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dc:title
results
n5:contains
_:vb29673224 _:vb29673225 _:vb29673220 _:vb29673221 _:vb29673222 _:vb29673223 _:vb29673216 _:vb29673217 _:vb29673218 _:vb29673219 _:vb29673212 _:vb29673213 _:vb29673214 _:vb29673215 _:vb29673211
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the fact that the CNS is not directly connected to draining lymphatics, two major outflow pathways to the CLNs have been described for CNS interstitial fluid and cerebrospinal fluid via which antigens and APCs may reach the periphery>>17<<. Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs1314181920, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive
n2:mentions
n3:23695293
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Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs>>13<<14181920, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive autoimmune responses is not fully understood.
n2:mentions
n3:15294948
Subject Item
_:vb29673213
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Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs>>1314<<181920, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive autoimmune responses is not fully understood.
n2:mentions
n3:12965258
Subject Item
_:vb29673214
rdf:type
n2:Context
rdf:value
Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs>>131418<<1920, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive autoimmune responses is not fully understood.
n2:mentions
n3:7223890
Subject Item
_:vb29673215
rdf:type
n2:Context
rdf:value
Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs>>13141819<<20, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive autoimmune responses is not fully understood.
n2:mentions
n3:16204309
Subject Item
_:vb29673216
rdf:type
n2:Context
rdf:value
Further, although it has been shown that cerebrally injected soluble proteins and DCs reach the CLNs>>1314181920<<, how endogenous intracellular antigens that are originally sequestered behind the BBB might exit the CNS and induce adaptive autoimmune responses is not fully understood.
n2:mentions
n3:1928403
Subject Item
_:vb29673217
rdf:type
n2:Context
rdf:value
As CNP and PLP promoters have been shown to drive EGFP transgene expression in Schwann cells of peripheral nervous system (PNS) tissues as well>>21<<22, we performed western blotting to detect EGFP in the sciatic nerve tissue from CNP-OP and PLP-OP mice.
n2:mentions
n3:11826117
Subject Item
_:vb29673218
rdf:type
n2:Context
rdf:value
As CNP and PLP promoters have been shown to drive EGFP transgene expression in Schwann cells of peripheral nervous system (PNS) tissues as well>>2122<<, we performed western blotting to detect EGFP in the sciatic nerve tissue from CNP-OP and PLP-OP mice.
n2:mentions
n3:12404507
Subject Item
_:vb29673219
rdf:type
n2:Context
rdf:value
We have previously shown that intracerebral injection of both soluble and DC-bound antigens in naïve C57BL/6 mice induces peripheral activation of antigen-specific T cells and their homing to the CNS>>13<<1415. However, in these experiments the microtrauma caused by intracerebral injection might elaborate danger signals leading to the activation of APCs and potentially influence immunity to injected antigens.
n2:mentions
n3:15294948
Subject Item
_:vb29673220
rdf:type
n2:Context
rdf:value
We have previously shown that intracerebral injection of both soluble and DC-bound antigens in naïve C57BL/6 mice induces peripheral activation of antigen-specific T cells and their homing to the CNS>>1314<<15. However, in these experiments the microtrauma caused by intracerebral injection might elaborate danger signals leading to the activation of APCs and potentially influence immunity to injected antigens.
n2:mentions
n3:12965258
Subject Item
_:vb29673221
rdf:type
n2:Context
rdf:value
We have previously shown that intracerebral injection of both soluble and DC-bound antigens in naïve C57BL/6 mice induces peripheral activation of antigen-specific T cells and their homing to the CNS>>131415<<. However, in these experiments the microtrauma caused by intracerebral injection might elaborate danger signals leading to the activation of APCs and potentially influence immunity to injected antigens.
n2:mentions
n3:19129392
Subject Item
_:vb29673222
rdf:type
n2:Context
rdf:value
Tertiary lymphoid structures containing T cells and follicular DCs have been shown in the leptomeninges of mice with EAE and humans with multiple sclerosis>>23<<24 and may be critical in maintaining CNS autoimmune disease by providing a local niche for the presentation of CNS autoantigens2425.
