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n4:pmcid
PMC0
bibo:doi
10.1038%2Fni.1790
n5:contains
_:vb6102621 _:vb6102596 _:vb6102636 _:vb6102567
Subject Item
_:vb6102567
rdf:type
n5:Section
dc:title
introduction
n5:contains
_:vb6102592 _:vb6102593 _:vb6102594 _:vb6102595 _:vb6102584 _:vb6102585 _:vb6102586 _:vb6102587 _:vb6102588 _:vb6102589 _:vb6102590 _:vb6102591 _:vb6102576 _:vb6102577 _:vb6102578 _:vb6102579 _:vb6102580 _:vb6102581 _:vb6102582 _:vb6102583 _:vb6102568 _:vb6102569 _:vb6102570 _:vb6102571 _:vb6102572 _:vb6102573 _:vb6102574 _:vb6102575
Subject Item
_:vb6102568
rdf:type
n4:Context
rdf:value
afcsid=A000706) is a well established negative regulator of T cell function >>1<<. CTLA-4 is rapidly expressed on the surface of T cells following activation and is highly upregulated by engagement of the co-stimulatory molecule CD28 1.
n4:mentions
n2:7882171
Subject Item
_:vb6102569
rdf:type
n4:Context
rdf:value
CTLA-4 is rapidly expressed on the surface of T cells following activation and is highly upregulated by engagement of the co-stimulatory molecule CD28 >>1<<. CTLA-4 ligation antagonizes early T cell activation leading to decreased interleukin 2 (IL-2) production, inhibition of cell cycle progression and modulation of TCR signaling 2,3. CTLA-4 and CD28 shares the ligands B7-1 (also called
n4:mentions
n2:7882171
Subject Item
_:vb6102570
rdf:type
n4:Context
rdf:value
CTLA-4 ligation antagonizes early T cell activation leading to decreased interleukin 2 (IL-2) production, inhibition of cell cycle progression and modulation of TCR signaling >>2<<,3. CTLA-4 and CD28 shares the ligands B7-1 (also called CD80) and B7-2 (also called CD86) 4. Mice deficient for CTLA-4 develop lymphoproliferative disease and die within 3-4 weeks of birth 5, adding further evidence for the critical role
n4:mentions
n2:11035773
Subject Item
_:vb6102571
rdf:type
n4:Context
rdf:value
CTLA-4 ligation antagonizes early T cell activation leading to decreased interleukin 2 (IL-2) production, inhibition of cell cycle progression and modulation of TCR signaling 2,>>3<<. CTLA-4 and CD28 shares the ligands B7-1 (also called CD80) and B7-2 (also called CD86) 4. Mice deficient for CTLA-4 develop lymphoproliferative disease and die within 3-4 weeks of birth 5, adding further evidence for the critical role of
n4:mentions
n2:12515820
Subject Item
_:vb6102572
rdf:type
n4:Context
rdf:value
Mice deficient for CTLA-4 develop lymphoproliferative disease and die within 3-4 weeks of birth >>5<<, adding further evidence for the critical role of CTLA-4 in controlling T cell responses and immune homeostasis.
n4:mentions
n2:7584144
Subject Item
_:vb6102573
rdf:type
n4:Context
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Programmed death-1 (PD-1) is an inhibitory molecule found on the surface of activated B and T cells and has been implicated in immune tolerance >>6<<. PD-1 (also called CD279) is a member of the CD28 and CTLA-4 immunoglobulin superfamily and interacts with two B7 family ligands, PD-L1 (also called CD274) and PD-L2 (also called CD273) 4.
