text] [PDF]
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E. O. Kvale, J. Dalgaard, F. Lund-Johansen, H. Rollag, L. Farkas, K. Midtvedt, F. L. Jahnsen, J. E. Brinchmann, and J. Olweus CD11c+ dendritic cells and plasmacytoid DCs are activated by human cytomegalovirus and retain efficient T cell-stimulatory capability upon infection Blood, March 1, 2006; 107(5): 2022 - 2029. [Abstract] [Full Text] [PDF]
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A. Suto, H. Nakajima, N. Tokumasa, H. Takatori, S.-i. Kagami, K. Suzuki, and I. Iwamoto Murine Plasmacytoid Dendritic Cells Produce IFN-{gamma} upon IL-4 Stimulation J. Immunol., November 1, 2005; 175(9): 5681 - 5689. [Abstract] [Full Text] [PDF]
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 F. Fallarino, C. Orabona, C. Vacca, R. Bianchi, S. Gizzi, C. Asselin-Paturel, M. C. Fioretti, G. Trinchieri, U. Grohmann, and P. Puccetti Ligand and cytokine dependence of the immunosuppressive pathway of tryptophan catabolism in plasmacytoid dendritic cells Int. Immunol., November 1, 2005; 17(11): 1429 - 1438. [Abstract] [Full Text] [PDF]
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R. Tussiwand, N. Onai, L. Mazzucchelli, and M. G. Manz Inhibition of Natural Type I IFN-Producing and Dendritic Cell Development by a Small Molecule Receptor Tyrosine Kinase Inhibitor with Flt3 Affinity J. Immunol., September 15, 2005; 175(6): 3674 - 3680. [Abstract] [Full Text] [PDF]
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A. Chahroudi, R. Chavan, N. Koyzr, E. K. Waller, G. Silvestri, and M. B. Feinberg Vaccinia Virus Tropism for Primary Hematolymphoid Cells Is Determined by Restricted Expression of a Unique Virus Receptor J. Virol., August 15, 2005; 79(16): 10397 - 10407. [Abstract] [Full Text] [PDF]
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M. Rossi and J. W. Young Human Dendritic Cells: Potent Antigen-Presenting Cells at the Crossroads of Innate and Adaptive Immunity J. Immunol., August 1, 2005; 175(3): 1373 - 1381. [Abstract] [Full Text] [PDF]
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G. S. Angelov, M. Tomkowiak, A. Marcais, Y. Leverrier, and J. Marvel Flt3 Ligand-Generated Murine Plasmacytoid and Conventional Dendritic Cells Differ in Their Capacity to Prime Naive CD8 T Cells and to Generate Memory Cells In Vivo J. Immunol., July 1, 2005; 175(1): 189 - 195. [Abstract] [Full Text] [PDF]
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Y. Omatsu, T. Iyoda, Y. Kimura, A. Maki, M. Ishimori, N. Toyama-Sorimachi, and K. Inaba Development of Murine Plasmacytoid Dendritic Cells Defined by Increased Expression of an Inhibitory NK Receptor, Ly49Q J. Immunol., June 1, 2005; 174(11): 6657 - 6662. [Abstract] [Full Text] [PDF]
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 C. Asselin-Paturel, G. Brizard, K. Chemin, A. Boonstra, A. O'Garra, A. Vicari, and G. Trinchieri Type I interferon dependence of plasmacytoid dendritic cell activation and migration J. Exp. Med., April 4, 2005; 201(7): 1157 - 1167. [Abstract] [Full Text] [PDF]
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Y. Kamogawa-Schifter, J. Ohkawa, S. Namiki, N. Arai, K.-i. Arai, and Y. Liu Ly49Q defines 2 pDC subsets in mice Blood, April 1, 2005; 105(7): 2787 - 2792. [Abstract] [Full Text] [PDF]
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G.-X. Yang, Z.-X. Lian, K. Kikuchi, Y.-J. Liu, A. A. Ansari, S. Ikehara, and M. E. Gershwin CD4- Plasmacytoid Dendritic Cells (pDCs) Migrate in Lymph Nodes by CpG Inoculation and Represent a Potent Functional Subset of pDCs J. Immunol., March 15, 2005; 174(6): 3197 - 3203. [Abstract] [Full Text] [PDF]