n2:mentions
n3:22177922
Subject Item
_:vb29673223
rdf:type
n2:Context
rdf:value
Tertiary lymphoid structures containing T cells and follicular DCs have been shown in the leptomeninges of mice with EAE and humans with multiple sclerosis>>2324<< and may be critical in maintaining CNS autoimmune disease by providing a local niche for the presentation of CNS autoantigens2425.
n2:mentions
n3:15193029
Subject Item
_:vb29673224
rdf:type
n2:Context
rdf:value
and follicular DCs have been shown in the leptomeninges of mice with EAE and humans with multiple sclerosis2324 and may be critical in maintaining CNS autoimmune disease by providing a local niche for the presentation of CNS autoantigens>>24<<25. Thus, we studied whether the OVA-specific Thy1.1+ T cells associated with DCs in the CNS of CNP-OP mice with EAE (day 15 post-induction) by immunofluorescent microscopy.
n2:mentions
n3:15193029
Subject Item
_:vb29673225
rdf:type
n2:Context
rdf:value
follicular DCs have been shown in the leptomeninges of mice with EAE and humans with multiple sclerosis2324 and may be critical in maintaining CNS autoimmune disease by providing a local niche for the presentation of CNS autoantigens>>2425<<. Thus, we studied whether the OVA-specific Thy1.1+ T cells associated with DCs in the CNS of CNP-OP mice with EAE (day 15 post-induction) by immunofluorescent microscopy.
n2:mentions
n3:16498451
Subject Item
_:vb29673226
rdf:type
n5:Section
dc:title
methods
n5:contains
_:vb29673228 _:vb29673229 _:vb29673230 _:vb29673231 _:vb29673227
Subject Item
_:vb29673227
rdf:type
n2:Context
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One (#147) was crossed with either PLP/CreERT mice>>46<< or with Cnp1Cre/Cre mice47 to obtain the double transgenic PLP-OP or CNP-OP offspring having either tamoxifen-inducible or constitutive Cre-mediated myelinating glial cell-specific transgene expression, respectively.
n2:mentions
n3:12481300
Subject Item
_:vb29673228
rdf:type
n2:Context
rdf:value
One (#147) was crossed with either PLP/CreERT mice46 or with Cnp1Cre/Cre mice>>47<< to obtain the double transgenic PLP-OP or CNP-OP offspring having either tamoxifen-inducible or constitutive Cre-mediated myelinating glial cell-specific transgene expression, respectively.
n2:mentions
n3:12590258
Subject Item
_:vb29673229
rdf:type
n2:Context
rdf:value
As described by Leone et al.>>48<<, tamoxifen (TM) (Sigma, St. Louis, MO) was dissolved in a 9:1 sunflower oil:ethanol mixture at a concentration of 10 mg/mL.
n2:mentions
n3:12727441
Subject Item
_:vb29673230
rdf:type
n2:Context
rdf:value
Oligodendrocytes (ODCs) were prepared from CNS tissues according to the following protocol adapted from Ji et al.>>49<<. Briefly, CNS tissues were minced finely in 5 mL digestion solution containing 1 mg/mL papain (Worthington Biochemical), 20 ng/mL DNase (Sigma), 1.1 mM EDTA, 0.067 mM mercaptoethanol, and 5.5 mM cysteine-HCl in HBSS.
n2:mentions
n3:23291597
Subject Item
_:vb29673231
rdf:type
n2:Context
rdf:value
Lymphocytes from pooled lymph node preparation from OT-I Thy1.1 and OT-II Thy1.1 transgenic mice were CFSE labeled in 1 μM CFSE as previously described>>50<<. 5 × 105 CFSE-labeled naïve (i.e., CD44low) OT-I CD8+ Thy1.1+ and OT-II CD4+ Thy1.1+ T cells were adoptively transferred intravenously (i.