n4:mentions
n2:9796923
Subject Item
_:vb6102574
rdf:type
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rdf:value
PD-L1 is widely distributed on leukocytes, non-hematopoietic cells, and in non-lymphoid tissues including pancreatic islets, while PD-L2 is expressed exclusively on dendritic cells (DC) and monocytes >>7<<,8. The PD-1 signaling pathway limits viral clearance during chronic infections by creating an unresponsive state in virus-specific CD8+ T cells termed “exhaustion” 9. In an autoimmune setting, PD-1 is essential for maintaining T cell
n4:mentions
n2:12421930
Subject Item
_:vb6102575
rdf:type
n4:Context
rdf:value
PD-L1 is widely distributed on leukocytes, non-hematopoietic cells, and in non-lymphoid tissues including pancreatic islets, while PD-L2 is expressed exclusively on dendritic cells (DC) and monocytes 7,>>8<<. The PD-1 signaling pathway limits viral clearance during chronic infections by creating an unresponsive state in virus-specific CD8+ T cells termed “exhaustion” 9. In an autoimmune setting, PD-1 is essential for maintaining T cell anergy
n4:mentions
n2:12161284
Subject Item
_:vb6102576
rdf:type
n4:Context
rdf:value
The PD-1 signaling pathway limits viral clearance during chronic infections by creating an unresponsive state in virus-specific CD8+ T cells termed “exhaustion” >>9<<. In an autoimmune setting, PD-1 is essential for maintaining T cell anergy and preventing autoimmunity 10,11. Genetic deletion of PD-1 results in profound and complex multi-organ autoimmune destruction 4. Similarly, blocking PD-1-PD-L1
n4:mentions
n2:16382236
Subject Item
_:vb6102577
rdf:type
n4:Context
rdf:value
In an autoimmune setting, PD-1 is essential for maintaining T cell anergy and preventing autoimmunity >>10<<,11. Genetic deletion of PD-1 results in profound and complex multi-organ autoimmune destruction 4. Similarly, blocking PD-1-PD-L1 interactions accelerates spontaneous autoimmune diabetes 12 and reverses immune tolerance 10,11. Together
n4:mentions
n2:17116737
Subject Item
_:vb6102578
rdf:type
n4:Context
rdf:value
In an autoimmune setting, PD-1 is essential for maintaining T cell anergy and preventing autoimmunity 10,>>11<<. Genetic deletion of PD-1 results in profound and complex multi-organ autoimmune destruction 4. Similarly, blocking PD-1-PD-L1 interactions accelerates spontaneous autoimmune diabetes 12 and reverses immune tolerance 10,11. Together these
n4:mentions
n2:16606670
Subject Item
_:vb6102579
rdf:type
n4:Context
rdf:value
Similarly, blocking PD-1-PD-L1 interactions accelerates spontaneous autoimmune diabetes >>12<< and reverses immune tolerance 10,11.
n4:mentions
n2:12847137
Subject Item
_:vb6102580
rdf:type
n4:Context
rdf:value
Similarly, blocking PD-1-PD-L1 interactions accelerates spontaneous autoimmune diabetes 12 and reverses immune tolerance >>10<<,11. Together these data support a central role for PD-1 as a major inhibitory pathway controlling immunity and suggest that exhaustion and peripheral tolerance are linked mechanistically.
n4:mentions
n2:17116737
Subject Item
_:vb6102581
rdf:type
n4:Context
rdf:value
Similarly, blocking PD-1-PD-L1 interactions accelerates spontaneous autoimmune diabetes 12 and reverses immune tolerance 10,>>11<<. Together these data support a central role for PD-1 as a major inhibitory pathway controlling immunity and suggest that exhaustion and peripheral tolerance are linked mechanistically.
n4:mentions
n2:16606670
Subject Item
_:vb6102582
rdf:type
n4:Context
rdf:value
PD-L1 may act passively by competing directly with CD28 for B7-1 binding >>13<<. In addition, PD-1 may directly recruit phosphatases such as SHP-1, SHP-2 and PP2A, which interfere with TCR signaling 4,14-16.
n4:mentions
n2:17629517
Subject Item
_:vb6102583
rdf:type
n4:Context
rdf:value
In addition, PD-1 may directly recruit phosphatases such as SHP-1, SHP-2 and PP2A, which interfere with TCR signaling 4,>>14<<-16. However, the effects of PD-1 engagement during T cell activation are unclear as in vitro models do not adequately demonstrate the inhibitory activity of PD-1 17. In vivo studies of PD-1–mediated inhibition have been limited to global
n4:mentions
n2:16227604 n2:10694513 n2:14713717
Subject Item
_:vb6102584
rdf:type
n4:Context
rdf:value
However, the effects of PD-1 engagement during T cell activation are unclear as in vitro models do not adequately demonstrate the inhibitory activity of PD-1 >>17<<. In vivo studies of PD-1–mediated inhibition have been limited to global effects on immunity with little direct evidence that PD-1 engagement directly controls T cell signaling.
n4:mentions
n2:18759926
Subject Item
_:vb6102585
rdf:type
n4:Context
rdf:value
Previously, we have used a well-defined, antigen-specific tolerance model to examine the cellular basis of immunological tolerance using antigen-pulsed and fixed antigen-presenting cells (APC) >>10<<. We determined that the administration of islet antigen peptide mimic, p31, coupled to chemically fixed APCs reversed diabetes and induced a robust, long-term inactivation of islet-specific, BDC2.5 TCR transgenic T cells.
n4:mentions
n2:17116737
Subject Item
_:vb6102586
rdf:type
n4:Context
rdf:value
Although both PD-1 and CTLA-4 interactions were critical for the induction of tolerance, the long term maintenance of the anergic state depended on PD-1-PD-L1 interactions, but not CTLA-4 interactions within the inflamed tissue >>10<<. These results implied that a cell-intrinsic mechanism maintains tolerance in the setting of continual exposure to autoantigen.
