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Botelho)] TJ ET 0.271 0.267 0.267 rg BT 90.200 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 95.621 638.299 Td /F1 9.8 Tf [(Elizabeth Wang)] TJ ET 0.271 0.267 0.267 rg BT 163.901 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 169.322 638.299 Td /F1 9.8 Tf [(Jane Y. Chen)] TJ ET 0.271 0.267 0.267 rg BT 228.397 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 233.818 638.299 Td /F1 9.8 Tf [(Sandra Holley)] TJ ET 0.271 0.267 0.267 rg BT 295.048 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 300.469 638.299 Td /F1 9.8 Tf [(Veronique Andre)] TJ ET 0.271 0.267 0.267 rg BT 373.633 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 379.054 638.299 Td /F1 9.8 Tf [(Carlos Cepeda)] TJ ET 0.271 0.267 0.267 rg BT 444.077 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 449.498 638.299 Td /F1 9.8 Tf [(Michael S. Levine)] TJ ET 0.271 0.267 0.267 rg BT 26.250 626.394 Td /F1 9.8 Tf [(Botelho EP, Wang E, Chen JY, Holley S, Andre V, Cepeda C, Levine MS. Differential Synaptic and Extrasynaptic Glutamate-)] TJ ET BT 26.250 614.490 Td /F1 9.8 Tf [(Receptor Alterations in Striatal Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice. PLOS Currents )] TJ ET BT 26.250 602.585 Td /F1 9.8 Tf [(Huntington Disease. 2014 May 21 . Edition 1. doi: 10.1371/currents.hd.34957c4f8bd7cb1f5ec47381dfc811c3.)] TJ ET q 15.000 31.834 577.500 568.370 re W n 0.271 0.267 0.267 rg BT 26.250 573.482 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 553.528 Td /F1 9.8 Tf [(Huntingtons disease \(HD\) is a late-onset, slowly progressing neurodegenerative disorder caused by an expansion of glutamine )] TJ ET BT 26.250 541.623 Td /F1 9.8 Tf [(repeats. The YAC128 mouse model has been widely used to study the progression of HD symptoms, but little is known about )] TJ ET BT 26.250 529.719 Td /F1 9.8 Tf [(synaptic alterations in very old animals. The present experiments examined synaptic properties of striatal medium-sized spiny )] TJ ET BT 26.250 517.814 Td /F1 9.8 Tf [(neurons \(MSNs\) in 16 month-old YAC128 mice. These mice were crossed with mice expressing enhanced green fluorescent )] TJ ET BT 26.250 505.909 Td /F1 9.8 Tf [(protein \(EGFP\) under the control of either D1 or D2 dopamine receptor promoters to identify MSNs originating the direct and )] TJ ET BT 26.250 494.004 Td /F1 9.8 Tf [(indirect pathways, respectively. The input-output curves of evoked excitatory postsynaptic currents mediated by activation of the )] TJ ET BT 26.250 482.100 Td /F1 9.8 Tf [(?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid \(AMPA\) or N-methyl-D-aspartate \(NMDA\) receptors were reduced in )] TJ ET BT 26.250 470.195 Td /F1 9.8 Tf [(MSNs in both pathways. In the presence of DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA\), a glutamate transporter blocker )] TJ ET BT 26.250 458.290 Td /F1 9.8 Tf [(used to increase activation of extrasynaptic receptors, NMDA receptor-mediated currents displayed altered amplitudes, longer )] TJ ET BT 26.250 446.385 Td /F1 9.8 Tf [(decay times, and greater charge \(response areas\) in both direct and indirect pathway MSNs in YAC128 mice compared to )] TJ ET BT 26.250 434.481 Td /F1 9.8 Tf [(wildtype controls. Amplitudes were significantly increased, primarily in direct pathway MSNs while normalized areas were )] TJ ET BT 26.250 422.576 Td /F1 9.8 Tf [(significantly increased only in indirect pathway MSNs, suggesting that the two types of MSNs are affected in different ways. It )] TJ ET BT 26.250 410.671 Td /F1 9.8 Tf [(may be that indirect pathway neurons are more susceptible to changes in glutamate transport. Taken together, the present )] TJ ET BT 26.250 398.766 Td /F1 9.8 Tf [(findings demonstrate differential alterations in synaptic versus extrasynaptic NMDA receptors in both direct and indirect pathway )] TJ ET BT 26.250 386.862 Td /F1 9.8 Tf [(MSNs in late HD, which may contribute to the dysfunction and degeneration in both pathways.)] TJ ET BT 26.250 350.259 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 330.305 Td /F1 9.8 Tf [(Supported by USPHS NS41574. The funders had no role in study design, data collection and analysis, decision to publish, or )] TJ ET BT 26.250 318.400 Td /F1 9.8 Tf [(preparation of the manuscript. The authors have declared that no competing interests exist.)] TJ ET BT 26.250 289.298 Td /F4 12.0 Tf [(Notice of Correction)] TJ ET BT 26.250 269.343 Td /F4 9.8 Tf [(11 July 2017:)] TJ ET BT 86.953 269.343 Td /F1 9.8 Tf [( PLOS Currents Correction: Differential Synaptic and Extrasynaptic Glutamate-Receptor Alterations in Striatal )] TJ ET BT 26.250 257.439 Td /F1 9.8 Tf [(Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice.. PLOS Currents Huntington Disease. 2017 Jul 11 . )] TJ ET BT 26.250 245.534 Td /F1 9.8 Tf [(Edition 1. doi: 10.1371/currents.hd.2f36af340a70a0d248b48d4cc9c3e58e. )] TJ ET 0.267 0.267 0.267 rg BT 349.297 245.534 Td /F1 9.8 Tf [(View Correction.)] TJ ET 0.271 0.267 0.267 rg BT 26.250 208.931 Td /F4 12.0 Tf [(Introduction)] TJ ET BT 26.250 188.977 Td /F1 9.8 Tf [(Huntingtons disease \(HD\), a slowly progressing neurodegenerative disorder caused by an expansion of glutamine \(CAG\) )] TJ ET BT 26.250 177.072 Td /F1 9.8 Tf [(repeats, is characterized by motor, psychiatric and cognitive symptoms)] TJ ET 0.267 0.267 0.267 rg BT 331.854 178.580 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 336.673 177.072 Td /F1 9.8 Tf [(. The YAC128 mouse model has been widely used to )] TJ ET BT 26.250 165.168 Td /F1 9.8 Tf [(examine mechanisms of HD. YAC128 mice carry the entire human HD gene containing 128 CAG repeats on a yeast artificial )] TJ ET BT 26.250 153.263 Td /F1 9.8 Tf [(chromosome \(line 53\) and express full-length human mutant huntingtin protein)] TJ ET 0.267 0.267 0.267 rg BT 363.853 154.770 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 368.672 153.263 Td /F1 9.8 Tf [(. These mice display a hyperkinetic phenotype )] TJ ET BT 26.250 141.358 Td /F1 9.8 Tf [(at 3 months of age, followed by motor deficits at 6 months, with eventual progression to hypokinesia by 12 months)] TJ ET 0.267 0.267 0.267 rg BT 517.777 142.865 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 522.595 141.358 Td /F1 9.8 Tf [(. The )] TJ ET BT 26.250 129.453 Td /F1 9.8 Tf [(behavioral changes are associated with biphasic alterations in excitatory synaptic transmission to striatal medium-sized spiny )] TJ ET BT 26.250 117.549 Td /F1 9.8 Tf [(neurons \(MSNs\) of the direct \(expressing dopamine D1 receptors\) and indirect \(expressing dopamine D2 receptors\) striatal )] TJ ET BT 26.250 105.644 Td /F1 9.8 Tf [(output pathways)] TJ ET 0.267 0.267 0.267 rg BT 97.249 107.151 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 102.068 109.532 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 104.478 107.151 Td /F4 8.7 Tf [(5)] TJ ET 0.271 0.267 0.267 rg BT 109.296 105.644 Td /F1 9.8 Tf [(. At the early, pre-symptomatic stage, glutamate transmission is increased in both direct and indirect )] TJ ET BT 26.250 93.739 Td /F1 9.8 Tf [(pathways. In contrast, at 12 months of age, the more symptomatic phase, glutamatergic transmission is markedly reduced in )] TJ ET BT 26.250 81.834 Td /F1 9.8 Tf [(direct pathway MSNs, while only a trend towards a reduction in glutamatergic inputs is observed in D2 MSNs)] TJ ET 0.267 0.267 0.267 rg BT 494.962 83.342 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 499.780 81.834 Td /F1 9.8 Tf [(. Such biphasic )] TJ ET BT 26.250 69.930 Td /F1 9.8 Tf [(alterations are also observed in evoked ?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor \(AMPAR\)-mediated )] TJ ET BT 26.250 58.025 Td /F1 9.8 Tf [(currents, demonstrated by an increase in the peak amplitude in indirect pathway MSNs at 1.5 months and a reduction in direct )] TJ ET BT 26.250 46.120 Td /F1 9.8 Tf [(pathway MSNs at 12 months)] TJ ET 0.267 0.267 0.267 rg BT 150.884 47.627 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 155.703 46.120 Td /F1 9.8 Tf [(. Alterations in synaptic communication lead to a progressive disconnection between cortex and )] TJ ET Q q 15.000 659.406 577.500 78.594 re W n 0.267 0.267 0.267 rg BT 15.000 718.042 Td /F2 21.0 Tf [(Differential Synaptic and Extrasynaptic Glutamate-Receptor )] TJ ET BT 15.000 693.094 Td /F2 21.0 Tf [(Alterations in Striatal Medium-Sized Spiny Neurons of Aged )] TJ ET BT 15.000 668.146 Td /F2 21.0 Tf [(YAC128 Huntingtons Disease Mice)] TJ ET Q 0.271 0.267 0.267 rg BT 15.000 650.140 Td /F3 9.8 Tf [(May 21, 2014)] TJ ET BT 73.763 650.140 Td /F3 9.8 Tf [()] TJ ET 0.267 0.267 0.267 rg BT 78.638 650.140 Td /F3 9.8 Tf [(HD Models)] TJ ET BT 26.250 638.299 Td /F1 9.8 Tf [(Eli P. Botelho)] TJ ET 0.271 0.267 0.267 rg BT 90.200 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 95.621 638.299 Td /F1 9.8 Tf [(Elizabeth Wang)] TJ ET 0.271 0.267 0.267 rg BT 163.901 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 169.322 638.299 Td /F1 9.8 Tf [(Jane Y. Chen)] TJ ET 0.271 0.267 0.267 rg BT 228.397 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 233.818 638.299 Td /F1 9.8 Tf [(Sandra Holley)] TJ ET 0.271 0.267 0.267 rg BT 295.048 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 300.469 638.299 Td /F1 9.8 Tf [(Veronique Andre)] TJ ET 0.271 0.267 0.267 rg BT 373.633 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 379.054 638.299 Td /F1 9.8 Tf [(Carlos Cepeda)] TJ ET 0.271 0.267 0.267 rg BT 444.077 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 449.498 638.299 Td /F1 9.8 Tf [(Michael S. Levine)] TJ ET 0.271 0.267 0.267 rg BT 26.250 626.394 Td /F1 9.8 Tf [(Botelho EP, Wang E, Chen JY, Holley S, Andre V, Cepeda C, Levine MS. Differential Synaptic and Extrasynaptic Glutamate-)] TJ ET BT 26.250 614.490 Td /F1 9.8 Tf [(Receptor Alterations in Striatal Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice. PLOS Currents )] TJ ET BT 26.250 602.585 Td /F1 9.8 Tf [(Huntington Disease. 2014 May 21 . Edition 1. doi: 10.1371/currents.hd.34957c4f8bd7cb1f5ec47381dfc811c3.)] TJ ET q 15.000 31.834 577.500 568.370 re W n 0.271 0.267 0.267 rg BT 26.250 573.482 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 553.528 Td /F1 9.8 Tf [(Huntingtons disease \(HD\) is a late-onset, slowly progressing neurodegenerative disorder caused by an expansion of glutamine )] TJ ET BT 26.250 541.623 Td /F1 9.8 Tf [(repeats. The YAC128 mouse model has been widely used to study the progression of HD symptoms, but little is known about )] TJ ET BT 26.250 529.719 Td /F1 9.8 Tf [(synaptic alterations in very old animals. The present experiments examined synaptic properties of striatal medium-sized spiny )] TJ ET BT 26.250 517.814 Td /F1 9.8 Tf [(neurons \(MSNs\) in 16 month-old YAC128 mice. These mice were crossed with mice expressing enhanced green fluorescent )] TJ ET BT 26.250 505.909 Td /F1 9.8 Tf [(protein \(EGFP\) under the control of either D1 or D2 dopamine receptor promoters to identify MSNs originating the direct and )] TJ ET BT 26.250 494.004 Td /F1 9.8 Tf [(indirect pathways, respectively. The input-output curves of evoked excitatory postsynaptic currents mediated by activation of the )] TJ ET BT 26.250 482.100 Td /F1 9.8 Tf [(?