n2:mentions
n3:20625513
Subject Item
_:vb29673232
rdf:type
n5:Section
dc:title
discussion
n5:contains
_:vb29673252 _:vb29673253 _:vb29673254 _:vb29673255 _:vb29673248 _:vb29673249 _:vb29673250 _:vb29673251 _:vb29673260 _:vb29673261 _:vb29673256 _:vb29673257 _:vb29673258 _:vb29673259 _:vb29673236 _:vb29673237 _:vb29673238 _:vb29673239 _:vb29673233 _:vb29673234 _:vb29673235 _:vb29673244 _:vb29673245 _:vb29673246 _:vb29673247 _:vb29673240 _:vb29673241 _:vb29673242 _:vb29673243
Subject Item
_:vb29673233
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modeled a scenario in which T cells with high affinity for the neoantigens bypassed central tolerance mechanisms but were kept in check by peripheral tolerance mechanisms, which are thought to be dysregulated in patients with MS>>26<<.
n2:mentions
n3:21437244
Subject Item
_:vb29673234
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>22<<272829303132.
n2:mentions
n3:12404507
Subject Item
_:vb29673235
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>2227<<2829303132.
n2:mentions
n3:1724062
Subject Item
_:vb29673236
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>222728<<29303132. Our findings that T cells were not stimulated by CNP-OP sciatic nerve tissue lysates in vitro and were not recruited to the sciatic nerves of CNP-OP mice with EAE in vivo suggest that neoantigen expression in peripheral myelin is
n2:mentions
n3:21488900
Subject Item
_:vb29673237
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>22272829<<303132. Our findings that T cells were not stimulated by CNP-OP sciatic nerve tissue lysates in vitro and were not recruited to the sciatic nerves of CNP-OP mice with EAE in vivo suggest that neoantigen expression in peripheral myelin is
n2:mentions
n3:9710259
Subject Item
_:vb29673238
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>2227282930<<3132. Our findings that T cells were not stimulated by CNP-OP sciatic nerve tissue lysates in vitro and were not recruited to the sciatic nerves of CNP-OP mice with EAE in vivo suggest that neoantigen expression in peripheral myelin is
n2:mentions
n3:2441390
Subject Item
_:vb29673239
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>222728293031<<32. Our findings that T cells were not stimulated by CNP-OP sciatic nerve tissue lysates in vitro and were not recruited to the sciatic nerves of CNP-OP mice with EAE in vivo suggest that neoantigen expression in peripheral myelin is below
n2:mentions
n3:3040924
Subject Item
_:vb29673240
rdf:type
n2:Context
rdf:value
Although PLP and CNPase proteins are most abundant in central myelin, they are also expressed at significantly lower levels in peripheral myelin>>22272829303132<<. Our findings that T cells were not stimulated by CNP-OP sciatic nerve tissue lysates in vitro and were not recruited to the sciatic nerves of CNP-OP mice with EAE in vivo suggest that neoantigen expression in peripheral myelin is below
n2:mentions
n3:7516688
Subject Item
_:vb29673241
rdf:type
n2:Context
rdf:value
This might be due to either little or no CNP- and PLP-driven transgene expression in the PNS after P>>3033<<. Additionally, CNPase is weakly expressed in non-neural cells and tissues, including circulating lymphocytes and the thymus31. However, we did not observe EGFP expression in secondary lymphoid organs of the CNP-OP and PLP-OP mice. While
n2:mentions
n3:11756744
Subject Item
_:vb29673242
rdf:type
n2:Context
rdf:value
Additionally, CNPase is weakly expressed in non-neural cells and tissues, including circulating lymphocytes and the thymus>>31<<. However, we did not observe EGFP expression in secondary lymphoid organs of the CNP-OP and PLP-OP mice. While the possibility remains that the level of neoantigen expression could be below the detectable level in these tissues by FACS
n2:mentions
n3:3040924
Subject Item
_:vb29673243
rdf:type
n2:Context
rdf:value
expression could be below the detectable level in these tissues by FACS analysis, peripheral expression of OVA antigen by CD45neg lymph node stromal cells has been shown to result in deletional tolerance of transferred OT-I T cells>>34<<. Contrary to this, we observed activation of transferred OT-I T cells, which produced IFN-γ under normal conditions.