n4:mentions
n2:17116737
Subject Item
_:vb6102587
rdf:type
n4:Context
rdf:value
Recent imaging advances have allowed the use of multi-photon laser-scanning microscopy (MPLSM) to visualize T cells in lymph nodes (LN) and non-lymphoid tissues in vivo on the single cell level >>18<<. Several groups have described the highly dynamic movement of CD4+ and CD8+ T cells, as well as DCs during T cell priming and tolerance induction 19-23.
n4:mentions
n2:16799470
Subject Item
_:vb6102588
rdf:type
n4:Context
rdf:value
Several groups have described the highly dynamic movement of CD4+ and CD8+ T cells, as well as DCs during T cell priming and tolerance induction >>19<<-23. Naïve circulating T cells enter the LN through high endothelial venules and move on a network of stromal cells called fibroblastic reticular cells (FRCs), which define the potential location and migratory range of T cells 24. The FRC
n4:mentions
n2:15924144 n2:12016203 n2:15516925 n2:15357946 n2:14712275
Subject Item
_:vb6102589
rdf:type
n4:Context
rdf:value
Naïve circulating T cells enter the LN through high endothelial venules and move on a network of stromal cells called fibroblastic reticular cells (FRCs), which define the potential location and migratory range of T cells >>24<<. The FRC also provide a substrate for resident DCs to sample and display antigen to passing T cells. As T cells migrate along the FRC highway they probe DCs for signals and antigen 24. Initial interactions may be transient, but antigen
n4:mentions
n2:17112751
Subject Item
_:vb6102590
rdf:type
n4:Context
rdf:value
As T cells migrate along the FRC highway they probe DCs for signals and antigen >>24<<. Initial interactions may be transient, but antigen recognition results in T cell swarming, formation of stable T cell-DC conjugates, and T cell arrest 25,26. TCR ligation results in ‘stop signals’ that decrease T cell motility and are
n4:mentions
n2:17112751
Subject Item
_:vb6102591
rdf:type
n4:Context
rdf:value
Initial interactions may be transient, but antigen recognition results in T cell swarming, formation of stable T cell-DC conjugates, and T cell arrest >>25<<,26. TCR ligation results in ‘stop signals’ that decrease T cell motility and are required for stable T cell-DC conjugate formation and development of the immunological synapse 21,26. Prolonged T cell-DC interactions are critical for full
n4:mentions
n2:8630728
Subject Item
_:vb6102592
rdf:type
n4:Context
rdf:value
Initial interactions may be transient, but antigen recognition results in T cell swarming, formation of stable T cell-DC conjugates, and T cell arrest 25,>>26<<. TCR ligation results in ‘stop signals’ that decrease T cell motility and are required for stable T cell-DC conjugate formation and development of the immunological synapse 21,26. Prolonged T cell-DC interactions are critical for full T
n4:mentions
n2:9108078
Subject Item
_:vb6102593
rdf:type
n4:Context
rdf:value
TCR ligation results in ‘stop signals’ that decrease T cell motility and are required for stable T cell-DC conjugate formation and development of the immunological synapse >>21<<,26. Prolonged T cell-DC interactions are critical for full T cell activation, proliferation and cytokine production 27-29. However, T cell movement and velocity during the long term maintenance of tolerance has not been well described.
n4:mentions
n2:14712275
Subject Item
_:vb6102594
rdf:type
n4:Context
rdf:value
TCR ligation results in ‘stop signals’ that decrease T cell motility and are required for stable T cell-DC conjugate formation and development of the immunological synapse 21,>>26<<. Prolonged T cell-DC interactions are critical for full T cell activation, proliferation and cytokine production 27-29. However, T cell movement and velocity during the long term maintenance of tolerance has not been well described.
n4:mentions
n2:9108078
Subject Item
_:vb6102595
rdf:type
n4:Context
rdf:value
Prolonged T cell-DC interactions are critical for full T cell activation, proliferation and cytokine production >>27<<-29. However, T cell movement and velocity during the long term maintenance of tolerance has not been well described.
n4:mentions
n2:18275834 n2:12835480 n2:14688337
Subject Item
_:vb6102596
rdf:type
n5:Section
dc:title
results
n5:contains
_:vb6102612 _:vb6102613 _:vb6102614 _:vb6102615 _:vb6102608 _:vb6102609 _:vb6102610 _:vb6102611 _:vb6102620 _:vb6102616 _:vb6102617 _:vb6102618 _:vb6102619 _:vb6102597 _:vb6102598 _:vb6102599 _:vb6102604 _:vb6102605 _:vb6102606 _:vb6102607 _:vb6102600 _:vb6102601 _:vb6102602 _:vb6102603
Subject Item
_:vb6102597
rdf:type
n4:Context
rdf:value
Insulin-coupled, ethylene carbodiimide (ECDI) fixed APCs have a profound effect on the development of spontaneous type 1 diabetes (T1D) in the non-obese diabetic (NOD) mouse >>10<<. A single injection of the antigen-pulsed fixed splenocytes prevents and even reverses T1D after disease onset.