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid \(AMPA\) or N-methyl-D-aspartate \(NMDA\) receptors were reduced in )] TJ ET BT 26.250 470.195 Td /F1 9.8 Tf [(MSNs in both pathways. In the presence of DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA\), a glutamate transporter blocker )] TJ ET BT 26.250 458.290 Td /F1 9.8 Tf [(used to increase activation of extrasynaptic receptors, NMDA receptor-mediated currents displayed altered amplitudes, longer )] TJ ET BT 26.250 446.385 Td /F1 9.8 Tf [(decay times, and greater charge \(response areas\) in both direct and indirect pathway MSNs in YAC128 mice compared to )] TJ ET BT 26.250 434.481 Td /F1 9.8 Tf [(wildtype controls. Amplitudes were significantly increased, primarily in direct pathway MSNs while normalized areas were )] TJ ET BT 26.250 422.576 Td /F1 9.8 Tf [(significantly increased only in indirect pathway MSNs, suggesting that the two types of MSNs are affected in different ways. It )] TJ ET BT 26.250 410.671 Td /F1 9.8 Tf [(may be that indirect pathway neurons are more susceptible to changes in glutamate transport. Taken together, the present )] TJ ET BT 26.250 398.766 Td /F1 9.8 Tf [(findings demonstrate differential alterations in synaptic versus extrasynaptic NMDA receptors in both direct and indirect pathway )] TJ ET BT 26.250 386.862 Td /F1 9.8 Tf [(MSNs in late HD, which may contribute to the dysfunction and degeneration in both pathways.)] TJ ET BT 26.250 350.259 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 330.305 Td /F1 9.8 Tf [(Supported by USPHS NS41574. The funders had no role in study design, data collection and analysis, decision to publish, or )] TJ ET BT 26.250 318.400 Td /F1 9.8 Tf [(preparation of the manuscript. The authors have declared that no competing interests exist.)] TJ ET BT 26.250 289.298 Td /F4 12.0 Tf [(Notice of Correction)] TJ ET BT 26.250 269.343 Td /F4 9.8 Tf [(11 July 2017:)] TJ ET BT 86.953 269.343 Td /F1 9.8 Tf [( PLOS Currents Correction: Differential Synaptic and Extrasynaptic Glutamate-Receptor Alterations in Striatal )] TJ ET BT 26.250 257.439 Td /F1 9.8 Tf [(Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice.. PLOS Currents Huntington Disease. 2017 Jul 11 . )] TJ ET BT 26.250 245.534 Td /F1 9.8 Tf [(Edition 1. doi: 10.1371/currents.hd.2f36af340a70a0d248b48d4cc9c3e58e. )] TJ ET 0.267 0.267 0.267 rg BT 349.297 245.534 Td /F1 9.8 Tf [(View Correction.)] TJ ET 0.271 0.267 0.267 rg BT 26.250 208.931 Td /F4 12.0 Tf [(Introduction)] TJ ET BT 26.250 188.977 Td /F1 9.8 Tf [(Huntingtons disease \(HD\), a slowly progressing neurodegenerative disorder caused by an expansion of glutamine \(CAG\) )] TJ ET BT 26.250 177.072 Td /F1 9.8 Tf [(repeats, is characterized by motor, psychiatric and cognitive symptoms)] TJ ET 0.267 0.267 0.267 rg BT 331.854 178.580 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 336.673 177.072 Td /F1 9.8 Tf [(. The YAC128 mouse model has been widely used to )] TJ ET BT 26.250 165.168 Td /F1 9.8 Tf [(examine mechanisms of HD. YAC128 mice carry the entire human HD gene containing 128 CAG repeats on a yeast artificial )] TJ ET BT 26.250 153.263 Td /F1 9.8 Tf [(chromosome \(line 53\) and express full-length human mutant huntingtin protein)] TJ ET 0.267 0.267 0.267 rg BT 363.853 154.770 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 368.672 153.263 Td /F1 9.8 Tf [(. These mice display a hyperkinetic phenotype )] TJ ET BT 26.250 141.358 Td /F1 9.8 Tf [(at 3 months of age, followed by motor deficits at 6 months, with eventual progression to hypokinesia by 12 months)] TJ ET 0.267 0.267 0.267 rg BT 517.777 142.865 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 522.595 141.358 Td /F1 9.8 Tf [(. The )] TJ ET BT 26.250 129.453 Td /F1 9.8 Tf [(behavioral changes are associated with biphasic alterations in excitatory synaptic transmission to striatal medium-sized spiny )] TJ ET BT 26.250 117.549 Td /F1 9.8 Tf [(neurons \(MSNs\) of the direct \(expressing dopamine D1 receptors\) and indirect \(expressing dopamine D2 receptors\) striatal )] TJ ET BT 26.250 105.644 Td /F1 9.8 Tf [(output pathways)] TJ ET 0.267 0.267 0.267 rg BT 97.249 107.151 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 102.068 109.532 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 104.478 107.151 Td /F4 8.7 Tf [(5)] TJ ET 0.271 0.267 0.267 rg BT 109.296 105.644 Td /F1 9.8 Tf [(. At the early, pre-symptomatic stage, glutamate transmission is increased in both direct and indirect )] TJ ET BT 26.250 93.739 Td /F1 9.8 Tf [(pathways. In contrast, at 12 months of age, the more symptomatic phase, glutamatergic transmission is markedly reduced in )] TJ ET BT 26.250 81.834 Td /F1 9.8 Tf [(direct pathway MSNs, while only a trend towards a reduction in glutamatergic inputs is observed in D2 MSNs)] TJ ET 0.267 0.267 0.267 rg BT 494.962 83.342 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 499.780 81.834 Td /F1 9.8 Tf [(. Such biphasic )] TJ ET BT 26.250 69.930 Td /F1 9.8 Tf [(alterations are also observed in evoked ?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor \(AMPAR\)-mediated )] TJ ET BT 26.250 58.025 Td /F1 9.8 Tf [(currents, demonstrated by an increase in the peak amplitude in indirect pathway MSNs at 1.5 months and a reduction in direct )] TJ ET BT 26.250 46.120 Td /F1 9.8 Tf [(pathway MSNs at 12 months)] TJ ET 0.267 0.267 0.267 rg BT 150.884 47.627 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 155.703 46.120 Td /F1 9.8 Tf [(. Alterations in synaptic communication lead to a progressive disconnection between cortex and )] TJ ET Q q 15.000 659.406 577.500 78.594 re W n 0.267 0.267 0.267 rg BT 15.000 718.042 Td /F2 21.0 Tf [(Differential Synaptic and Extrasynaptic Glutamate-Receptor )] TJ ET BT 15.000 693.094 Td /F2 21.0 Tf [(Alterations in Striatal Medium-Sized Spiny Neurons of Aged )] TJ ET BT 15.000 668.146 Td /F2 21.0 Tf [(YAC128 Huntingtons Disease Mice)] TJ ET Q 0.271 0.267 0.267 rg BT 15.000 650.140 Td /F3 9.8 Tf [(May 21, 2014)] TJ ET BT 73.763 650.140 Td /F3 9.8 Tf [()] TJ ET 0.267 0.267 0.267 rg BT 78.638 650.140 Td /F3 9.8 Tf [(HD Models)] TJ ET BT 26.250 638.299 Td /F1 9.8 Tf [(Eli P. Botelho)] TJ ET 0.271 0.267 0.267 rg BT 90.200 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 95.621 638.299 Td /F1 9.8 Tf [(Elizabeth Wang)] TJ ET 0.271 0.267 0.267 rg BT 163.901 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 169.322 638.299 Td /F1 9.8 Tf [(Jane Y. Chen)] TJ ET 0.271 0.267 0.267 rg BT 228.397 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 233.818 638.299 Td /F1 9.8 Tf [(Sandra Holley)] TJ ET 0.271 0.267 0.267 rg BT 295.048 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 300.469 638.299 Td /F1 9.8 Tf [(Veronique Andre)] TJ ET 0.271 0.267 0.267 rg BT 373.633 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 379.054 638.299 Td /F1 9.8 Tf [(Carlos Cepeda)] TJ ET 0.271 0.267 0.267 rg BT 444.077 638.299 Td /F1 9.8 Tf [(, )] TJ ET 0.267 0.267 0.267 rg BT 449.498 638.299 Td /F1 9.8 Tf [(Michael S. Levine)] TJ ET 0.271 0.267 0.267 rg BT 26.250 626.394 Td /F1 9.8 Tf [(Botelho EP, Wang E, Chen JY, Holley S, Andre V, Cepeda C, Levine MS. Differential Synaptic and Extrasynaptic Glutamate-)] TJ ET BT 26.250 614.490 Td /F1 9.8 Tf [(Receptor Alterations in Striatal Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice. PLOS Currents )] TJ ET BT 26.250 602.585 Td /F1 9.8 Tf [(Huntington Disease. 2014 May 21 . Edition 1. doi: 10.1371/currents.hd.34957c4f8bd7cb1f5ec47381dfc811c3.)] TJ ET q 15.000 31.834 577.500 568.370 re W n 0.271 0.267 0.267 rg BT 26.250 573.482 Td /F4 12.0 Tf [(Abstract)] TJ ET BT 26.250 553.528 Td /F1 9.8 Tf [(Huntingtons disease \(HD\) is a late-onset, slowly progressing neurodegenerative disorder caused by an expansion of glutamine )] TJ ET BT 26.250 541.623 Td /F1 9.8 Tf [(repeats. The YAC128 mouse model has been widely used to study the progression of HD symptoms, but little is known about )] TJ ET BT 26.250 529.719 Td /F1 9.8 Tf [(synaptic alterations in very old animals. The present experiments examined synaptic properties of striatal medium-sized spiny )] TJ ET BT 26.250 517.814 Td /F1 9.8 Tf [(neurons \(MSNs\) in 16 month-old YAC128 mice. These mice were crossed with mice expressing enhanced green fluorescent )] TJ ET BT 26.250 505.909 Td /F1 9.8 Tf [(protein \(EGFP\) under the control of either D1 or D2 dopamine receptor promoters to identify MSNs originating the direct and )] TJ ET BT 26.250 494.004 Td /F1 9.8 Tf [(indirect pathways, respectively. The input-output curves of evoked excitatory postsynaptic currents mediated by activation of the )] TJ ET BT 26.250 482.100 Td /F1 9.8 Tf [(?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid \(AMPA\) or N-methyl-D-aspartate \(NMDA\) receptors were reduced in )] TJ ET BT 26.250 470.195 Td /F1 9.8 Tf [(MSNs in both pathways. In the presence of DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA\), a glutamate transporter blocker )] TJ ET BT 26.250 458.290 Td /F1 9.8 Tf [(used to increase activation of extrasynaptic receptors, NMDA receptor-mediated currents displayed altered amplitudes, longer )] TJ ET BT 26.250 446.385 Td /F1 9.8 Tf [(decay times, and greater charge \(response areas\) in both direct and indirect pathway MSNs in YAC128 mice compared to )] TJ ET BT 26.250 434.481 Td /F1 9.8 Tf [(wildtype controls. Amplitudes were significantly increased, primarily in direct pathway MSNs while normalized areas were )] TJ ET BT 26.250 422.576 Td /F1 9.8 Tf [(significantly increased only in indirect pathway MSNs, suggesting that the two types of MSNs are affected in different ways. It )] TJ ET BT 26.250 410.671 Td /F1 9.8 Tf [(may be that indirect pathway neurons are more susceptible to changes in glutamate transport. Taken together, the present )] TJ ET BT 26.250 398.766 Td /F1 9.8 Tf [(findings demonstrate differential alterations in synaptic versus extrasynaptic NMDA receptors in both direct and indirect pathway )] TJ ET BT 26.250 386.862 Td /F1 9.8 Tf [(MSNs in late HD, which may contribute to the dysfunction and degeneration in both pathways.)] TJ ET BT 26.250 350.259 Td /F4 12.0 Tf [(Funding Statement)] TJ ET BT 26.250 330.305 Td /F1 9.8 Tf [(Supported by USPHS NS41574. The funders had no role in study design, data collection and analysis, decision to publish, or )] TJ ET BT 26.250 318.400 Td /F1 9.8 Tf [(preparation of the manuscript. The authors have declared that no competing interests exist.)] TJ ET BT 26.250 289.298 Td /F4 12.0 Tf [(Notice of Correction)] TJ ET BT 26.250 269.343 Td /F4 9.8 Tf [(11 July 2017:)] TJ ET BT 86.953 269.343 Td /F1 9.8 Tf [( PLOS Currents Correction: Differential Synaptic and Extrasynaptic Glutamate-Receptor Alterations in Striatal )] TJ ET BT 26.250 257.439 Td /F1 9.8 Tf [(Medium-Sized Spiny Neurons of Aged YAC128 Huntingtons Disease Mice.. PLOS Currents Huntington Disease. 