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B. Pulendran Variegation of the Immune Response with Dendritic Cells and Pathogen Recognition Receptors J. Immunol., March 1, 2005; 174(5): 2457 - 2465. [Abstract] [Full Text] [PDF]
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I. J. Fugier-Vivier, F. Rezzoug, Y. Huang, A. J. Graul-Layman, C. L. Schanie, H. Xu, P. M. Chilton, and S. T. Ildstad Plasmacytoid precursor dendritic cells facilitate allogeneic hematopoietic stem cell engraftment J. Exp. Med., February 7, 2005; 201(3): 373 - 383. [Abstract] [Full Text] [PDF]
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K. McKenna, A.-S. Beignon, and N. Bhardwaj Plasmacytoid Dendritic Cells: Linking Innate and Adaptive Immunity J. Virol., January 1, 2005; 79(1): 17 - 27. [Full Text] [PDF]
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M. A. Cannarile, N. Decanis, J. P. M. van Meerwijk, and T. Brocker The Role of Dendritic Cells in Selection of Classical and Nonclassical CD8+ T Cells In Vivo J. Immunol., October 15, 2004; 173(8): 4799 - 4805. [Abstract] [Full Text] [PDF]
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W. P. Halford, J. W. Balliet, and B. M. Gebhardt Re-Evaluating Natural Resistance to Herpes Simplex Virus Type 1 J. Virol., September 15, 2004; 78(18): 10086 - 10095. [Abstract] [Full Text] [PDF]
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F. Fallarino, C. Asselin-Paturel, C. Vacca, R. Bianchi, S. Gizzi, M. C. Fioretti, G. Trinchieri, U. Grohmann, and P. Puccetti Murine Plasmacytoid Dendritic Cells Initiate the Immunosuppressive Pathway of Tryptophan Catabolism in Response to CD200 Receptor Engagement J. Immunol., September 15, 2004; 173(6): 3748 - 3754. [Abstract] [Full Text] [PDF]
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G. Schlecht, S. Garcia, N. Escriou, A. A. Freitas, C. Leclerc, and G. Dadaglio Murine plasmacytoid dendritic cells induce effector/memory CD8+ T-cell responses in vivo after viral stimulation Blood, September 15, 2004; 104(6): 1808 - 1815. [Abstract] [Full Text] [PDF]
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F. Palamara, S. Meindl, M. Holcmann, P. Luhrs, G. Stingl, and M. Sibilia Identification and Characterization of pDC-Like Cells in Normal Mouse Skin and Melanomas Treated with Imiquimod J. Immunol., September 1, 2004; 173(5): 3051 - 3061. [Abstract] [Full Text] [PDF]
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F. Burke, A. J. Stagg, P. A. Bedford, N. English, and S. C. Knight IL-10-Producing B220+CD11c- APC in Mouse Spleen J. Immunol., August 15, 2004; 173(4): 2362 - 2372. [Abstract] [Full Text] [PDF]
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J. Diao, E. Winter, W. Chen, C. Cantin, and M. S. Cattral Characterization of Distinct Conventional and Plasmacytoid Dendritic Cell-Committed Precursors in Murine Bone Marrow J. Immunol., August 1, 2004; 173(3): 1826 - 1833. [Abstract] [Full Text] [PDF]
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S. Suzuki, K. Honma, T. Matsuyama, K. Suzuki, K. Toriyama, I. Akitoyo, K. Yamamoto, T. Suematsu, M. Nakamura, K. Yui, et al. From the Cover: Critical roles of interferon regulatory factor 4 in CD11bhighCD8{alpha}- dendritic cell development PNAS, June 15, 2004; 101(24): 8981 - 8986. [Abstract] [Full Text] [PDF]