n2:mentions
n3:17195844
Subject Item
_:vb29673244
rdf:type
n2:Context
rdf:value
of adoptively transferred LCMV antigen-specific CD8+ T cells in all peripheral lymphoid tissues, including both proximal CNS-draining lymph nodes and distal non-draining lymph nodes and spleen, using another inducible PLP driver strain>>35<<. However, our result differs from that of Na et al. who did not observe peripheral OVA-specific CD8+ T cell proliferation after P10 in mice with MBP promoter-driven OVA in ODCs36.
n2:mentions
n3:19462379
Subject Item
_:vb29673245
rdf:type
n2:Context
rdf:value
However, our result differs from that of Na et al. who did not observe peripheral OVA-specific CD8+ T cell proliferation after P10 in mice with MBP promoter-driven OVA in ODCs>>36<<. Their results may indicate OT-I T cell deletion in response to increased levels of OVA, as MBP-specific T cell deletion increases after two weeks of age and correlates with MBP protein expression28.
n2:mentions
n3:18669487
Subject Item
_:vb29673246
rdf:type
n2:Context
rdf:value
Their results may indicate OT-I T cell deletion in response to increased levels of OVA, as MBP-specific T cell deletion increases after two weeks of age and correlates with MBP protein expression>>28<<.
n2:mentions
n3:21488900
Subject Item
_:vb29673247
rdf:type
n2:Context
rdf:value
Our laboratory has previously shown that intracerebrally injected soluble and DC-bound OVA proteins elicit the systemic activation of neoantigen-specific CD8+ T cells and their recruitment to the brain in naïve animals>>13<<14. However, in the PLP-OP and CNP-OP models the peripherally activated OT-I T cells did not accumulate in the CNS.
n2:mentions
n3:15294948
Subject Item
_:vb29673248
rdf:type
n2:Context
rdf:value
Our laboratory has previously shown that intracerebrally injected soluble and DC-bound OVA proteins elicit the systemic activation of neoantigen-specific CD8+ T cells and their recruitment to the brain in naïve animals>>1314<<. However, in the PLP-OP and CNP-OP models the peripherally activated OT-I T cells did not accumulate in the CNS.
n2:mentions
n3:12965258
Subject Item
_:vb29673249
rdf:type
n2:Context
rdf:value
the physiological levels of the neoantigens are lower in the transgenic mice of the present study, resulting in fewer MHC-peptide complexes and, thus, lower OT-I T cell avidity and incomplete activation in the peripheral lymphoid tissues>>37<<. In support of this, a previous study showed weaker OT-I T cell proliferation that remained restricted to gut-associated lymphoid tissues (GALT) in mice fed lower amounts of OVA but showed stronger proliferation that disseminated to LNs
n2:mentions
n3:19444307
Subject Item
_:vb29673250
rdf:type
n2:Context
rdf:value
weaker OT-I T cell proliferation that remained restricted to gut-associated lymphoid tissues (GALT) in mice fed lower amounts of OVA but showed stronger proliferation that disseminated to LNs beyond GALT in mice fed higher amounts of OVA>>34<<. Consistent with this is our finding of weaker peripheral OT-I T cell activation in naive PLP-OP mice with lower neoantigen expression compared to that observed in CNP-OP mice with higher neoantigen expression.
n2:mentions
n3:17195844
Subject Item
_:vb29673251
rdf:type
n2:Context
rdf:value
In agreement with this, Na et al. found that OT-I T cells only induced disease in Rag−/− MBP-OVA mice but not in immune-sufficient hosts>>36<<. However, another study showed that ODC antigen-specific CD8+ T cells induced CNS disease in immune-sufficient host animals when pre-activated prior to transfer38. Collectively, these results argue that the context of antigen presentation
n2:mentions
n3:18669487
Subject Item
_:vb29673252
rdf:type
n2:Context
rdf:value
However, another study showed that ODC antigen-specific CD8+ T cells induced CNS disease in immune-sufficient host animals when pre-activated prior to transfer>>38<<. Collectively, these results argue that the context of antigen presentation and T cell activation regulate CNS immune privilege and disease initiation.