2017 Jul 11 . )] TJ ET BT 26.250 245.534 Td /F1 9.8 Tf [(Edition 1. doi: 10.1371/currents.hd.2f36af340a70a0d248b48d4cc9c3e58e. )] TJ ET 0.267 0.267 0.267 rg BT 349.297 245.534 Td /F1 9.8 Tf [(View Correction.)] TJ ET 0.271 0.267 0.267 rg BT 26.250 208.931 Td /F4 12.0 Tf [(Introduction)] TJ ET BT 26.250 188.977 Td /F1 9.8 Tf [(Huntingtons disease \(HD\), a slowly progressing neurodegenerative disorder caused by an expansion of glutamine \(CAG\) )] TJ ET BT 26.250 177.072 Td /F1 9.8 Tf [(repeats, is characterized by motor, psychiatric and cognitive symptoms)] TJ ET 0.267 0.267 0.267 rg BT 331.854 178.580 Td /F4 8.7 Tf [(1)] TJ ET 0.271 0.267 0.267 rg BT 336.673 177.072 Td /F1 9.8 Tf [(. The YAC128 mouse model has been widely used to )] TJ ET BT 26.250 165.168 Td /F1 9.8 Tf [(examine mechanisms of HD. YAC128 mice carry the entire human HD gene containing 128 CAG repeats on a yeast artificial )] TJ ET BT 26.250 153.263 Td /F1 9.8 Tf [(chromosome \(line 53\) and express full-length human mutant huntingtin protein)] TJ ET 0.267 0.267 0.267 rg BT 363.853 154.770 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 368.672 153.263 Td /F1 9.8 Tf [(. These mice display a hyperkinetic phenotype )] TJ ET BT 26.250 141.358 Td /F1 9.8 Tf [(at 3 months of age, followed by motor deficits at 6 months, with eventual progression to hypokinesia by 12 months)] TJ ET 0.267 0.267 0.267 rg BT 517.777 142.865 Td /F4 8.7 Tf [(3)] TJ ET 0.271 0.267 0.267 rg BT 522.595 141.358 Td /F1 9.8 Tf [(. The )] TJ ET BT 26.250 129.453 Td /F1 9.8 Tf [(behavioral changes are associated with biphasic alterations in excitatory synaptic transmission to striatal medium-sized spiny )] TJ ET BT 26.250 117.549 Td /F1 9.8 Tf [(neurons \(MSNs\) of the direct \(expressing dopamine D1 receptors\) and indirect \(expressing dopamine D2 receptors\) striatal )] TJ ET BT 26.250 105.644 Td /F1 9.8 Tf [(output pathways)] TJ ET 0.267 0.267 0.267 rg BT 97.249 107.151 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 102.068 109.532 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 104.478 107.151 Td /F4 8.7 Tf [(5)] TJ ET 0.271 0.267 0.267 rg BT 109.296 105.644 Td /F1 9.8 Tf [(. At the early, pre-symptomatic stage, glutamate transmission is increased in both direct and indirect )] TJ ET BT 26.250 93.739 Td /F1 9.8 Tf [(pathways. In contrast, at 12 months of age, the more symptomatic phase, glutamatergic transmission is markedly reduced in )] TJ ET BT 26.250 81.834 Td /F1 9.8 Tf [(direct pathway MSNs, while only a trend towards a reduction in glutamatergic inputs is observed in D2 MSNs)] TJ ET 0.267 0.267 0.267 rg BT 494.962 83.342 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 499.780 81.834 Td /F1 9.8 Tf [(. Such biphasic )] TJ ET BT 26.250 69.930 Td /F1 9.8 Tf [(alterations are also observed in evoked ?-amino-3-hydroxy-5-methyl-4-isoxazolepropionic acid receptor \(AMPAR\)-mediated )] TJ ET BT 26.250 58.025 Td /F1 9.8 Tf [(currents, demonstrated by an increase in the peak amplitude in indirect pathway MSNs at 1.5 months and a reduction in direct )] TJ ET BT 26.250 46.120 Td /F1 9.8 Tf [(pathway MSNs at 12 months)] TJ ET 0.267 0.267 0.267 rg BT 150.884 47.627 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 155.703 46.120 Td /F1 9.8 Tf [(. 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BT 26.250 767.476 Td /F1 9.8 Tf [(striatum and a loss of synaptic markets)] TJ ET 0.267 0.267 0.267 rg BT 194.769 768.983 Td /F4 8.7 Tf [(21)] TJ ET 0.271 0.267 0.267 rg BT 204.406 767.476 Td /F1 9.8 Tf [(. In these conditions, release of glutamate is more likely to activate extrasynaptic N-)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(methyl-D-aspartate receptors NMDARs\), which are known to mediate apoptotic cascades)] TJ ET 0.267 0.267 0.267 rg BT 411.502 757.079 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 421.139 755.571 Td /F1 9.8 Tf [(. Indeed, an increase in pro-)] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(apoptotic extrasynaptic NMDAR function occurs in YAC128 MSNs at the presymptomatic and symptomatic stages)] TJ ET 0.267 0.267 0.267 rg BT 517.718 745.174 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 522.537 743.667 Td /F1 9.8 Tf [(. However, )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(differences in extrasynaptic NMDARs from the direct and indirect pathways were not examined. This is important as studies )] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(have shown that MSNs of the indirect pathway are more vulnerable in HD)] TJ ET 0.267 0.267 0.267 rg BT 343.252 721.364 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 348.070 719.857 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(In the present study, we examined synaptic versus extrasynaptic electrophysiological changes induced by activation of AMPA )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(and NMDA receptors in direct and indirect pathway MSNs of 16 month-old YAC128 mice and their age-matched wild type \(WT\) )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(controls. Results revealed differential changes in synaptic and extrasynaptic activity of MSNs which could have important )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(implications to understand the differential vulnerability of MSNs in these two striatal output pathways in HD.)] TJ ET BT 26.250 628.136 Td /F4 12.0 Tf [(Materials and Methods)] TJ ET BT 26.250 608.181 Td /F1 9.8 Tf [(All experimental procedures were performed in accordance with the United States Public Health Service Guide for Care and )] TJ ET BT 26.250 596.277 Td /F1 9.8 Tf [(Use of Laboratory Animals and were approved by the Institutional Animal Care and Use Committee at the University of )] TJ ET BT 26.250 584.372 Td /F1 9.8 Tf [(California, Los Angeles \(UCLA\). Experiments were conducted in WT and YAC128 mice crossbred to FVB/N mice expressing )] TJ ET BT 26.250 572.467 Td /F1 9.8 Tf [(enhanced green fluorescent protein \(EGFP\) under the control of the D1 or D2 dopamine receptor promoter. All mice were )] TJ ET BT 26.250 560.562 Td /F1 9.8 Tf [(obtained from our breeding colonies at UCLA. Each experimental or control group consisted of 5-10 mice.)] TJ ET BT 26.250 541.158 Td /F1 9.8 Tf [(Detailed procedures for mouse slice preparation and electrophysiology have been published previously)] TJ ET 0.267 0.267 0.267 rg BT 470.070 542.665 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 474.889 541.158 Td /F1 9.8 Tf [(. Briefly, the mice were )] TJ ET BT 26.250 529.253 Td /F1 9.8 Tf [(deeply anaesthetized with isoflurane and decapitated. The brains were quickly removed and placed in oxygenated ice-cold low-)] TJ ET BT 26.250 517.348 Td /F1 9.8 Tf [(Ca)] TJ ET BT 38.710 521.236 Td /F1 8.7 Tf [(2+ )] TJ ET BT 51.000 517.348 Td /F1 9.8 Tf [(artificial cerebrospinal fluid \(ACSF\) containing \(in mM\): 130 NaCl, 3 KCl, 1.25 NaH)] TJ ET BT 405.890 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 410.709 517.348 Td /F1 9.8 Tf [(PO)] TJ ET BT 424.798 515.284 Td /F1 8.7 Tf [(4)] TJ ET BT 429.616 517.348 Td /F1 9.8 Tf [(, 26 NaHCO)] TJ ET BT 482.715 515.284 Td /F1 8.7 Tf [(3)] TJ ET BT 487.533 517.348 Td /F1 9.8 Tf [(, 5 MgCl)] TJ ET BT 523.833 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 528.651 517.348 Td /F1 9.8 Tf [(, 1 CaCl)] TJ ET BT 563.868 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 568.687 517.348 Td /F1 9.8 Tf [(, )] TJ ET BT 26.250 505.443 Td /F1 9.8 Tf [(and 10 glucose. Coronal slices \(350 ?m\) were cut and transferred to an incubating chamber containing ACSF \(with 2 mM CaCl)] TJ ET BT 571.918 503.379 Td /F1 8.7 Tf [(2)] TJ ET BT 26.250 493.539 Td /F1 9.8 Tf [(and 2 mM MgCl)] TJ ET BT 95.056 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 99.874 493.539 Td /F1 9.8 Tf [(\) oxygenated with 95% O)] TJ ET BT 208.801 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 213.620 493.539 Td /F1 9.8 Tf [(-5% CO)] TJ ET BT 248.291 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 253.110 493.539 Td /F1 9.8 Tf [( \(pH 7.2-7.4, 290-310 mOsm, 252C\).)] TJ ET BT 26.250 474.134 Td /F1 9.8 Tf [(After 1 h incubation, direct and indirect pathway MSNs were selected for recordings. Cells were visualized using a 40x water-)] TJ ET BT 26.250 462.229 Td /F1 9.8 Tf [(immersion lens on an Olympus microscope \(BX50WI\) equipped with differential interference contrast optics and fluorescence. )] TJ ET BT 26.250 450.324 Td /F1 9.8 Tf [(After identifying MSNs with infrared videomicroscopy \(QICAM-IR Fast 1394, Qimaging\), the filter was switched to fluorescence )] TJ ET BT 26.250 438.420 Td /F1 9.8 Tf [(to determine whether the cell was labeled with EGFP. The digitized infrared image was superimposed over the fluorescence )] TJ ET BT 26.250 426.515 Td /F1 9.8 Tf [(image, and electrophysiological recordings proceeded only if the cell identified with infrared light overlapped with EGFP )] TJ ET BT 26.250 414.610 Td /F1 9.8 Tf [(fluorescence.)] TJ ET BT 26.250 395.205 Td /F1 9.8 Tf [(All experiments were performed in voltage clamp mode, with patch electrodes \(3-5 M?\) filled with the following internal solution )] TJ ET BT 26.250 383.301 Td /F1 9.8 Tf [(\(in mM\): 130 Cs-methanesulfonate, 10 CsCl, 4 NaCl, 1 MgCl)] TJ ET BT 287.950 381.236 Td /F1 8.7 Tf [(2)] TJ ET BT 292.768 383.301 Td /F1 9.8 Tf [(, 5 MgATP, 5 EGTA, 10 HEPES, 0.5 GTP, 10 phosphocreatine, )] TJ ET BT 26.250 371.396 Td /F1 9.8 Tf [(\(pH 7.257.3, osmolality, 280290 mOsm\). QX-314 \(4 mM\) was added to the intracellular solution to prevent Na)] TJ ET BT 509.070 375.284 Td /F1 8.7 Tf [(+ )] TJ ET BT 516.541 371.396 Td /F1 9.8 Tf [(channel )] TJ ET BT 26.250 359.491 Td /F1 9.8 Tf [(activation when stimulating or holding the membrane at depolarized potentials.)] TJ ET BT 26.250 340.086 Td /F1 9.8 Tf [(Synaptic stimulation: To assess input-output functions, a monopolar glass electrode \(impedance 1 M?