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F.-X. Hubert, C. Voisine, C. Louvet, M. Heslan, and R. Josien Rat Plasmacytoid Dendritic Cells Are an Abundant Subset of MHC Class II+ CD4+CD11b-OX62- and Type I IFN-Producing Cells That Exhibit Selective Expression of Toll-Like Receptors 7 and 9 and Strong Responsiveness to CpG J. Immunol., June 15, 2004; 172(12): 7485 - 7494. [Abstract] [Full Text] [PDF]
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A. Blasius, W. Vermi, A. Krug, F. Facchetti, M. Cella, and M. Colonna A cell-surface molecule selectively expressed on murine natural interferon-producing cells that blocks secretion of interferon-alpha Blood, June 1, 2004; 103(11): 4201 - 4206. [Abstract] [Full Text] [PDF]
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M. Franchini, H. Hefti, S. Vollstedt, B. Glanzmann, M. Riesen, M. Ackermann, P. Chaplin, K. Shortman, and M. Suter Dendritic Cells from Mice Neonatally Vaccinated with Modified Vaccinia Virus Ankara Transfer Resistance against Herpes Simplex Virus Type I to Naive One-Week-Old Mice J. Immunol., May 15, 2004; 172(10): 6304 - 6312. [Abstract] [Full Text] [PDF]
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A. Mazzoni and D. M. Segal Controlling the Toll road to dendritic cell polarization J. Leukoc. Biol., May 1, 2004; 75(5): 721 - 730. [Abstract] [Full Text] [PDF]
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P. Bjorck Dendritic Cells Exposed to Herpes Simplex Virus In Vivo Do Not Produce IFN-{alpha} after Rechallenge with Virus In Vitro and Exhibit Decreased T Cell Alloreactivity J. Immunol., May 1, 2004; 172(9): 5396 - 5404. [Abstract] [Full Text] [PDF]
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B. J. Masten, G. K. Olson, D. F. Kusewitt, and M. F. Lipscomb Flt3 Ligand Preferentially Increases the Number of Functionally Active Myeloid Dendritic Cells in the Lungs of Mice J. Immunol., April 1, 2004; 172(7): 4077 - 4083. [Abstract] [Full Text] [PDF]
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G. Schiavoni, F. Mattei, P. Borghi, P. Sestili, M. Venditti, H. C. Morse III, F. Belardelli, and L. Gabriele ICSBP is critically involved in the normal development and trafficking of Langerhans cells and dermal dendritic cells Blood, March 15, 2004; 103(6): 2221 - 2228. [Abstract] [Full Text] [PDF]
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M. Salio, M. J. Palmowski, A. Atzberger, I. F. Hermans, and V. Cerundolo CpG-matured Murine Plasmacytoid Dendritic Cells Are Capable of In Vivo Priming of Functional CD8 T Cell Responses to Endogenous but Not Exogenous Antigens J. Exp. Med., February 17, 2004; 199(4): 567 - 579. [Abstract] [Full Text] [PDF]
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A. Dakic, Q.-x. Shao, A. D'Amico, M. O'Keeffe, W.-f. Chen, K. Shortman, and L. Wu Development of the Dendritic Cell System during Mouse Ontogeny J. Immunol., January 15, 2004; 172(2): 1018 - 1027. [Abstract] [Full Text] [PDF]
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C. Asselin-Paturel, G. Brizard, J.-J. Pin, F. Briere, and G. Trinchieri Mouse Strain Differences in Plasmacytoid Dendritic Cell Frequency and Function Revealed by a Novel Monoclonal Antibody J. Immunol., December 15, 2003; 171(12): 6466 - 6477. [Abstract] [Full Text] [PDF]
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G. F. Hoyne Notch signaling in the immune system J. Leukoc. Biol., December 1, 2003; 74(6): 971 - 981. [Abstract] [Full Text]