n4:mentions
n2:17116737
Subject Item
_:vb6102598
rdf:type
n4:Context
rdf:value
We previously identified a critical role for PD-1 but not CTLA-4 in the maintenance of tolerance induced by injection of insulin-coupled fixed splenocytes >>10<<. Herein, we adapted this model using islet antigen-specific BDC2.5 TCR transgenic T cells to define the role of PD-1 and CTLA-4 in the maintenance of peripheral tolerance.
n4:mentions
n2:17116737
Subject Item
_:vb6102599
rdf:type
n4:Context
rdf:value
Injection of splenocytes coupled to the p31 peptide (Ag-SP, p31-SP), an islet antigen mimetope recognized by the BDC2.5 TCR >>30<<, prevented autoimmune diabetes and resulted in decreased T cell proliferation and cytokine production (Fig.
n4:mentions
n2:11145667
Subject Item
_:vb6102600
rdf:type
n4:Context
rdf:value
Previous work demonstrated that TCR-induced calcium signals are associated with T cell motility and stable interactions with APCs >>25<<. BDC2.5 T cells isolated from p31-SP-tolerized, but not SHAM-SP-injected mice, did not flux calcium in response to strong secondary TCR stimulation with anti-CD3 (Fig. 1b). Furthermore, phosphorylation of the TCR ζ-chain and the MAP
n4:mentions
n2:8630728
Subject Item
_:vb6102601
rdf:type
n4:Context
rdf:value
Antigen-specific T cells swarm and cluster when they enter antigen-containing PLNs >>31<<. Thus, we compared the migration behavior of activated control BDC2.5 T cells with that of tolerized BDC2.5 T cells. BDC2.5 Thy1.1 CD4+ T cells from SHAM-SP or p31-SP-treated mice were harvested, CMTMR-labeled and transferred into NOD
n4:mentions
n2:16311599
Subject Item
_:vb6102602
rdf:type
n4:Context
rdf:value
This CD11c-YFP mouse allowed tracking of the in vivo movements of CD11c+ DC >>32<<. The velocity and displacement of control and tolerized T cells in islet antigen-bearing PLN 18-24 hours following T cell transfer was compared.
n4:mentions
n2:15543150
Subject Item
_:vb6102603
rdf:type
n4:Context
rdf:value
2a and Supplementary Movie 1), similar to what has been previously reported >>31<<. In contrast, the tolerant T cells did not swarm in antigen-containing PLN, and exhibited a mean velocity of 5.2 ± 0.4 μm/min (Fig.
n4:mentions
n2:16311599
Subject Item
_:vb6102604
rdf:type
n4:Context
rdf:value
2a and Supplementary Movie 2), similar to what has been observed in antigen non-bearing LN >>31<<.
n4:mentions
n2:16311599
Subject Item
_:vb6102605
rdf:type
n4:Context
rdf:value
As PD-L1, but not PD-L2, is required for the maintenance of tolerance, we injected PD-L1-blocking antibodies >>10<<. PD-L1 blockade led the tolerant T cells to slow and swarm with a velocity of 1.4 ± 0.1 μm/min, as compared to the 5.2 μm/min velocity observed after isotype control antibody treatment (P<0.0001) (Fig. 2a and Supplementary Movie 3). PD-1
n4:mentions
n2:17116737
Subject Item
_:vb6102606
rdf:type
n4:Context
rdf:value
2b, M = 0.32 ± 0.16 μm2 min-1) >>22<<,33,34. In contrast, the displacement of tolerized BDC2.5 T cells was significantly higher (M = 6.65 ± 0.21 μm2 min-1, Fig.
n4:mentions
n2:12016203
Subject Item
_:vb6102607
rdf:type
n4:Context
rdf:value
2b, M = 0.32 ± 0.16 μm2 min-1) 22,>>33<<,34. In contrast, the displacement of tolerized BDC2.5 T cells was significantly higher (M = 6.65 ± 0.21 μm2 min-1, Fig.
n4:mentions
n2:15357943
Subject Item
_:vb6102608
rdf:type
n4:Context
rdf:value
2b, M = 0.32 ± 0.16 μm2 min-1) 22,33,>>34<<. In contrast, the displacement of tolerized BDC2.5 T cells was significantly higher (M = 6.65 ± 0.21 μm2 min-1, Fig.
n4:mentions
n2:12415310
Subject Item
_:vb6102609
rdf:type
n4:Context
rdf:value
This is likely due to the fact that tolerized T cells express high amounts of PD-1 on their surface, while naïve T cells do not >>10<<,35. These results reinforce the notion that antigen is required for the stop signal and that PD-1 normally functions to inhibit T cell activation by preventing T cell arrest.