\) was placed in the corpus )] TJ ET BT 26.250 328.182 Td /F1 9.8 Tf [(callosum 200-300 ?m from the MSN. To examine AMPAR-mediated currents, stimuli of increasing intensities \(0.01-0.2 mA\) )] TJ ET BT 26.250 316.277 Td /F1 9.8 Tf [(were applied every 10 s, while for NMDAR-mediated currents stimuli were applied every 20 s. To record AMPAR-mediated )] TJ ET BT 26.250 304.372 Td /F1 9.8 Tf [(currents, the membrane potential was clamped at -70mV in the presence of bicuculline \(BIC, 10 ?M\). NMDAR-mediated )] TJ ET BT 26.250 292.467 Td /F1 9.8 Tf [(currents were recorded at a holding potential of +40 mV in the presence of BIC \(10 ?M\) and 6-Cyano-7nitroquinoxaline-2,3-)] TJ ET BT 26.250 280.563 Td /F1 9.8 Tf [(dione \(CNQX, 10 ?M\), which blocks AMPAR-mediated currents. Glycine \(20 ?M\) and strychnine \(10 ?M\) also were included to )] TJ ET BT 26.250 268.658 Td /F1 9.8 Tf [(augment NMDAR function and block inhibitory glycine receptors, respectively)] TJ ET 0.267 0.267 0.267 rg BT 358.949 270.165 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 363.768 268.658 Td /F1 9.8 Tf [(. DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA, )] TJ ET BT 26.250 256.753 Td /F1 9.8 Tf [(30 ?M\), a blocker of excitatory amino acid transporters, also was used to promote spillover of glutamate from synaptic receptors )] TJ ET BT 26.250 244.848 Td /F1 9.8 Tf [(onto extrasynaptic NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 148.700 246.356 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 153.519 244.848 Td /F1 9.8 Tf [(. At this concentration, the blocker has no direct effects on NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 447.228 246.356 Td /F4 8.7 Tf [(22)] TJ ET 0.271 0.267 0.267 rg BT 456.865 244.848 Td /F1 9.8 Tf [(. For this experiment, the )] TJ ET BT 26.250 232.944 Td /F1 9.8 Tf [(current intensity that evoked 75% of the peak amplitude response was used. Peak amplitude, area, and decay time were )] TJ ET BT 26.250 221.039 Td /F1 9.8 Tf [(determined using ClampFit software. Decay times were assessed using the 90-10% portion of the decay phase, where 100% )] TJ ET BT 26.250 209.134 Td /F1 9.8 Tf [(was the peak amplitude and 0% was at baseline. Thus, the decay time is the absolute time measurement of the decay phase of )] TJ ET BT 26.250 197.229 Td /F1 9.8 Tf [(the response between 90% and 10% peak amplitude values.)] TJ ET BT 26.250 177.825 Td /F1 9.8 Tf [(Values in figures and text are means SEMs. Differences between group means were assessed with appropriate )] TJ ET BT 518.030 177.825 Td /F5 9.8 Tf [(t)] TJ ET BT 520.741 177.825 Td /F1 9.8 Tf [(-tests and )] TJ ET BT 26.250 165.920 Td /F1 9.8 Tf [(two-way analyses of variance \(ANOVA\) with one repeated measure, followed by Bonferroni )] TJ ET BT 422.363 165.920 Td /F5 9.8 Tf [(post hoc)] TJ ET BT 459.218 165.920 Td /F1 9.8 Tf [( tests. Differences were )] TJ ET BT 26.250 154.015 Td /F1 9.8 Tf [(considered statistically significant if )] TJ ET BT 180.134 154.015 Td /F5 9.8 Tf [(p)] TJ ET BT 185.555 154.015 Td /F1 9.8 Tf [(<0.05.)] TJ ET BT 26.250 117.413 Td /F4 12.0 Tf [(Results)] TJ ET BT 26.250 97.458 Td /F4 9.8 Tf [(Evoked excitatory currents were decreased in YAC128 direct and indirect pathway MSNs.)] TJ ET BT 26.250 78.054 Td /F1 9.8 Tf [(We first examined evoked AMPAR- and NMDAR-mediated currents in WT and YAC128 direct and indirect pathway MSNs in )] TJ ET BT 26.250 66.149 Td /F1 9.8 Tf [(response to increasing stimulus intensities \(Fig. 1\). In YAC128 direct pathway MSNs \(n=9\), there was a significant decrease in )] TJ ET BT 26.250 54.244 Td /F1 9.8 Tf [(peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=8\) \(Genotype: )] TJ ET BT 415.051 54.244 Td /F5 9.8 Tf [(F)] TJ ET BT 421.008 54.244 Td /F1 9.8 Tf [(\(1,15\)=5.029, )] TJ ET BT 481.984 54.244 Td /F5 9.8 Tf [(p)] TJ ET BT 487.405 54.244 Td /F1 9.8 Tf [(=0.04; Interaction of )] TJ ET BT 26.250 42.339 Td /F1 9.8 Tf [(genotype and stimulation intensity: )] TJ ET BT 178.535 42.339 Td /F5 9.8 Tf [(F)] TJ ET BT 184.492 42.339 Td /F1 9.8 Tf [(\(6,90\)=2.952, )] TJ ET BT 245.469 42.339 Td /F5 9.8 Tf [(p)] TJ ET BT 250.890 42.339 Td /F1 9.8 Tf [(=0.011\) \(Fig. 1B, left\). Similarly, YAC128 indirect pathway MSNs \(n=10\) )] TJ ET Q q 15.000 28.054 577.500 748.946 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(striatum and a loss of synaptic markets)] TJ ET 0.267 0.267 0.267 rg BT 194.769 768.983 Td /F4 8.7 Tf [(21)] TJ ET 0.271 0.267 0.267 rg BT 204.406 767.476 Td /F1 9.8 Tf [(. In these conditions, release of glutamate is more likely to activate extrasynaptic N-)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(methyl-D-aspartate receptors NMDARs\), which are known to mediate apoptotic cascades)] TJ ET 0.267 0.267 0.267 rg BT 411.502 757.079 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 421.139 755.571 Td /F1 9.8 Tf [(. Indeed, an increase in pro-)] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(apoptotic extrasynaptic NMDAR function occurs in YAC128 MSNs at the presymptomatic and symptomatic stages)] TJ ET 0.267 0.267 0.267 rg BT 517.718 745.174 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 522.537 743.667 Td /F1 9.8 Tf [(. However, )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(differences in extrasynaptic NMDARs from the direct and indirect pathways were not examined. This is important as studies )] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(have shown that MSNs of the indirect pathway are more vulnerable in HD)] TJ ET 0.267 0.267 0.267 rg BT 343.252 721.364 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 348.070 719.857 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(In the present study, we examined synaptic versus extrasynaptic electrophysiological changes induced by activation of AMPA )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(and NMDA receptors in direct and indirect pathway MSNs of 16 month-old YAC128 mice and their age-matched wild type \(WT\) )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(controls. Results revealed differential changes in synaptic and extrasynaptic activity of MSNs which could have important )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(implications to understand the differential vulnerability of MSNs in these two striatal output pathways in HD.)] TJ ET BT 26.250 628.136 Td /F4 12.0 Tf [(Materials and Methods)] TJ ET BT 26.250 608.181 Td /F1 9.8 Tf [(All experimental procedures were performed in accordance with the United States Public Health Service Guide for Care and )] TJ ET BT 26.250 596.277 Td /F1 9.8 Tf [(Use of Laboratory Animals and were approved by the Institutional Animal Care and Use Committee at the University of )] TJ ET BT 26.250 584.372 Td /F1 9.8 Tf [(California, Los Angeles \(UCLA\). Experiments were conducted in WT and YAC128 mice crossbred to FVB/N mice expressing )] TJ ET BT 26.250 572.467 Td /F1 9.8 Tf [(enhanced green fluorescent protein \(EGFP\) under the control of the D1 or D2 dopamine receptor promoter. All mice were )] TJ ET BT 26.250 560.562 Td /F1 9.8 Tf [(obtained from our breeding colonies at UCLA. Each experimental or control group consisted of 5-10 mice.)] TJ ET BT 26.250 541.158 Td /F1 9.8 Tf [(Detailed procedures for mouse slice preparation and electrophysiology have been published previously)] TJ ET 0.267 0.267 0.267 rg BT 470.070 542.665 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 474.889 541.158 Td /F1 9.8 Tf [(. Briefly, the mice were )] TJ ET BT 26.250 529.253 Td /F1 9.8 Tf [(deeply anaesthetized with isoflurane and decapitated. The brains were quickly removed and placed in oxygenated ice-cold low-)] TJ ET BT 26.250 517.348 Td /F1 9.8 Tf [(Ca)] TJ ET BT 38.710 521.236 Td /F1 8.7 Tf [(2+ )] TJ ET BT 51.000 517.348 Td /F1 9.8 Tf [(artificial cerebrospinal fluid \(ACSF\) containing \(in mM\): 130 NaCl, 3 KCl, 1.25 NaH)] TJ ET BT 405.890 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 410.709 517.348 Td /F1 9.8 Tf [(PO)] TJ ET BT 424.798 515.284 Td /F1 8.7 Tf [(4)] TJ ET BT 429.616 517.348 Td /F1 9.8 Tf [(, 26 NaHCO)] TJ ET BT 482.715 515.284 Td /F1 8.7 Tf [(3)] TJ ET BT 487.533 517.348 Td /F1 9.8 Tf [(, 5 MgCl)] TJ ET BT 523.833 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 528.651 517.348 Td /F1 9.8 Tf [(, 1 CaCl)] TJ ET BT 563.868 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 568.687 517.348 Td /F1 9.8 Tf [(, )] TJ ET BT 26.250 505.443 Td /F1 9.8 Tf [(and 10 glucose. Coronal slices \(350 ?m\) were cut and transferred to an incubating chamber containing ACSF \(with 2 mM CaCl)] TJ ET BT 571.918 503.379 Td /F1 8.7 Tf [(2)] TJ ET BT 26.250 493.539 Td /F1 9.8 Tf [(and 2 mM MgCl)] TJ ET BT 95.056 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 99.874 493.539 Td /F1 9.8 Tf [(\) oxygenated with 95% O)] TJ ET BT 208.801 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 213.620 493.539 Td /F1 9.8 Tf [(-5% CO)] TJ ET BT 248.291 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 253.110 493.539 Td /F1 9.8 Tf [( \(pH 7.2-7.4, 290-310 mOsm, 252C\).)] TJ ET BT 26.250 474.134 Td /F1 9.8 Tf [(After 1 h incubation, direct and indirect pathway MSNs were selected for recordings. Cells were visualized using a 40x water-)] TJ ET BT 26.250 462.229 Td /F1 9.8 Tf [(immersion lens on an Olympus microscope \(BX50WI\) equipped with differential interference contrast optics and fluorescence. )] TJ ET BT 26.250 450.324 Td /F1 9.8 Tf [(After identifying MSNs with infrared videomicroscopy \(QICAM-IR Fast 1394, Qimaging\), the filter was switched to fluorescence )] TJ ET BT 26.250 438.420 Td /F1 9.8 Tf [(to determine whether the cell was labeled with EGFP. The digitized infrared image was superimposed over the fluorescence )] TJ ET BT 26.250 426.515 Td /F1 9.8 Tf [(image, and electrophysiological recordings proceeded only if the cell identified with infrared light overlapped with EGFP )] TJ ET BT 26.250 414.610 Td /F1 9.8 Tf [(fluorescence.)] TJ ET BT 26.250 395.205 Td /F1 9.8 Tf [(All experiments were performed in voltage clamp mode, with patch electrodes \(3-5 M?\) filled with the following internal solution )] TJ ET BT 26.250 383.301 Td /F1 9.8 Tf [(\(in mM\): 130 Cs-methanesulfonate, 10 CsCl, 4 NaCl, 1 MgCl)] TJ ET BT 287.950 381.236 Td /F1 8.7 Tf [(2)] TJ ET BT 292.768 383.301 Td /F1 9.8 Tf [(, 5 MgATP, 5 EGTA, 10 HEPES, 0.5 GTP, 10 phosphocreatine, )] TJ ET BT 26.250 371.396 Td /F1 9.8 Tf [(\(pH 7.257.3, osmolality, 280290 mOsm\). QX-314 \(4 mM\) was added to the intracellular solution to prevent Na)] TJ ET BT 509.070 375.284 Td /F1 8.7 Tf [(+ )] TJ ET BT 516.541 371.396 Td /F1 9.8 Tf [(channel )] TJ ET BT 26.250 359.491 Td /F1 9.8 Tf [(activation when stimulating or holding the membrane at depolarized potentials.)] TJ ET BT 26.250 340.086 Td /F1 9.8 Tf [(Synaptic stimulation: To assess input-output functions, a monopolar glass electrode \(impedance 1 M?