n2:mentions
n3:18641296
Subject Item
_:vb29673253
rdf:type
n2:Context
rdf:value
Walter and Albert showed that antigen from intracerebrally injected MHC class I-deficient mOVA/Kb−/− splenocytes is first cross-presented to OT-I T cells in the CLNs prior to their accumulation in the brain>>39<<. It has also been shown that cell-associated OVA is more efficiently cross-presented to OT-I T cells than to OT-II T cells, but that soluble OVA stimulates both OT-I and OT-II T cells equally40. Taken together with our data showing a
n2:mentions
n3:17475827
Subject Item
_:vb29673254
rdf:type
n2:Context
rdf:value
It has also been shown that cell-associated OVA is more efficiently cross-presented to OT-I T cells than to OT-II T cells, but that soluble OVA stimulates both OT-I and OT-II T cells equally>>40<<. Taken together with our data showing a strong OT-I response, these findings support the idea that the ODC-associated OVA is being efficiently cross-presented to the OT-I T cells in the peripheral lymphoid tissues of CNP-OP and PLP-OP
n2:mentions
n3:11342628
Subject Item
_:vb29673255
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be due to increased antigen drainage from the CNS as was suggested previously in a study showing that myelin proteins and degradation products from brain lesions of monkeys with EAE and humans with MS can be found in the draining CLNs>>41<<. The fact that CNS-infiltrating OT-II T cells were only observed in mice with both high neoantigen expression and active neuroinflammation suggests that the levels of both antigen expression and antigen release/drainage contribute to CNS
n2:mentions
n3:12421916
Subject Item
_:vb29673256
rdf:type
n2:Context
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not observably contribute to EAE onset and is in agreement with that reported by Schildknecht et al., who observed that non-tolerized myelinating glial cell antigen-specific CD8+ T cells did not induce CNS autoimmunity or inflammation>>35<<. As the initiation of EAE is determined by accumulation of DCs42, further studies are needed to determine the role of the neoantigen-specific T cells in the chronic phase of EAE disease in the CNP-OP and PLP-OP mice.
n2:mentions
n3:19462379
Subject Item
_:vb29673257
rdf:type
n2:Context
rdf:value
As the initiation of EAE is determined by accumulation of DCs>>42<<, further studies are needed to determine the role of the neoantigen-specific T cells in the chronic phase of EAE disease in the CNP-OP and PLP-OP mice.
n2:mentions
n3:15735653
Subject Item
_:vb29673258
rdf:type
n2:Context
rdf:value
of CD4+ T cells, as CCL20, the ligand for the receptor CCR6 expressed on Th17 cells, is constitutively expressed on epithelial cells of the choroid plexus; this interaction facilitates Th17 cell entry into the brain at this location>>43<< and is a required step for EAE induction. It has been shown that adhesion of CD8+ T cells to inflamed brain microvessels is critically dependent on P-selectin glycoprotein ligand-1 (PSGL-1)44.
n2:mentions
n3:19305396
Subject Item
_:vb29673259
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It has been shown that adhesion of CD8+ T cells to inflamed brain microvessels is critically dependent on P-selectin glycoprotein ligand-1 (PSGL-1)>>44<<. Interestingly, the receptor for PSGL-1, P-selectin, has been shown to be upregulated more in the spinal cord than in the brain during EAE45. Whether P-selectin is also upregulated more in the spinal cord of the PLP-OP and CNP-OP models
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Interestingly, the receptor for PSGL-1, P-selectin, has been shown to be upregulated more in the spinal cord than in the brain during EAE>>45<<. Whether P-selectin is also upregulated more in the spinal cord of the PLP-OP and CNP-OP models under neuroinflammatory conditions remains to be determined.
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At one time it was thought that the CNS was completely isolated from immune surveillance because of lack of draining lymphatics, which prevented escape of CNS antigens, and prevention of immune cell entry due to the BBB>>3<<. Our data, instead, suggest that there is continuous surveillance of CNS antigens in peripheral lymphoid tissues, which indicates that immune privilege is not the result of limited antigen sampling. Additionally, neuroinflammation induces
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