n4:mentions
n2:17116737
Subject Item
_:vb6102610
rdf:type
n4:Context
rdf:value
This is likely due to the fact that tolerized T cells express high amounts of PD-1 on their surface, while naïve T cells do not 10,>>35<<. These results reinforce the notion that antigen is required for the stop signal and that PD-1 normally functions to inhibit T cell activation by preventing T cell arrest.
n4:mentions
n2:8671666
Subject Item
_:vb6102611
rdf:type
n4:Context
rdf:value
PD-1–mediated control of immune responses depends on interactions between PD-1 on T cells and PD-L1 ligand in tissues >>10<<,11. Thus, we sought to determine if the effects of PD-1-PD-L1 blockade had similar effects in the autoimmune target tissue, the pancreatic islets, as was the case in antigen draining PLN.
n4:mentions
n2:17116737
Subject Item
_:vb6102612
rdf:type
n4:Context
rdf:value
PD-1–mediated control of immune responses depends on interactions between PD-1 on T cells and PD-L1 ligand in tissues 10,>>11<<. Thus, we sought to determine if the effects of PD-1-PD-L1 blockade had similar effects in the autoimmune target tissue, the pancreatic islets, as was the case in antigen draining PLN.
n4:mentions
n2:16606670
Subject Item
_:vb6102613
rdf:type
n4:Context
rdf:value
The slope of these two curves is a straight line representing free and random walking within the target tissue and does not correspond to directed migration >>33<<. PD-L1 blockade, however, attenuated T cell motility and decreased T cell displacement when compared to isotype control antibody-treated animals (Fig. 4b, M= 0.5 ± 0.3 μm2 min-1). Tracking individual cells confirmed these results (Fig.
n4:mentions
n2:15357943
Subject Item
_:vb6102614
rdf:type
n4:Context
rdf:value
Prolonged interactions between T cells and DCs is critical for full T cell activation >>27<<,29. We hypothesized that PD-1 normally functions to prevent the stable T cell-DC contacts that are required for full T cell activation. To test this notion, we used the adoptive transfer and imaging system described above to measure the
n4:mentions
n2:12835480
Subject Item
_:vb6102615
rdf:type
n4:Context
rdf:value
Prolonged interactions between T cells and DCs is critical for full T cell activation 27,>>29<<. We hypothesized that PD-1 normally functions to prevent the stable T cell-DC contacts that are required for full T cell activation. To test this notion, we used the adoptive transfer and imaging system described above to measure the
n4:mentions
n2:18275834
Subject Item
_:vb6102616
rdf:type
n4:Context
rdf:value
1) and Erk phosphorylation >>10<<,36. However, treatment of anergic T cells with PMA and ionophore can reverse tolerance and promote T cell activation (Fig.
n4:mentions
n2:17116737
Subject Item
_:vb6102617
rdf:type
n4:Context
rdf:value
1) and Erk phosphorylation 10,>>36<<. However, treatment of anergic T cells with PMA and ionophore can reverse tolerance and promote T cell activation (Fig.
n4:mentions
n2:12086671
Subject Item
_:vb6102618
rdf:type
n4:Context
rdf:value
Tolerized T cells did not exhibit Er phoksphorylation after in vivo APC engagement >>36<<-38 (Fig. 5e). In contrast, treatment with anti-PD-L1 but not isotype control antibody during exposure of the tolerized T cells to antigen-bearing APCs resulted in substantial Erk phosphorylation in tolerized T cells (Fig. 5f). These
n4:mentions
n2:18056342 n2:8638107 n2:12086671
Subject Item
_:vb6102619
rdf:type
n4:Context
rdf:value
Ag-SP tolerance resulted in a decrease in inflammatory cytokines including IL-2 and IFN-γ within the PLN >>10<<. Here we bred antigen-specific NOD.BDC2.5 TCR transgenic mice to Yeti mice, which express a YFP reporter cassette under the control of the IFN-γ promoter 31. Tolerized BDC2.5.Thy1.1.Yeti T cells were transferred to recipient NOD mice and
n4:mentions
n2:17116737
Subject Item
_:vb6102620
rdf:type
n4:Context
rdf:value
BDC2.5 TCR transgenic mice to Yeti mice, which express a YFP reporter cassette under the control of the IFN-γ promoter >>31<<. Tolerized BDC2.5.Thy1.1.Yeti T cells were transferred to recipient NOD mice and received isotype control antibody, anti-PD-L1, or anti-CTLA-4. PD-L1 blockade abrogated tolerance, and the majority of CD4+ T cells isolated directly from
n4:mentions
n2:16311599
Subject Item
_:vb6102621
rdf:type
n5:Section
dc:title
methods
n5:contains
_:vb6102622 _:vb6102623 _:vb6102624 _:vb6102625 _:vb6102626 _:vb6102627 _:vb6102628 _:vb6102629 _:vb6102630 _:vb6102631 _:vb6102632 _:vb6102633 _:vb6102634 _:vb6102635
Subject Item
_:vb6102622
rdf:type
n4:Context
rdf:value
Bell (University of Chicago, Chicago, IL) >>46<< and were backcrossed for more than 12 generations to NOD mice.