\) was placed in the corpus )] TJ ET BT 26.250 328.182 Td /F1 9.8 Tf [(callosum 200-300 ?m from the MSN. To examine AMPAR-mediated currents, stimuli of increasing intensities \(0.01-0.2 mA\) )] TJ ET BT 26.250 316.277 Td /F1 9.8 Tf [(were applied every 10 s, while for NMDAR-mediated currents stimuli were applied every 20 s. To record AMPAR-mediated )] TJ ET BT 26.250 304.372 Td /F1 9.8 Tf [(currents, the membrane potential was clamped at -70mV in the presence of bicuculline \(BIC, 10 ?M\). NMDAR-mediated )] TJ ET BT 26.250 292.467 Td /F1 9.8 Tf [(currents were recorded at a holding potential of +40 mV in the presence of BIC \(10 ?M\) and 6-Cyano-7nitroquinoxaline-2,3-)] TJ ET BT 26.250 280.563 Td /F1 9.8 Tf [(dione \(CNQX, 10 ?M\), which blocks AMPAR-mediated currents. Glycine \(20 ?M\) and strychnine \(10 ?M\) also were included to )] TJ ET BT 26.250 268.658 Td /F1 9.8 Tf [(augment NMDAR function and block inhibitory glycine receptors, respectively)] TJ ET 0.267 0.267 0.267 rg BT 358.949 270.165 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 363.768 268.658 Td /F1 9.8 Tf [(. DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA, )] TJ ET BT 26.250 256.753 Td /F1 9.8 Tf [(30 ?M\), a blocker of excitatory amino acid transporters, also was used to promote spillover of glutamate from synaptic receptors )] TJ ET BT 26.250 244.848 Td /F1 9.8 Tf [(onto extrasynaptic NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 148.700 246.356 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 153.519 244.848 Td /F1 9.8 Tf [(. At this concentration, the blocker has no direct effects on NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 447.228 246.356 Td /F4 8.7 Tf [(22)] TJ ET 0.271 0.267 0.267 rg BT 456.865 244.848 Td /F1 9.8 Tf [(. For this experiment, the )] TJ ET BT 26.250 232.944 Td /F1 9.8 Tf [(current intensity that evoked 75% of the peak amplitude response was used. Peak amplitude, area, and decay time were )] TJ ET BT 26.250 221.039 Td /F1 9.8 Tf [(determined using ClampFit software. Decay times were assessed using the 90-10% portion of the decay phase, where 100% )] TJ ET BT 26.250 209.134 Td /F1 9.8 Tf [(was the peak amplitude and 0% was at baseline. Thus, the decay time is the absolute time measurement of the decay phase of )] TJ ET BT 26.250 197.229 Td /F1 9.8 Tf [(the response between 90% and 10% peak amplitude values.)] TJ ET BT 26.250 177.825 Td /F1 9.8 Tf [(Values in figures and text are means SEMs. Differences between group means were assessed with appropriate )] TJ ET BT 518.030 177.825 Td /F5 9.8 Tf [(t)] TJ ET BT 520.741 177.825 Td /F1 9.8 Tf [(-tests and )] TJ ET BT 26.250 165.920 Td /F1 9.8 Tf [(two-way analyses of variance \(ANOVA\) with one repeated measure, followed by Bonferroni )] TJ ET BT 422.363 165.920 Td /F5 9.8 Tf [(post hoc)] TJ ET BT 459.218 165.920 Td /F1 9.8 Tf [( tests. Differences were )] TJ ET BT 26.250 154.015 Td /F1 9.8 Tf [(considered statistically significant if )] TJ ET BT 180.134 154.015 Td /F5 9.8 Tf [(p)] TJ ET BT 185.555 154.015 Td /F1 9.8 Tf [(<0.05.)] TJ ET BT 26.250 117.413 Td /F4 12.0 Tf [(Results)] TJ ET BT 26.250 97.458 Td /F4 9.8 Tf [(Evoked excitatory currents were decreased in YAC128 direct and indirect pathway MSNs.)] TJ ET BT 26.250 78.054 Td /F1 9.8 Tf [(We first examined evoked AMPAR- and NMDAR-mediated currents in WT and YAC128 direct and indirect pathway MSNs in )] TJ ET BT 26.250 66.149 Td /F1 9.8 Tf [(response to increasing stimulus intensities \(Fig. 1\). In YAC128 direct pathway MSNs \(n=9\), there was a significant decrease in )] TJ ET BT 26.250 54.244 Td /F1 9.8 Tf [(peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=8\) \(Genotype: )] TJ ET BT 415.051 54.244 Td /F5 9.8 Tf [(F)] TJ ET BT 421.008 54.244 Td /F1 9.8 Tf [(\(1,15\)=5.029, )] TJ ET BT 481.984 54.244 Td /F5 9.8 Tf [(p)] TJ ET BT 487.405 54.244 Td /F1 9.8 Tf [(=0.04; Interaction of )] TJ ET BT 26.250 42.339 Td /F1 9.8 Tf [(genotype and stimulation intensity: )] TJ ET BT 178.535 42.339 Td /F5 9.8 Tf [(F)] TJ ET BT 184.492 42.339 Td /F1 9.8 Tf [(\(6,90\)=2.952, )] TJ ET BT 245.469 42.339 Td /F5 9.8 Tf [(p)] TJ ET BT 250.890 42.339 Td /F1 9.8 Tf [(=0.011\) \(Fig. 1B, left\). Similarly, YAC128 indirect pathway MSNs \(n=10\) )] TJ ET Q q 15.000 28.054 577.500 748.946 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(striatum and a loss of synaptic markets)] TJ ET 0.267 0.267 0.267 rg BT 194.769 768.983 Td /F4 8.7 Tf [(21)] TJ ET 0.271 0.267 0.267 rg BT 204.406 767.476 Td /F1 9.8 Tf [(. In these conditions, release of glutamate is more likely to activate extrasynaptic N-)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(methyl-D-aspartate receptors NMDARs\), which are known to mediate apoptotic cascades)] TJ ET 0.267 0.267 0.267 rg BT 411.502 757.079 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 421.139 755.571 Td /F1 9.8 Tf [(. Indeed, an increase in pro-)] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(apoptotic extrasynaptic NMDAR function occurs in YAC128 MSNs at the presymptomatic and symptomatic stages)] TJ ET 0.267 0.267 0.267 rg BT 517.718 745.174 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 522.537 743.667 Td /F1 9.8 Tf [(. However, )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(differences in extrasynaptic NMDARs from the direct and indirect pathways were not examined. This is important as studies )] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(have shown that MSNs of the indirect pathway are more vulnerable in HD)] TJ ET 0.267 0.267 0.267 rg BT 343.252 721.364 Td /F4 8.7 Tf [(2)] TJ ET 0.271 0.267 0.267 rg BT 348.070 719.857 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(In the present study, we examined synaptic versus extrasynaptic electrophysiological changes induced by activation of AMPA )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(and NMDA receptors in direct and indirect pathway MSNs of 16 month-old YAC128 mice and their age-matched wild type \(WT\) )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(controls. Results revealed differential changes in synaptic and extrasynaptic activity of MSNs which could have important )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(implications to understand the differential vulnerability of MSNs in these two striatal output pathways in HD.)] TJ ET BT 26.250 628.136 Td /F4 12.0 Tf [(Materials and Methods)] TJ ET BT 26.250 608.181 Td /F1 9.8 Tf [(All experimental procedures were performed in accordance with the United States Public Health Service Guide for Care and )] TJ ET BT 26.250 596.277 Td /F1 9.8 Tf [(Use of Laboratory Animals and were approved by the Institutional Animal Care and Use Committee at the University of )] TJ ET BT 26.250 584.372 Td /F1 9.8 Tf [(California, Los Angeles \(UCLA\). Experiments were conducted in WT and YAC128 mice crossbred to FVB/N mice expressing )] TJ ET BT 26.250 572.467 Td /F1 9.8 Tf [(enhanced green fluorescent protein \(EGFP\) under the control of the D1 or D2 dopamine receptor promoter. All mice were )] TJ ET BT 26.250 560.562 Td /F1 9.8 Tf [(obtained from our breeding colonies at UCLA. Each experimental or control group consisted of 5-10 mice.)] TJ ET BT 26.250 541.158 Td /F1 9.8 Tf [(Detailed procedures for mouse slice preparation and electrophysiology have been published previously)] TJ ET 0.267 0.267 0.267 rg BT 470.070 542.665 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 474.889 541.158 Td /F1 9.8 Tf [(. Briefly, the mice were )] TJ ET BT 26.250 529.253 Td /F1 9.8 Tf [(deeply anaesthetized with isoflurane and decapitated. The brains were quickly removed and placed in oxygenated ice-cold low-)] TJ ET BT 26.250 517.348 Td /F1 9.8 Tf [(Ca)] TJ ET BT 38.710 521.236 Td /F1 8.7 Tf [(2+ )] TJ ET BT 51.000 517.348 Td /F1 9.8 Tf [(artificial cerebrospinal fluid \(ACSF\) containing \(in mM\): 130 NaCl, 3 KCl, 1.25 NaH)] TJ ET BT 405.890 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 410.709 517.348 Td /F1 9.8 Tf [(PO)] TJ ET BT 424.798 515.284 Td /F1 8.7 Tf [(4)] TJ ET BT 429.616 517.348 Td /F1 9.8 Tf [(, 26 NaHCO)] TJ ET BT 482.715 515.284 Td /F1 8.7 Tf [(3)] TJ ET BT 487.533 517.348 Td /F1 9.8 Tf [(, 5 MgCl)] TJ ET BT 523.833 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 528.651 517.348 Td /F1 9.8 Tf [(, 1 CaCl)] TJ ET BT 563.868 515.284 Td /F1 8.7 Tf [(2)] TJ ET BT 568.687 517.348 Td /F1 9.8 Tf [(, )] TJ ET BT 26.250 505.443 Td /F1 9.8 Tf [(and 10 glucose. Coronal slices \(350 ?m\) were cut and transferred to an incubating chamber containing ACSF \(with 2 mM CaCl)] TJ ET BT 571.918 503.379 Td /F1 8.7 Tf [(2)] TJ ET BT 26.250 493.539 Td /F1 9.8 Tf [(and 2 mM MgCl)] TJ ET BT 95.056 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 99.874 493.539 Td /F1 9.8 Tf [(\) oxygenated with 95% O)] TJ ET BT 208.801 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 213.620 493.539 Td /F1 9.8 Tf [(-5% CO)] TJ ET BT 248.291 491.474 Td /F1 8.7 Tf [(2)] TJ ET BT 253.110 493.539 Td /F1 9.8 Tf [( \(pH 7.2-7.4, 290-310 mOsm, 252C\).)] TJ ET BT 26.250 474.134 Td /F1 9.8 Tf [(After 1 h incubation, direct and indirect pathway MSNs were selected for recordings. Cells were visualized using a 40x water-)] TJ ET BT 26.250 462.229 Td /F1 9.8 Tf [(immersion lens on an Olympus microscope \(BX50WI\) equipped with differential interference contrast optics and fluorescence. )] TJ ET BT 26.250 450.324 Td /F1 9.8 Tf [(After identifying MSNs with infrared videomicroscopy \(QICAM-IR Fast 1394, Qimaging\), the filter was switched to fluorescence )] TJ ET BT 26.250 438.420 Td /F1 9.8 Tf [(to determine whether the cell was labeled with EGFP. The digitized infrared image was superimposed over the fluorescence )] TJ ET BT 26.250 426.515 Td /F1 9.8 Tf [(image, and electrophysiological recordings proceeded only if the cell identified with infrared light overlapped with EGFP )] TJ ET BT 26.250 414.610 Td /F1 9.8 Tf [(fluorescence.)] TJ ET BT 26.250 395.205 Td /F1 9.8 Tf [(All experiments were performed in voltage clamp mode, with patch electrodes \(3-5 M?\) filled with the following internal solution )] TJ ET BT 26.250 383.301 Td /F1 9.8 Tf [(\(in mM\): 130 Cs-methanesulfonate, 10 CsCl, 4 NaCl, 1 MgCl)] TJ ET BT 287.950 381.236 Td /F1 8.7 Tf [(2)] TJ ET BT 292.768 383.301 Td /F1 9.8 Tf [(, 5 MgATP, 5 EGTA, 10 HEPES, 0.5 GTP, 10 phosphocreatine, )] TJ ET BT 26.250 371.396 Td /F1 9.8 Tf [(\(pH 7.257.3, osmolality, 280290 mOsm\). QX-314 \(4 mM\) was added to the intracellular solution to prevent Na)] TJ ET BT 509.070 375.284 Td /F1 8.7 Tf [(+ )] TJ ET BT 516.541 371.396 Td /F1 9.8 Tf [(channel )] TJ ET BT 26.250 359.491 Td /F1 9.8 Tf [(activation when stimulating or holding the membrane at depolarized potentials.)] TJ ET BT 26.250 340.086 Td /F1 9.8 Tf [(Synaptic stimulation: To assess input-output functions, a monopolar glass electrode \(impedance 1 M?\) was placed in the corpus )] TJ ET BT 26.250 328.182 Td /F1 9.8 Tf [(callosum 200-300 ?m from the MSN. To examine AMPAR-mediated currents, stimuli of increasing intensities \(0.01-0.2 mA\) )] TJ ET BT 26.250 316.277 Td /F1 9.8 Tf [(were applied every 10 s, while for NMDAR-mediated currents stimuli were applied every 20 s. To record AMPAR-mediated )] TJ ET BT 26.250 304.372 Td /F1 9.8 Tf [(currents, the membrane potential was clamped at -70mV in the presence of bicuculline \(BIC, 10 ?M\). NMDAR-mediated )] TJ ET BT 26.250 292.467 Td /F1 9.8 Tf [(currents were recorded at a holding potential of +40 mV in the presence of BIC \(10 ?M\) and 6-Cyano-7nitroquinoxaline-2,3-)] TJ ET BT 26.250 280.563 Td /F1 9.8 Tf [(dione \(CNQX, 10 ?M\), which blocks AMPAR-mediated currents. Glycine \(20 ?M\) and strychnine \(10 ?M\) also were included to )] TJ ET BT 26.250 268.658 Td /F1 9.8 Tf [(augment NMDAR function and block inhibitory glycine receptors, respectively)] TJ ET 0.267 0.267 0.267 rg BT 358.949 270.165 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 363.768 268.658 Td /F1 9.8 Tf [(. DL-threo-?