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 F. J. Hernandez-Ilizaliturri, V. Jupudy, J. Ostberg, E. Oflazoglu, A. Huberman, E. Repasky, and M. S. Czuczman Neutrophils Contribute to the Biological Antitumor Activity of Rituximab in a Non-Hodgkin's Lymphoma Severe Combined Immunodeficiency Mouse Model Clin. Cancer Res., December 1, 2003; 9(16): 5866 - 5873. [Abstract] [Full Text] [PDF]
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E. Hartmann, B. Wollenberg, S. Rothenfusser, M. Wagner, D. Wellisch, B. Mack, T. Giese, O. Gires, S. Endres, and G. Hartmann Identification and Functional Analysis of Tumor-Infiltrating Plasmacytoid Dendritic Cells in Head and Neck Cancer Cancer Res., October 1, 2003; 63(19): 6478 - 6487. [Abstract] [Full Text] [PDF]
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A. D'Amico and L. Wu The Early Progenitors of Mouse Dendritic Cells and Plasmacytoid Predendritic Cells Are within the Bone Marrow Hemopoietic Precursors Expressing Flt3 J. Exp. Med., July 21, 2003; 198(2): 293 - 303. [Abstract] [Full Text] [PDF]
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H. Karsunky, M. Merad, A. Cozzio, I. L. Weissman, and M. G. Manz Flt3 Ligand Regulates Dendritic Cell Development from Flt3+ Lymphoid and Myeloid-committed Progenitors to Flt3+ Dendritic Cells In Vivo J. Exp. Med., July 21, 2003; 198(2): 305 - 313. [Abstract] [Full Text] [PDF]
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A. D. Edwards, D. Chaussabel, S. Tomlinson, O. Schulz, A. Sher, and C. Reis e Sousa Relationships Among Murine CD11chigh Dendritic Cell Subsets as Revealed by Baseline Gene Expression Patterns J. Immunol., July 1, 2003; 171(1): 47 - 60. [Abstract] [Full Text] [PDF]
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 E. Muraille, C. De Trez, B. Pajak, F. A. Torrentera, P. De Baetselier, O. Leo, and Y. Carlier Amastigote Load and Cell Surface Phenotype of Infected Cells from Lesions and Lymph Nodes of Susceptible and Resistant Mice Infected with Leishmania major Infect. Immun., May 1, 2003; 71(5): 2704 - 2715. [Abstract] [Full Text] [PDF]
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C. M. Smith, G. T. Belz, N. S. Wilson, J. A. Villadangos, K. Shortman, F. R. Carbone, and W. R. Heath Cutting Edge: Conventional CD8{alpha}+ Dendritic Cells Are Preferentially Involved in CTL Priming After Footpad Infection with Herpes Simplex Virus-1 J. Immunol., May 1, 2003; 170(9): 4437 - 4440. [Abstract] [Full Text] [PDF]
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J.-F. Fonteneau, M. Gilliet, M. Larsson, I. Dasilva, C. Munz, Y.-J. Liu, and N. Bhardwaj Activation of influenza virus-specific CD4+ and CD8+ T cells: a new role for plasmacytoid dendritic cells in adaptive immunity Blood, May 1, 2003; 101(9): 3520 - 3526. [Abstract] [Full Text] [PDF]
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A. Krug, R. Veeraswamy, A. Pekosz, O. Kanagawa, E. R. Unanue, M. Colonna, and M. Cella Interferon-producing Cells Fail to Induce Proliferation of Naive T Cells but Can Promote Expansion and T Helper 1 Differentiation of Antigen-experienced Unpolarized T Cells J. Exp. Med., April 7, 2003; 197(7): 899 - 906. [Abstract] [Full Text] [PDF]
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M. Dalod, T. Hamilton, R. Salomon, T. P. Salazar-Mather, S. C. Henry, J. D. Hamilton, and C. A. Biron Dendritic Cell Responses to Early Murine Cytomegalovirus Infection: Subset Functional Specialization and Differential Regulation by Interferon {alpha}/{beta} J. Exp. Med., April 7, 2003; 197(7): 885 - 898. [Abstract] [Full Text] [PDF]