n4:mentions
n2:12388130
Subject Item
_:vb6102623
rdf:type
n4:Context
rdf:value
Nussenzweig (The Rockefeller University, New York, NY) >>32<< mice were backcrossed for more than 10 generations to NOD.
n4:mentions
n2:15543150
Subject Item
_:vb6102624
rdf:type
n4:Context
rdf:value
Sharpe (Harvard, Boston, MA) >>11<< and were backcrossed for 10 generations to NOD. Female NOD mice were purchased from Taconic.
n4:mentions
n2:16606670
Subject Item
_:vb6102625
rdf:type
n4:Context
rdf:value
Benoist and D. Mathis (Harvard Medical School, Boston, MA) >>47<< and were crossed to NOD.Thy1.1.
n4:mentions
n2:8402882
Subject Item
_:vb6102626
rdf:type
n4:Context
rdf:value
BDC2.5.Thy1.1.Yeti mice >>31<<. NOD.SCID mice were purchased from The Jackson Laboratory.
n4:mentions
n2:16311599
Subject Item
_:vb6102627
rdf:type
n4:Context
rdf:value
Anti-Phospho-p44/42 MAPK (Thr202/Tyr204) (197G2) rabbit mAb was purchased from Cell Signaling Technology >>48<<. Anti–PD-1 (RMP1-14), anti–PD-L1 (MIH5, MIH6), and anti–PD-L2 (TY25) were made as described previously 7,10. Anti-CTLA-4 (UC4F10), and anti-CTLA-4 Fab fragments were made as described previously 1. Mice were injected intraperitoneally
n4:mentions
n2:16774041
Subject Item
_:vb6102628
rdf:type
n4:Context
rdf:value
Anti–PD-1 (RMP1-14), anti–PD-L1 (MIH5, MIH6), and anti–PD-L2 (TY25) were made as described previously >>7<<,10. Anti-CTLA-4 (UC4F10), and anti-CTLA-4 Fab fragments were made as described previously 1. Mice were injected intraperitoneally with 250 μg anti–PD-L1, anti–PD-L2, anti-B7-1, anti-PD-1, or anti-CTLA-4 as indicated.
n4:mentions
n2:12421930
Subject Item
_:vb6102629
rdf:type
n4:Context
rdf:value
Anti–PD-1 (RMP1-14), anti–PD-L1 (MIH5, MIH6), and anti–PD-L2 (TY25) were made as described previously 7,>>10<<. Anti-CTLA-4 (UC4F10), and anti-CTLA-4 Fab fragments were made as described previously 1. Mice were injected intraperitoneally with 250 μg anti–PD-L1, anti–PD-L2, anti-B7-1, anti-PD-1, or anti-CTLA-4 as indicated.
n4:mentions
n2:17116737
Subject Item
_:vb6102630
rdf:type
n4:Context
rdf:value
Anti-CTLA-4 (UC4F10), and anti-CTLA-4 Fab fragments were made as described previously >>1<<. Mice were injected intraperitoneally with 250 μg anti–PD-L1, anti–PD-L2, anti-B7-1, anti-PD-1, or anti-CTLA-4 as indicated.
n4:mentions
n2:7882171
Subject Item
_:vb6102631
rdf:type
n4:Context
rdf:value
injections of 50 × 106 chemically treated antigen-coupled syngeneic splenocytes (p31 or SHAM control), as described previously >>10<<.
n4:mentions
n2:17116737
Subject Item
_:vb6102632
rdf:type
n4:Context
rdf:value
molecule and cytokine expression, cells were labeled with predetermined optimal antibody concentrations according to the manufacturer's staining protocol and 0.5 × 106 events in the CD4+ gate were acquired, as described previously >>10<<. Data acquisition was performed on an LSRII flow cytometer and analyzed using FACSDiva software (Becton Dickinson).
n4:mentions
n2:17116737
Subject Item
_:vb6102633
rdf:type
n4:Context
rdf:value
Islets were isolated as previously reported >>49<<. Briefly, a 3-ml collagenase P (Roche) solution (0.75 mg/ml) was injected into the pancreatic duct of 4-wk-old NOD.