-Benzyloxyaspartic acid \(DL-TBOA, )] TJ ET BT 26.250 256.753 Td /F1 9.8 Tf [(30 ?M\), a blocker of excitatory amino acid transporters, also was used to promote spillover of glutamate from synaptic receptors )] TJ ET BT 26.250 244.848 Td /F1 9.8 Tf [(onto extrasynaptic NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 148.700 246.356 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 153.519 244.848 Td /F1 9.8 Tf [(. At this concentration, the blocker has no direct effects on NMDARs)] TJ ET 0.267 0.267 0.267 rg BT 447.228 246.356 Td /F4 8.7 Tf [(22)] TJ ET 0.271 0.267 0.267 rg BT 456.865 244.848 Td /F1 9.8 Tf [(. For this experiment, the )] TJ ET BT 26.250 232.944 Td /F1 9.8 Tf [(current intensity that evoked 75% of the peak amplitude response was used. Peak amplitude, area, and decay time were )] TJ ET BT 26.250 221.039 Td /F1 9.8 Tf [(determined using ClampFit software. Decay times were assessed using the 90-10% portion of the decay phase, where 100% )] TJ ET BT 26.250 209.134 Td /F1 9.8 Tf [(was the peak amplitude and 0% was at baseline. Thus, the decay time is the absolute time measurement of the decay phase of )] TJ ET BT 26.250 197.229 Td /F1 9.8 Tf [(the response between 90% and 10% peak amplitude values.)] TJ ET BT 26.250 177.825 Td /F1 9.8 Tf [(Values in figures and text are means SEMs. Differences between group means were assessed with appropriate )] TJ ET BT 518.030 177.825 Td /F5 9.8 Tf [(t)] TJ ET BT 520.741 177.825 Td /F1 9.8 Tf [(-tests and )] TJ ET BT 26.250 165.920 Td /F1 9.8 Tf [(two-way analyses of variance \(ANOVA\) with one repeated measure, followed by Bonferroni )] TJ ET BT 422.363 165.920 Td /F5 9.8 Tf [(post hoc)] TJ ET BT 459.218 165.920 Td /F1 9.8 Tf [( tests. Differences were )] TJ ET BT 26.250 154.015 Td /F1 9.8 Tf [(considered statistically significant if )] TJ ET BT 180.134 154.015 Td /F5 9.8 Tf [(p)] TJ ET BT 185.555 154.015 Td /F1 9.8 Tf [(<0.05.)] TJ ET BT 26.250 117.413 Td /F4 12.0 Tf [(Results)] TJ ET BT 26.250 97.458 Td /F4 9.8 Tf [(Evoked excitatory currents were decreased in YAC128 direct and indirect pathway MSNs.)] TJ ET BT 26.250 78.054 Td /F1 9.8 Tf [(We first examined evoked AMPAR- and NMDAR-mediated currents in WT and YAC128 direct and indirect pathway MSNs in )] TJ ET BT 26.250 66.149 Td /F1 9.8 Tf [(response to increasing stimulus intensities \(Fig. 1\). In YAC128 direct pathway MSNs \(n=9\), there was a significant decrease in )] TJ ET BT 26.250 54.244 Td /F1 9.8 Tf [(peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=8\) \(Genotype: )] TJ ET BT 415.051 54.244 Td /F5 9.8 Tf [(F)] TJ ET BT 421.008 54.244 Td /F1 9.8 Tf [(\(1,15\)=5.029, )] TJ ET BT 481.984 54.244 Td /F5 9.8 Tf [(p)] TJ ET BT 487.405 54.244 Td /F1 9.8 Tf [(=0.04; Interaction of )] TJ ET BT 26.250 42.339 Td /F1 9.8 Tf [(genotype and stimulation intensity: )] TJ ET BT 178.535 42.339 Td /F5 9.8 Tf [(F)] TJ ET BT 184.492 42.339 Td /F1 9.8 Tf [(\(6,90\)=2.952, )] TJ ET BT 245.469 42.339 Td /F5 9.8 Tf [(p)] TJ ET BT 250.890 42.339 Td /F1 9.8 Tf [(=0.011\) \(Fig. 1B, left\). Similarly, YAC128 indirect pathway MSNs \(n=10\) )] TJ ET Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(2)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Huntington Disease)] TJ ET Q endstream endobj 130 0 obj << /Type /Font /Subtype /Type1 /Name /F5 /BaseFont /Helvetica-Oblique /Encoding /WinAnsiEncoding >> endobj 131 0 obj << /Type /Annot /Subtype /Link /A 132 0 R /Border [0 0 0] /H /I /Rect [ 194.7690 768.1817 204.4063 777.0000 ] >> endobj 132 0 obj << /Type /Action >> endobj 133 0 obj << /Type /Annot /Subtype /Link /A 134 0 R /Border [0 0 0] /H /I /Rect [ 411.5018 756.2769 421.1391 765.0952 ] >> endobj 134 0 obj << /Type /Action >> endobj 135 0 obj << /Type /Annot /Subtype /Link /A 136 0 R /Border [0 0 0] /H /I /Rect [ 517.7183 744.3722 522.5369 753.1905 ] >> endobj 136 0 obj << /Type /Action >> endobj 137 0 obj << /Type /Annot /Subtype /Link /A 138 0 R /Border [0 0 0] /H /I /Rect [ 343.2518 720.5627 348.0704 729.3810 ] >> endobj 138 0 obj << /Type /Action >> endobj 139 0 obj << /Type /Annot /Subtype /Link /A 140 0 R /Border [0 0 0] /H /I /Rect [ 470.0700 541.8632 474.8887 550.6815 ] >> endobj 140 0 obj << /Type /Action >> endobj 141 0 obj << /Type /Annot /Subtype /Link /A 142 0 R /Border [0 0 0] /H /I /Rect [ 358.9493 269.3634 363.7679 278.1818 ] >> endobj 142 0 obj << /Type /Action >> endobj 143 0 obj << /Type /Annot /Subtype /Link /A 144 0 R /Border [0 0 0] /H /I /Rect [ 148.7002 245.5539 153.5189 254.3723 ] >> endobj 144 0 obj << /Type /Action >> endobj 145 0 obj << /Type /Annot /Subtype /Link /A 146 0 R /Border [0 0 0] /H /I /Rect [ 447.2279 245.5539 456.8653 254.3723 ] >> endobj 146 0 obj << /Type /Action >> endobj 147 0 obj << /Type /Annot /Subtype /Link /A 148 0 R /Border [0 0 0] /H /I /Rect [ 194.7690 768.1817 204.4063 777.0000 ] >> endobj 148 0 obj << /Type /Action >> endobj 149 0 obj << /Type /Annot /Subtype /Link /A 150 0 R /Border [0 0 0] /H /I /Rect [ 411.5018 756.2769 421.1391 765.0952 ] >> endobj 150 0 obj << /Type /Action >> endobj 151 0 obj << /Type /Annot /Subtype /Link /A 152 0 R /Border [0 0 0] /H /I /Rect [ 517.7183 744.3722 522.5369 753.1905 ] >> endobj 152 0 obj << /Type /Action >> endobj 153 0 obj << /Type /Annot /Subtype /Link /A 154 0 R /Border [0 0 0] /H /I /Rect [ 343.2518 720.5627 348.0704 729.3810 ] >> endobj 154 0 obj << /Type /Action >> endobj 155 0 obj << /Type /Annot /Subtype /Link /A 156 0 R /Border [0 0 0] /H /I /Rect [ 470.0700 541.8632 474.8887 550.6815 ] >> endobj 156 0 obj << /Type /Action >> endobj 157 0 obj << /Type /Annot /Subtype /Link /A 158 0 R /Border [0 0 0] /H /I /Rect [ 358.9493 269.3634 363.7679 278.1818 ] >> endobj 158 0 obj << /Type /Action >> endobj 159 0 obj << /Type /Annot /Subtype /Link /A 160 0 R /Border [0 0 0] /H /I /Rect [ 148.7002 245.5539 153.5189 254.3723 ] >> endobj 160 0 obj << /Type /Action >> endobj 161 0 obj << /Type /Annot /Subtype /Link /A 162 0 R /Border [0 0 0] /H /I /Rect [ 447.2279 245.5539 456.8653 254.3723 ] >> endobj 162 0 obj << /Type /Action >> endobj 163 0 obj << /Type /Annot /Subtype /Link /A 164 0 R /Border [0 0 0] /H /I /Rect [ 194.7690 768.1817 204.4063 777.0000 ] >> endobj 164 0 obj << /Type /Action >> endobj 165 0 obj << /Type /Annot /Subtype /Link /A 166 0 R /Border [0 0 0] /H /I /Rect [ 411.5018 756.2769 421.1391 765.0952 ] >> endobj 166 0 obj << /Type /Action >> endobj 167 0 obj << /Type /Annot /Subtype /Link /A 168 0 R /Border [0 0 0] /H /I /Rect [ 517.7183 744.3722 522.5369 753.1905 ] >> endobj 168 0 obj << /Type /Action >> endobj 169 0 obj << /Type /Annot /Subtype /Link /A 170 0 R /Border [0 0 0] /H /I /Rect [ 343.2518 720.5627 348.0704 729.3810 ] >> endobj 170 0 obj << /Type /Action >> endobj 171 0 obj << /Type /Annot /Subtype /Link /A 172 0 R /Border [0 0 0] /H /I /Rect [ 470.0700 541.8632 474.8887 550.6815 ] >> endobj 172 0 obj << /Type /Action >> endobj 173 0 obj << /Type /Annot /Subtype /Link /A 174 0 R /Border [0 0 0] /H /I /Rect [ 358.9493 269.3634 363.7679 278.1818 ] >> endobj 174 0 obj << /Type /Action >> endobj 175 0 obj << /Type /Annot /Subtype /Link /A 176 0 R /Border [0 0 0] /H /I /Rect [ 148.7002 245.5539 153.5189 254.3723 ] >> endobj 176 0 obj << /Type /Action >> endobj 177 0 obj << /Type /Annot /Subtype /Link /A 178 0 R /Border [0 0 0] /H /I /Rect [ 447.2279 245.5539 456.8653 254.3723 ] >> endobj 178 0 obj << /Type /Action >> endobj 179 0 obj << /Type /Page /Parent 3 0 R /Annots [ 181 0 R 185 0 R 189 0 R ] /Contents 180 0 R >> endobj 180 0 obj << /Length 11360 >> stream 0.271 0.267 0.267 rg q 15.000 20.548 577.500 756.452 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(displayed significant decreases in peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=15\) \(Genotype: )] TJ ET BT 568.409 767.476 Td /F5 9.8 Tf [(F)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(\(1,20\)=7.877, )] TJ ET BT 87.227 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 92.648 755.571 Td /F1 9.8 Tf [(=0.011; Interaction of genotype and stimulation intensity: )] TJ ET BT 339.518 755.571 Td /F5 9.8 Tf [(F)] TJ ET BT 345.475 755.571 Td /F1 9.8 Tf [(\(6,120\)=4.200, )] TJ ET BT 411.872 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 417.293 755.571 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1B, right\). Post-hoc )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(analysis revealed that decreases in AMPAR- mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 543.195 743.667 Td /F5 9.8 Tf [(p)] TJ ET BT 548.616 743.667 Td /F1 9.8 Tf [(=0.05-)] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(0.008\) for direct pathway MSNs and at the 0.04-0.20 mA intensities \()] TJ ET BT 322.133 731.762 Td /F5 9.8 Tf [(p)] TJ ET BT 327.554 731.762 Td /F1 9.8 Tf [(=0.019-0.001\) for indirect pathway MSNs. Peak NMDAR-)] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(mediated currents also were significantly decreased for direct pathway MSNs from YAC128 mice \(Genotype: )] TJ ET BT 497.126 719.857 Td /F5 9.8 Tf [(F)] TJ ET BT 503.084 719.857 Td /F1 9.8 Tf [(\(1,17\)=5.285, )] TJ ET BT 564.060 719.857 Td /F5 9.8 Tf [(p)] TJ ET BT 26.250 707.952 Td /F1 9.8 Tf [(=0.034; Interaction of genotype and stimulation intensity: )] TJ ET BT 273.120 707.952 Td /F5 9.8 Tf [(F)] TJ ET BT 279.077 707.952 Td /F1 9.8 Tf [(\(6,102\)=4.874, )] TJ ET BT 345.475 707.952 Td /F5 9.8 Tf [(p)] TJ ET BT 350.896 707.952 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1D, left\). )] TJ ET BT 448.708 707.952 Td /F5 9.8 Tf [(Post hoc)] TJ ET BT 486.645 707.952 Td /F1 9.8 Tf [( analysis revealed )] TJ ET BT 26.250 696.048 Td /F1 9.8 Tf [(that decreases in NMDAR-mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 463.528 696.048 Td /F5 9.8 Tf [(p)] TJ ET BT 468.949 696.048 Td /F1 9.8 Tf [(=0.017-0.004\). There )] TJ ET BT 26.250 684.143 Td /F1 9.8 Tf [(was a non-significant but strong trend for a decrease in peak NMDAR-mediated currents between WT and YAC128 indirect )] TJ ET BT 26.250 672.238 Td /F1 9.8 Tf [(pathway MSNs \(Genotype: )] TJ ET BT 145.463 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 151.421 672.238 Td /F1 9.8 Tf [(\(1,23\)=3.844, )] TJ ET BT 212.397 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 217.818 672.238 Td /F1 9.8 Tf [(=0.062; Interaction of genotype and stimulation intensity: )] TJ ET BT 464.688 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 470.645 672.238 Td /F1 9.8 Tf [(\(6,138\)=1.955, )] TJ ET BT 537.043 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 542.464 672.238 Td /F1 9.8 Tf [(<0.076\). )] TJ ET BT 26.250 660.333 Td /F1 9.8 Tf [(The)] TJ ET BT 43.049 660.333 Td /F5 9.8 Tf [( post hoc)] TJ ET BT 82.615 660.333 Td /F1 9.8 Tf [( analysis revealed a significant difference in the peak NMDAR-mediated currents at the 0.04 mA stimulation )] TJ ET BT 26.250 648.429 Td /F1 9.8 Tf [(intensity \()] TJ ET BT 67.970 648.429 Td /F5 9.8 Tf [(t)] TJ ET BT 70.681 648.429 Td /F1 9.8 Tf [(=2.787, )] TJ ET BT 106.190 648.429 Td /F5 9.8 Tf [(p)] TJ ET BT 111.611 648.429 Td /F1 9.8 Tf [(=0.008\) \(Fig. 1D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 20.548 555.000 618.000 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 638.548 m 581.250 638.548 l 581.250 637.798 l 26.250 637.798 l f q 450.000 0 0 602.250 35.250 26.548 cm /I4 Do Q q 35.250 20.548 537.000 0.000 re W n Q Q q 15.000 20.548 577.500 756.452 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(displayed significant decreases in peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=15\) \(Genotype: )] TJ ET BT 568.409 767.476 Td /F5 9.8 Tf [(F)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(\(1,20\)=7.877, )] TJ ET BT 87.227 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 92.648 755.571 Td /F1 9.8 Tf [(=0.011; Interaction of genotype and stimulation intensity: )] TJ ET BT 339.518 755.571 Td /F5 9.8 Tf [(F)] TJ ET BT 345.475 755.571 Td /F1 9.8 Tf [(\(6,120\)=4.200, )] TJ ET BT 411.872 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 417.293 755.571 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1B, right\). Post-hoc )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(analysis revealed that decreases in AMPAR- mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 543.195 743.667 Td /F5 9.8 Tf [(p)] TJ ET BT 548.616 743.667 Td /F1 9.8 Tf [(=0.05-)] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(0.008\) for direct pathway MSNs and at the 0.04-0.20 mA intensities \()] TJ ET BT 322.133 731.762 Td /F5 9.8 Tf [(p)] TJ ET BT 327.554 731.762 Td /F1 9.8 Tf [(=0.019-0.001\) for indirect pathway MSNs. Peak NMDAR-)] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(mediated currents also were significantly decreased for direct pathway MSNs from YAC128 mice \(Genotype: )] TJ ET BT 497.126 719.857 Td /F5 9.8 Tf [(F)] TJ ET BT 503.084 719.857 Td /F1 9.8 Tf [(\(1,17\)=5.285, )] TJ ET BT 564.060 719.857 Td /F5 9.8 Tf [(p)] TJ ET BT 26.250 707.952 Td /F1 9.8 Tf [(=0.034; Interaction of genotype and stimulation intensity: )] TJ ET BT 273.120 707.952 Td /F5 9.8 Tf [(F)] TJ ET BT 279.077 707.952 Td /F1 9.8 Tf [(\(6,102\)=4.874, )] TJ ET BT 345.475 707.952 Td /F5 9.8 Tf [(p)] TJ ET BT 350.896 707.952 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1D, left\). )] TJ ET BT 448.708 707.952 Td /F5 9.8 Tf [(Post hoc)] TJ ET BT 486.645 707.952 Td /F1 9.8 Tf [( analysis revealed )] TJ ET BT 26.250 696.048 Td /F1 9.8 Tf [(that decreases in NMDAR-mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 463.528 696.048 Td /F5 9.8 Tf [(p)] TJ ET BT 468.949 696.048 Td /F1 9.8 Tf [(=0.017-0.004\). There )] TJ ET BT 26.250 684.143 Td /F1 9.8 Tf [(was a non-significant but strong trend for a decrease in peak NMDAR-mediated currents between WT and YAC128 indirect )] TJ ET BT 26.250 672.238 Td /F1 9.8 Tf [(pathway MSNs \(Genotype: )] TJ ET BT 145.463 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 151.421 672.238 Td /F1 9.8 Tf [(\(1,23\)=3.844, )] TJ ET BT 212.397 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 217.818 672.238 Td /F1 9.8 Tf [(=0.062; Interaction of genotype and stimulation intensity: )] TJ ET BT 464.688 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 470.645 672.238 Td /F1 9.8 Tf [(\(6,138\)=1.955, )] TJ ET BT 537.043 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 542.464 672.238 Td /F1 9.8 Tf [(<0.076\). )] TJ ET BT 26.250 660.333 Td /F1 9.8 Tf [(The)] TJ ET BT 43.049 660.333 Td /F5 9.8 Tf [( post hoc)] TJ ET BT 82.615 660.333 Td /F1 9.8 Tf [( analysis revealed a significant difference in the peak NMDAR-mediated currents at the 0.04 mA stimulation )] TJ ET BT 26.250 648.429 Td /F1 9.8 Tf [(intensity \()] TJ ET BT 67.970 648.429 Td /F5 9.8 Tf [(t)] TJ ET BT 70.681 648.429 Td /F1 9.8 Tf [(=2.787, )] TJ ET BT 106.190 648.429 Td /F5 9.8 Tf [(p)] TJ ET BT 111.611 648.429 Td /F1 9.8 Tf [(=0.008\) \(Fig. 1D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 20.548 555.000 618.000 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 638.548 m 581.250 638.548 l 581.250 637.798 l 26.250 637.798 l f q 450.000 0 0 602.250 35.250 26.548 cm /I6 Do Q q 35.250 20.548 537.000 0.000 re W n Q Q q 15.000 20.548 577.500 756.452 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(displayed significant decreases in peak amplitude of evoked AMPAR-mediated currents compared to WTs \(n=15\) \(Genotype: )] TJ ET BT 568.409 767.476 Td /F5 9.8 Tf [(F)] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(\(1,20\)=7.877, )] TJ ET BT 87.227 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 92.648 755.571 Td /F1 9.8 Tf [(=0.011; Interaction of genotype and stimulation intensity: )] TJ ET BT 339.518 755.571 Td /F5 9.8 Tf [(F)] TJ ET BT 345.475 755.571 Td /F1 9.8 Tf [(\(6,120\)=4.200, )] TJ ET BT 411.872 755.571 Td /F5 9.8 Tf [(p)] TJ ET BT 417.293 755.571 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1B, right\). Post-hoc )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(analysis revealed that decreases in AMPAR- mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 543.195 743.667 Td /F5 9.8 Tf [(p)] TJ ET BT 548.616 743.667 Td /F1 9.8 Tf [(=0.05-)] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(0.008\) for direct pathway MSNs and at the 0.04-0.20 mA intensities \()] TJ ET BT 322.133 731.762 Td /F5 9.8 Tf [(p)] TJ ET BT 327.554 731.762 Td /F1 9.8 Tf [(=0.019-0.001\) for indirect pathway MSNs. Peak NMDAR-)] TJ ET BT 26.250 719.857 Td /F1 9.8 Tf [(mediated currents also were significantly decreased for direct pathway MSNs from YAC128 mice \(Genotype: )] TJ ET BT 497.126 719.857 Td /F5 9.8 Tf [(F)] TJ ET BT 503.084 719.857 Td /F1 9.8 Tf [(\(1,17\)=5.285, )] TJ ET BT 564.060 719.857 Td /F5 9.8 Tf [(p)] TJ ET BT 26.250 707.952 Td /F1 9.8 Tf [(=0.034; Interaction of genotype and stimulation intensity: )] TJ ET BT 273.120 707.952 Td /F5 9.8 Tf [(F)] TJ ET BT 279.077 707.952 Td /F1 9.8 Tf [(\(6,102\)=4.874, )] TJ ET BT 345.475 707.952 Td /F5 9.8 Tf [(p)] TJ ET BT 350.896 707.952 Td /F1 9.8 Tf [(<0.001\) \(Fig. 1D, left\). )] TJ ET BT 448.708 707.952 Td /F5 9.8 Tf [(Post hoc)] TJ ET BT 486.645 707.952 Td /F1 9.8 Tf [( analysis revealed )] TJ ET BT 26.250 696.048 Td /F1 9.8 Tf [(that decreases in NMDAR-mediated currents were significant at 0.06-0.20 mA stimulation intensities \()] TJ ET BT 463.528 696.048 Td /F5 9.8 Tf [(p)] TJ ET BT 468.949 696.048 Td /F1 9.8 Tf [(=0.017-0.004\). There )] TJ ET BT 26.250 684.143 Td /F1 9.8 Tf [(was a non-significant but strong trend for a decrease in peak NMDAR-mediated currents between WT and YAC128 indirect )] TJ ET BT 26.250 672.238 Td /F1 9.8 Tf [(pathway MSNs \(Genotype: )] TJ ET BT 145.463 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 151.421 672.238 Td /F1 9.8 Tf [(\(1,23\)=3.844, )] TJ ET BT 212.397 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 217.818 672.238 Td /F1 9.8 Tf [(=0.062; Interaction of genotype and stimulation intensity: )] TJ ET BT 464.688 672.238 Td /F5 9.8 Tf [(F)] TJ ET BT 470.645 672.238 Td /F1 9.8 Tf [(\(6,138\)=1.955, )] TJ ET BT 537.043 672.238 Td /F5 9.8 Tf [(p)] TJ ET BT 542.464 672.238 Td /F1 9.8 Tf [(<0.076\). )] TJ ET BT 26.250 660.333 Td /F1 9.8 Tf [(The)] TJ ET BT 43.049 660.333 Td /F5 9.8 Tf [( post hoc)] TJ ET BT 82.615 660.333 Td /F1 9.8 Tf [( analysis revealed a significant difference in the peak NMDAR-mediated currents at the 0.04 mA stimulation )] TJ ET BT 26.250 648.429 Td /F1 9.8 Tf [(intensity \()] TJ ET BT 67.970 648.429 Td /F5 9.8 Tf [(t)] TJ ET BT 70.681 648.429 Td /F1 9.8 Tf [(=2.787, )] TJ ET BT 106.190 648.429 Td /F5 9.8 Tf [(p)] TJ ET BT 111.611 648.429 Td /F1 9.8 Tf [(=0.008\) \(Fig. 1D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 20.548 555.000 618.000 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 638.548 m 581.250 638.548 l 581.250 637.798 l 26.250 637.798 l f q 450.000 0 0 602.250 35.250 26.548 cm /I8 Do Q q 35.250 20.548 537.000 0.000 re W n Q Q q 450.000 0 0 602.250 35.250 26.548 cm /I10 Do Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(3)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Huntington Disease)] TJ ET Q endstream endobj 181 0 obj << /Type /Annot /Subtype /Link /A 182 0 R /Border [0 0 0] /H /I /Rect [ 35.2500 26.5477 485.2500 628.7977 ] >> endobj 182 0 obj << /Type /Action /S /URI /URI (https://currents.plos.org/hd/files/2014/03/Figure1.png) >> endobj 183 0 obj << /Type /XObject /Subtype /Image /Width 600 /Height 803 /Filter /FlateDecode /DecodeParms << /Predictor 15 /Colors 1 /Columns 600 /BitsPerComponent 8>> /ColorSpace /DeviceGray /BitsPerComponent 8 /Length 41342>> stream xD;asLff$I$$I2I$I$I$I&I$I&IL$I$I$Ilֶ}mww},G O>1y+OOxm$[En|tp[>j Y Ŭ@:?2ˋL_/T'Ou}UܲZkii{*kvDXXsƃ>a ظlmiXzZB00P\}¢w볊VF7;T'o&y3~D^DX}P",:Cba|V?!M> ܪ>ȏ]jA nR }" n߅%w\&V# SAzr]FqCw],p.JN!uzF$aA?[r;OJ:(I!O 9[Iʚ$෈T'gкV]:Z] =ޅŬ<62[;3; 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In this and other figures * indicates p<0.05, ** indicates p<0.01, *** indicates p<0.001. See text for )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(details of statistics.