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A. M. Woltman and C. van Kooten Functional modulation of dendritic cells to suppress adaptive immune responses J. Leukoc. Biol., April 1, 2003; 73(4): 428 - 441. [Abstract] [Full Text] [PDF]
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V. Pullarkat, P. P. Lee, R. Scotland, V. Rubio, S. Groshen, C. Gee, R. Lau, J. Snively, S. Sian, S. L. Woulfe, et al. A Phase I Trial of SD-9427 (Progenipoietin) with a Multipeptide Vaccine for Resected Metastatic Melanoma Clin. Cancer Res., April 1, 2003; 9(4): 1301 - 1312. [Abstract] [Full Text] [PDF]
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S. Vollstedt, M. Franchini, H. P. Hefti, B. Odermatt, M. O'Keeffe, G. Alber, B. Glanzmann, M. Riesen, M. Ackermann, and M. Suter Flt3 Ligand-treated Neonatal Mice Have Increased Innate Immunity Against Intracellular Pathogens and Efficiently Control Virus Infections J. Exp. Med., March 3, 2003; 197(5): 575 - 584. [Abstract] [Full Text] [PDF]
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Z.-X. Lian, T. Okada, X.-S. He, H. Kita, Y.-J. Liu, A. A. Ansari, K. Kikuchi, S. Ikehara, and M. E. Gershwin Heterogeneity of Dendritic Cells in the Mouse Liver: Identification and Characterization of Four Distinct Populations J. Immunol., March 1, 2003; 170(5): 2323 - 2330. [Abstract] [Full Text] [PDF]
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T. Kambayashi, E. Assarsson, A. E. Lukacher, H.-G. Ljunggren, and P. E. Jensen Memory CD8+ T Cells Provide an Early Source of IFN-{gamma} J. Immunol., March 1, 2003; 170(5): 2399 - 2408. [Abstract] [Full Text] [PDF]
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T. Oida, X. Zhang, M. Goto, S. Hachimura, M. Totsuka, S. Kaminogawa, and H. L. Weiner CD4+CD25- T Cells That Express Latency-Associated Peptide on the Surface Suppress CD4+CD45RBhigh-Induced Colitis by a TGF-{beta}-Dependent Mechanism J. Immunol., March 1, 2003; 170(5): 2516 - 2522. [Abstract] [Full Text] [PDF]
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M. O'Keeffe, H. Hochrein, D. Vremec, B. Scott, P. Hertzog, L. Tatarczuch, and K. Shortman Dendritic cell precursor populations of mouse blood: identification of the murine homologues of human blood plasmacytoid pre-DC2 and CD11c+ DC1 precursors Blood, February 15, 2003; 101(4): 1453 - 1459. [Abstract] [Full Text] [PDF]
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A. Boonstra, C. Asselin-Paturel, M. Gilliet, C. Crain, G. Trinchieri, Y.-J. Liu, and A. O'Garra Flexibility of Mouse Classical and Plasmacytoid-derived Dendritic Cells in Directing T Helper Type 1 and 2 Cell Development: Dependency on Antigen Dose and Differential Toll-like Receptor Ligation J. Exp. Med., January 6, 2003; 197(1): 101 - 109. [Abstract] [Full Text] [PDF]
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J. Aliberti, O. Schulz, D. J. Pennington, H. Tsujimura, C. R. e Sousa, K. Ozato, and A. Sher Essential role for ICSBP in the in vivo development of murine CD8alpha + dendritic cells Blood, January 1, 2003; 101(1): 305 - 310. [Abstract] [Full Text] [PDF]
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P. Brawand, D. R. Fitzpatrick, B. W. Greenfield, K. Brasel, C. R. Maliszewski, and T. De Smedt Murine Plasmacytoid Pre-Dendritic Cells Generated from Flt3 Ligand-Supplemented Bone Marrow Cultures Are Immature APCs J. Immunol., December 15, 2002; 169(12): 6711 - 6719. [Abstract] [Full Text] [PDF]