n4:mentions
n2:7482727
Subject Item
_:vb6102634
rdf:type
n4:Context
rdf:value
A custom built resonant-scanning instrument based on published designs containing four–photomultiplier tubes operating at video rate was used for multi-photon microscopy >>31<<. For imaging of T cells, BDC2.5 CD4+ CD90.1 T cells were sorted and labeled with Cell Tracker™ Orange CMTMR (5-(and-6)-(((4-chloromethyl) benzoyl) amino) tetramethyl rhodamine) (Invitrogen) and transferred to recipient mice.
n4:mentions
n2:16311599
Subject Item
_:vb6102635
rdf:type
n4:Context
rdf:value
The motility coefficient M=x2/6t was calculated from the slope (x/t1/2) obtained by regression analysis of the mean displacement (x) versus the square root of time (t1/2) as described previously >>33<<.
n4:mentions
n2:15357943
Subject Item
_:vb6102636
rdf:type
n5:Section
dc:title
discussion
n5:contains
_:vb6102644 _:vb6102645 _:vb6102646 _:vb6102647 _:vb6102640 _:vb6102641 _:vb6102642 _:vb6102643 _:vb6102652 _:vb6102653 _:vb6102654 _:vb6102655 _:vb6102648 _:vb6102649 _:vb6102650 _:vb6102651 _:vb6102637 _:vb6102638 _:vb6102639 _:vb6102660 _:vb6102661 _:vb6102662 _:vb6102656 _:vb6102657 _:vb6102658 _:vb6102659
Subject Item
_:vb6102637
rdf:type
n4:Context
rdf:value
Several other reports illustrated that during the induction of T cell tolerance both transient and stable DC interactions can occur >>20<<,23. During the process of T cell activation the duration of T cell-DC contacts is highly variable in vivo, ranging from minutes to several hours 39. Early reports suggested that initial interactions during the first phase of T cell
n4:mentions
n2:15516925
Subject Item
_:vb6102638
rdf:type
n4:Context
rdf:value
Several other reports illustrated that during the induction of T cell tolerance both transient and stable DC interactions can occur 20,>>23<<. During the process of T cell activation the duration of T cell-DC contacts is highly variable in vivo, ranging from minutes to several hours 39. Early reports suggested that initial interactions during the first phase of T cell
n4:mentions
n2:15924144
Subject Item
_:vb6102639
rdf:type
n4:Context
rdf:value
During the process of T cell activation the duration of T cell-DC contacts is highly variable in vivo, ranging from minutes to several hours >>39<<. Early reports suggested that initial interactions during the first phase of T cell activation tend to be transient [5-10 minutes during the first 3-15 hours 20,21,40]. However, a more recent report found that long lived T cell-DC
n4:mentions
n2:16765578
Subject Item
_:vb6102640
rdf:type
n4:Context
rdf:value
Early reports suggested that initial interactions during the first phase of T cell activation tend to be transient [5-10 minutes during the first 3-15 hours >>20<<,21,40]. However, a more recent report found that long lived T cell-DC interactions could occur following the initial T cell-DC contact and that prolonged interactions were required for T cell activation 41. In this study, Erk
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Early reports suggested that initial interactions during the first phase of T cell activation tend to be transient [5-10 minutes during the first 3-15 hours 20,>>21<<,40]. However, a more recent report found that long lived T cell-DC interactions could occur following the initial T cell-DC contact and that prolonged interactions were required for T cell activation 41. In this study, Erk phosphorylation
n4:mentions
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Early reports suggested that initial interactions during the first phase of T cell activation tend to be transient [5-10 minutes during the first 3-15 hours 20,21,>>40<<]. However, a more recent report found that long lived T cell-DC interactions could occur following the initial T cell-DC contact and that prolonged interactions were required for T cell activation 41. In this study, Erk phosphorylation
n4:mentions
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However, a more recent report found that long lived T cell-DC interactions could occur following the initial T cell-DC contact and that prolonged interactions were required for T cell activation >>41<<. In this study, Erk phosphorylation occurred early after stable T cell-DC conjugates were formed, and IFN-γ production increased with longer T cell-DC interactions 41. These interactions required TCR engagement of peptide-MHCII complexes,
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In this study, Erk phosphorylation occurred early after stable T cell-DC conjugates were formed, and IFN-γ production increased with longer T cell-DC interactions >>41<<. These interactions required TCR engagement of peptide-MHCII complexes, as MHCII blockade in vivo terminated stable contacts, increased T cell motility, decreased T cell proliferation and prevented IFN-γ production. The second phase
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The second phase subsequently results in longer lived contacts where T cells form stable conjugates and begin to secrete cytokines which transitions into a third phase of high motility and rapid proliferation >>21<<. Our studies demonstrated that PD-1 blockade restored stable T cell-DC contacts, Erk phosphorylation, IFN-γ production and, most importantly, T1D. These findings suggest that PD-1 normally functions to prevent the T cell stop signal and
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Anergic T cells have been shown in vitro to form unstable immunological synapses with allogeneic APCs and failed to recruit the signaling proteins necessary to initiate T-cell activation >>42<<. We suggest that these transient interactions are required in our system (but impossible to discern in vivo), since the breakdown of tolerance following PD-1-PD-L1 blockade only occurs when antigen is present, as in the case of PLN and
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Recent reports suggested that B7-1 and PD-L1 interact directly with each other to negatively regulate T cells >>13<<. Thus far, however, in vivo data to support the functionality of this interaction have not been reported.
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Blockade of CTLA-4–B7 interactions can prevent induction of tolerance by peptide-pulsed fixed APCs but could not reverse tolerance once established >>10<<. It is important to note that the same dose of blocking CTLA-4–specific antibody was used in these MPLSM experiments as in a previous study that documented a critical role for CTLA-4 during the induction of tolerance 10.
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It is important to note that the same dose of blocking CTLA-4–specific antibody was used in these MPLSM experiments as in a previous study that documented a critical role for CTLA-4 during the induction of tolerance >>10<<. We detected no influence of CTLA-4 in the migratory behavior of tolerized T cells. Due to the complex nature of lattice formation with B7-1, we have also explored anti-CTLA-4 Fab fragments with similar results as shown using intact
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Upon PD-L engagement, PD-1 can bind SH2-domain containing tyrosine phosphatase 1 (SHP-1) and SHP-2 4,>>43<<. The binding of SHP-1 and SHP-2 can terminate early TCR signals by dephosphorylating key signaling intermediates including the kinases Akt, PI3K, ZAP-70, and PKC-θ. Like PD-1, CTLA-4 can interact with SHP-1 and SHP-2 14. Unlike PD-1,
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Like PD-1, CTLA-4 can interact with SHP-1 and SHP-2 >>14<<. Unlike PD-1, CTLA-4 can also interact with the phosphatase PP2a 15.
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Unlike PD-1, CTLA-4 can also interact with the phosphatase PP2a >>15<<. Another difference between these two inhibitory receptors is the structural motif used to bind phosphatases.
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CTLA-4 interacts through the immunoreceptor tyrosine-based inhibitory motifs (ITIM) while PD-1 contains an additional motif, the immunoreceptor tyrosine-based switch motif (ITSM) >>43<<. Mutation of the ITIM motif had little effect on signaling, while ITSM mutations abrogated the ability of PD-1 to limit T cell population expansion 43. This suggests that PD-1 and CTLA-4 use different structural motifs to bind and recruit
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Mutation of the ITIM motif had little effect on signaling, while ITSM mutations abrogated the ability of PD-1 to limit T cell population expansion >>43<<. This suggests that PD-1 and CTLA-4 use different structural motifs to bind and recruit phosphatases for signal blockade. Future work is necessary to determine the precise biochemical relationship between these two potent negative
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CTLA-4 deficiency results in rapid multi-organ tissue inflammation and death within 3-4 weeks of age, regardless of mouse genetic background >>5<<, whereas autoimmunity in PD-1-deficient mice is slower and tissue specific in a manner dependent on genetic background 4.
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Two recent reports investigated the role of CTLA-4 on T cell stop signals >>44<<,45. One study found that CTLA-4 positive T cells failed to stop or slow down in response to in vivo peptide challenge, and anti-CTLA-4 increased T cell motility 44.
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Two recent reports investigated the role of CTLA-4 on T cell stop signals 44,>>45<<. One study found that CTLA-4 positive T cells failed to stop or slow down in response to in vivo peptide challenge, and anti-CTLA-4 increased T cell motility 44.
n4:mentions
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One study found that CTLA-4 positive T cells failed to stop or slow down in response to in vivo peptide challenge, and anti-CTLA-4 increased T cell motility >>44<<. A follow-up study reported that CTLA-4-deficient T cells showed a marked resistance to a stop-signal induced by anti-CD3 45. It is difficult to explain the discrepancies between these previous studies, but it may be due to subtle
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A follow-up study reported that CTLA-4-deficient T cells showed a marked resistance to a stop-signal induced by anti-CD3 >>45<<. It is difficult to explain the discrepancies between these previous studies, but it may be due to subtle differences between sorted T cell subpopulations, CTLA-4 surface stability, blocking antibodies, or use of knockout T cells. Here we
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One major difference is the previous reports tracked the migration of naïve T cells during primary stimulation >>44<<,45 whereas the present studies focused on anergic T cells during the re-activation phase.
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One major difference is the previous reports tracked the migration of naïve T cells during primary stimulation 44,>>45<< whereas the present studies focused on anergic T cells during the re-activation phase.
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Finally, it is interesting to note that PD-1-PD-L1 blockade resulted in increased accumulation and/or enhanced proliferation of antigen-specific T cells within the target tissue >>10<<. This finding supports a key role for T cell-DC interactions during tissue-specific reactivation.
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