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G. Schiavoni, F. Mattei, P. Sestili, P. Borghi, M. Venditti, H. C. Morse III, F. Belardelli, and L. Gabriele ICSBP Is Essential for the Development of Mouse Type I Interferon-producing Cells and for the Generation and Activation of CD8{alpha}+ Dendritic Cells J. Exp. Med., December 2, 2002; 196(11): 1415 - 1425. [Abstract] [Full Text] [PDF]
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A. Krug, R. Uppaluri, F. Facchetti, B. G. Dorner, K. C. F. Sheehan, R. D. Schreiber, M. Cella, and M. Colonna Cutting Edge: IFN-Producing Cells Respond to CXCR3 Ligands in the Presence of CXCL12 and Secrete Inflammatory Chemokines upon Activation J. Immunol., December 1, 2002; 169(11): 6079 - 6083. [Abstract] [Full Text] [PDF]
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B. J. Weigel, N. Nath, P. A. Taylor, A. Panoskaltsis-Mortari, W. Chen, A. M. Krieg, K. Brasel, and B. R. Blazar Comparative analysis of murine marrow-derived dendritic cells generated by Flt3L or GM-CSF/IL-4 and matured with immune stimulatory agents on the in vivo induction of antileukemia responses Blood, December 1, 2002; 100(12): 4169 - 4176. [Abstract] [Full Text] [PDF]
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M. O'Keeffe, H. Hochrein, D. Vremec, I. Caminschi, J. L. Miller, E. M. Anders, L. Wu, M. H. Lahoud, S. Henri, B. Scott, et al. Mouse Plasmacytoid Cells: Long-lived Cells, Heterogeneous in Surface Phenotype and Function, that Differentiate Into CD8+ Dendritic Cells Only after Microbial Stimulus J. Exp. Med., November 18, 2002; 196(10): 1307 - 1319. [Abstract] [Full Text] [PDF]
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I. Ferrero, W. Held, A. Wilson, F. Tacchini-Cottier, F. Radtke, and H. R. MacDonald Mouse CD11c+ B220+ Gr1+ plasmacytoid dendritic cells develop independently of the T-cell lineage Blood, September 26, 2002; 100(8): 2852 - 2857. [Abstract] [Full Text] [PDF]
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J. M.M. den Haan and M. J. Bevan Constitutive versus Activation-dependent Cross-Presentation of Immune Complexes by CD8+ and CD8- Dendritic Cells In Vivo J. Exp. Med., September 16, 2002; 196(6): 817 - 827. [Abstract] [Full Text] [PDF]
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J. H. Ahn, Y. Lee, C. Jeon, S.-J. Lee, B.-H. Lee, K. D. Choi, and Y.-S. Bae Identification of the genes differentially expressed in human dendritic cell subsets by cDNA subtraction and microarray analysis Blood, August 13, 2002; 100(5): 1742 - 1754. [Abstract] [Full Text] [PDF]
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P. Martin, G. M. del Hoyo, F. Anjuere, C. F. Arias, H. H. Vargas, A. Fernandez-L, V. Parrillas, and C. Ardavin Characterization of a new subpopulation of mouse CD8alpha + B220+ dendritic cells endowed with type 1 interferon production capacity and tolerogenic potential Blood, June 28, 2002; 100(2): 383 - 390. [Abstract] [Full Text] [PDF]
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M. Gilliet, A. Boonstra, C. Paturel, S. Antonenko, X.-L. Xu, G. Trinchieri, A. O'Garra, and Y.-J. Liu The Development of Murine Plasmacytoid Dendritic Cell Precursors Is Differentially Regulated by FLT3-ligand and Granulocyte/Macrophage Colony-Stimulating Factor J. Exp. Med., April 1, 2002; 195(7): 953 - 958. [Abstract] [Full Text] [PDF]
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Top
Abstract
Introduction
plasmacytoid
(p)DC2
and IL-10.
IL-3, but not GM-CSF, also induces the maturation of pDC2. Once mature,
these APCs can promote TH2 cytokine production (ie, IL-4, IL-5, and
IL-10).
subclasses of dendritic cells direct the development of distinct T helper cells in vivo.
J Exp Med.
1999;189:587-592