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(There were no significant differences in peak AMPAR- or NMDAR-mediated currents between direct or indirect pathway MSNs )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(from either WT or YAC128 mice. Furthermore, there were no differences in NMDA/AMPA ratios among direct and indirect )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(pathway MSNs in WT and YAC128 mice \(0.510.074 vs. 0.440.08 for WTs and YAC128 from direct pathway MSNs, )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(respectively, p=0.542; 0.450.047 vs. 0.560.056 for WTs and YAC128 from indirect pathway MSNs, respectively, p=0.135\).)] TJ ET BT 26.250 560.144 Td /F4 9.8 Tf [(Glutamate transport block revealed alterations in extrasynaptic NMDAR-mediated responses in YAC128 MSNs.)] TJ ET BT 26.250 540.739 Td /F1 9.8 Tf [(We used DL-TBOA \(30 ?M\), a glutamate transport blocker which promotes activation of extrasynaptic NMDARs through )] TJ ET BT 26.250 528.834 Td /F1 9.8 Tf [(glutamate spillover, to assess differences in evoked NMDAR-mediated currents in direct and indirect pathway MSNs. We )] TJ ET BT 26.250 516.930 Td /F1 9.8 Tf [(examined alterations in four indices, peak amplitude, decay time, response area and response area normalized by peak )] TJ ET BT 26.250 505.025 Td /F1 9.8 Tf [(response separately for direct or indirect pathway MSNs \(Fig.2\). An increase in the contribution from extrasynaptic NMDA )] TJ ET BT 26.250 493.120 Td /F1 9.8 Tf [(receptors would be shown by changes in peak amplitude, decay time and response area of the currents. We only tested the )] TJ ET BT 26.250 481.215 Td /F1 9.8 Tf [(effects of TBOA at the stimulus intensity that produced 75% of the maximum response for each neuron. In WTs for both direct )] TJ ET BT 26.250 469.311 Td /F1 9.8 Tf [(and indirect pathway MSNs, peak amplitude decreased significantly in TBOA \()] TJ ET BT 363.298 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 366.008 469.311 Td /F1 9.8 Tf [(=2.177, )] TJ ET BT 401.518 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 406.939 469.311 Td /F1 9.8 Tf [(=0.047; )] TJ ET BT 442.448 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 445.159 469.311 Td /F1 9.8 Tf [(=2.684, )] TJ ET BT 480.668 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 486.089 469.311 Td /F1 9.8 Tf [(=0.016 for direct and )] TJ ET BT 26.250 457.406 Td /F1 9.8 Tf [(indirect pathway MSNs, respectively\) \(Fig. 2B\). In both direct and indirect pathway MSNs, almost all neurons \(6/7 and 9/10 for )] TJ ET BT 26.250 445.501 Td /F1 9.8 Tf [(direct and indirect pathway MSNs, respectively\) showed a decrease in the peak response amplitude although for each group )] TJ ET BT 26.250 433.596 Td /F1 9.8 Tf [(there was considerable variation in the absolute amplitudes of the responses of the neurons. In contrast, in direct pathway )] TJ ET BT 26.250 421.692 Td /F1 9.8 Tf [(MSNs of YAC128 mice, peak amplitude increased significantly \()] TJ ET BT 301.512 421.692 Td /F5 9.8 Tf [(t)] TJ ET BT 304.223 421.692 Td /F1 9.8 Tf [(=4.128, )] TJ ET BT 339.732 421.692 Td /F5 9.8 Tf [(p)] TJ ET BT 345.153 421.692 Td /F1 9.8 Tf [(=0.001; 8/9 neurons\) while in indirect pathway MSNs )] TJ ET BT 26.250 409.787 Td /F1 9.8 Tf [(peak amplitude decreased significantly \()] TJ ET BT 199.654 409.787 Td /F5 9.8 Tf [(t)] TJ ET BT 202.364 409.787 Td /F1 9.8 Tf [(=6.017, )] TJ ET BT 237.874 409.787 Td /F5 9.8 Tf [(p)] TJ ET BT 243.295 409.787 Td /F1 9.8 Tf [(<0.001; 7/7 neurons\) \(Fig. 2B\).)] TJ ET BT 26.250 390.382 Td /F1 9.8 Tf [(Decay time increased significantly for both direct and indirect pathway MSNs in WTs and YAC128 mice \()] TJ ET BT 477.070 390.382 Td /F5 9.8 Tf [(t)] TJ ET BT 479.781 390.382 Td /F1 9.8 Tf [(=2.817, )] TJ ET BT 515.291 390.382 Td /F5 9.8 Tf [(p)] TJ ET BT 520.712 390.382 Td /F1 9.8 Tf [(=0.014;)] TJ ET BT 553.510 390.382 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 378.477 Td /F1 9.8 Tf [(=3.131, )] TJ ET BT 61.760 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 378.477 Td /F1 9.8 Tf [(=0.006 for direct \(7/7 neurons\) and indirect pathway \(11/11 neurons\) MSNs in WTs, respectively; )] TJ ET BT 486.869 378.477 Td /F5 9.8 Tf [(t)] TJ ET BT 489.580 378.477 Td /F1 9.8 Tf [(=4.601, )] TJ ET BT 525.089 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 530.510 378.477 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 563.309 378.477 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 366.573 Td /F1 9.8 Tf [(=5.153, )] TJ ET BT 61.760 366.573 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 366.573 Td /F1 9.8 Tf [(<0.001 for direct \(9/9 neurons\) and indirect pathway \(7/7 neurons\) MSNs in YAC128 mice, respectively\) \(Fig. 2C\). )] TJ ET BT 26.250 354.668 Td /F1 9.8 Tf [(Response area increased significantly in direct pathway MSNs in only YAC128 mice \()] TJ ET BT 394.712 354.668 Td /F5 9.8 Tf [(t)] TJ ET BT 397.423 354.668 Td /F1 9.8 Tf [(=3.143, )] TJ ET BT 432.932 354.668 Td /F5 9.8 Tf [(p)] TJ ET BT 438.353 354.668 Td /F1 9.8 Tf [(=0.007; 9/9 neurons\) \(Fig. 2D\). )] TJ ET BT 26.250 342.763 Td /F1 9.8 Tf [(In contrast, in indirect pathway MSNs response area increased significantly in both WTs and YAC128 mice \()] TJ ET BT 492.261 342.763 Td /F5 9.8 Tf [(t)] TJ ET BT 494.972 342.763 Td /F1 9.8 Tf [(=3.371, )] TJ ET BT 530.481 342.763 Td /F5 9.8 Tf [(p)] TJ ET BT 535.902 342.763 Td /F1 9.8 Tf [(<0.004 )] TJ ET BT 26.250 330.858 Td /F1 9.8 Tf [(for WTs, 11/11 neurons; )] TJ ET BT 133.549 330.858 Td /F5 9.8 Tf [(t)] TJ ET BT 136.259 330.858 Td /F1 9.8 Tf [(=3.352, )] TJ ET BT 171.769 330.858 Td /F5 9.8 Tf [(p)] TJ ET BT 177.190 330.858 Td /F1 9.8 Tf [(=0.004 for YAC128, 7/7 neurons\).)] TJ ET BT 26.250 311.454 Td /F1 9.8 Tf [(Because DL-TBOA affected direct and indirect pathway MSN peak values in opposing ways, the area was normalized by peak )] TJ ET BT 26.250 299.549 Td /F1 9.8 Tf [(amplitude \(\(pA*ms\)/pA\) to verify a difference in the density of extrasynaptic NMDARs between direct and indirect pathway MSNs)] TJ ET 0.267 0.267 0.267 rg BT 26.250 289.151 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 31.069 287.644 Td /F1 9.8 Tf [(. In this way the relative area could be compared without the confounding influence of the peak amplitude. Normalized )] TJ ET BT 26.250 275.739 Td /F1 9.8 Tf [(response area increased significantly in all groups \()] TJ ET BT 247.877 275.739 Td /F5 9.8 Tf [(t)] TJ ET BT 250.588 275.739 Td /F1 9.8 Tf [(=5.090, )] TJ ET BT 286.097 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 291.518 275.739 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 324.317 275.739 Td /F5 9.8 Tf [( t)] TJ ET BT 329.738 275.739 Td /F1 9.8 Tf [(=2.64, )] TJ ET BT 359.827 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 365.248 275.739 Td /F1 9.8 Tf [(=0.017 for direct and indirect pathway MSNs in )] TJ ET BT 26.250 263.835 Td /F1 9.8 Tf [(WTs, respectively; )] TJ ET BT 108.599 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 111.309 263.835 Td /F1 9.8 Tf [(=4.125, )] TJ ET BT 146.819 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 152.240 263.835 Td /F1 9.8 Tf [(=0.001; )] TJ ET BT 187.749 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 190.460 263.835 Td /F1 9.8 Tf [(=6.622, )] TJ ET BT 225.969 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 231.390 263.835 Td /F1 9.8 Tf [(<0.001 for direct and indirect pathway MSNs in YAC128, respectively\) \(Fig. 2E\). )] TJ ET BT 26.250 251.930 Td /F1 9.8 Tf [(In addition, the normalized area increase was significantly greater for indirect pathway MSNs in YAC128 mice compared to WTs )] TJ ET BT 26.250 240.025 Td /F1 9.8 Tf [(\()] TJ ET BT 29.497 240.025 Td /F5 9.8 Tf [(t)] TJ ET BT 32.207 240.025 Td /F1 9.8 Tf [(=3.649, )] TJ ET BT 67.717 240.025 Td /F5 9.8 Tf [(p)] TJ ET BT 73.138 240.025 Td /F1 9.8 Tf [(=0.001\). These findings indicate that both response amplitude and decay are affected by TBOA but in slightly )] TJ ET BT 26.250 228.120 Td /F1 9.8 Tf [(different ways in direct and indirect pathway MSNs in YAC128 mice.)] TJ ET 0.965 0.965 0.965 rg 26.250 -201.011 555.000 419.250 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 218.239 m 581.250 218.239 l 581.250 217.489 l 26.250 217.489 l f q 450.000 0 0 403.500 35.250 -195.011 cm /I12 Do Q q 35.250 -201.011 537.000 0.000 re W n Q Q q 15.000 -201.011 577.500 978.011 re W n 0.965 0.965 0.965 rg 26.250 632.287 555.000 144.713 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 632.287 m 581.250 632.287 l 581.250 633.037 l 26.250 633.037 l f q 35.250 643.537 537.000 133.463 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 1: AMPAR- and NMDAR-mediated currents were significantly reduced in direct and indirect pathway MSNs in )] TJ ET BT 35.250 755.571 Td /F4 9.8 Tf [(YAC128 mice.)] TJ ET BT 35.250 736.201 Td /F4 9.8 Tf [(A)] TJ ET BT 42.290 736.201 Td /F1 9.8 Tf [(: Typical traces of AMPAR- mediated currents in WT and YAC128 direct and indirect pathway MSNs evoked at the 0.10 )] TJ ET BT 35.250 722.465 Td /F1 9.8 Tf [(mA stimulus intensity are shown. )] TJ ET BT 179.921 722.465 Td /F4 9.8 Tf [(B)] TJ ET BT 186.960 722.465 Td /F1 9.8 Tf [(: Input-output functions of evoked AMPAR-mediated currents were significantly )] TJ ET BT 35.250 708.729 Td /F1 9.8 Tf [(decreased in both direct and indirect pathway MSNs of YAC128 mice. )] TJ ET BT 339.255 708.729 Td /F4 9.8 Tf [(C)] TJ ET BT 346.294 708.729 Td /F1 9.8 Tf [(: Typical traces of NMDAR-mediated currents in )] TJ ET BT 35.250 694.993 Td /F1 9.8 Tf [(WT and YAC128 direct and indirect pathway MSNs evoked at 0.10 mA stimulus intensity are shown. )] TJ ET BT 470.383 694.993 Td /F4 9.8 Tf [(D)] TJ ET BT 477.422 694.993 Td /F1 9.8 Tf [(: Input-output )] TJ ET BT 35.250 681.256 Td /F1 9.8 Tf [(functions of evoked NMDAR-mediated currents also were significantly reduced in both direct and indirect pathway MSNs )] TJ ET BT 35.250 667.520 Td /F1 9.8 Tf [(from YAC128 mice. In this and other figures * indicates p<0.05, ** indicates p<0.01, *** indicates p<0.001. See text for )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(details of statistics.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(There were no significant differences in peak AMPAR- or NMDAR-mediated currents between direct or indirect pathway MSNs )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(from either WT or YAC128 mice. Furthermore, there were no differences in NMDA/AMPA ratios among direct and indirect )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(pathway MSNs in WT and YAC128 mice \(0.510.074 vs. 0.440.08 for WTs and YAC128 from direct pathway MSNs, )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(respectively, p=0.542; 0.450.047 vs. 0.560.056 for WTs and YAC128 from indirect pathway MSNs, respectively, p=0.135\).)] TJ ET BT 26.250 560.144 Td /F4 9.8 Tf [(Glutamate transport block revealed alterations in extrasynaptic NMDAR-mediated responses in YAC128 MSNs.)] TJ ET BT 26.250 540.739 Td /F1 9.8 Tf [(We used DL-TBOA \(30 ?M\), a glutamate transport blocker which promotes activation of extrasynaptic NMDARs through )] TJ ET BT 26.250 528.834 Td /F1 9.8 Tf [(glutamate spillover, to assess differences in evoked NMDAR-mediated currents in direct and indirect pathway MSNs. We )] TJ ET BT 26.250 516.930 Td /F1 9.8 Tf [(examined alterations in four indices, peak amplitude, decay time, response area and response area normalized by peak )] TJ ET BT 26.250 505.025 Td /F1 9.8 Tf [(response separately for direct or indirect pathway MSNs \(Fig.2\). An increase in the contribution from extrasynaptic NMDA )] TJ ET BT 26.250 493.120 Td /F1 9.8 Tf [(receptors would be shown by changes in peak amplitude, decay time and response area of the currents. We only tested the )] TJ ET BT 26.250 481.215 Td /F1 9.8 Tf [(effects of TBOA at the stimulus intensity that produced 75% of the maximum response for each neuron. In WTs for both direct )] TJ ET BT 26.250 469.311 Td /F1 9.8 Tf [(and indirect pathway MSNs, peak amplitude decreased significantly in TBOA \()] TJ ET BT 363.298 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 366.008 469.311 Td /F1 9.8 Tf [(=2.177, )] TJ ET BT 401.518 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 406.939 469.311 Td /F1 9.8 Tf [(=0.047; )] TJ ET BT 442.448 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 445.159 469.311 Td /F1 9.8 Tf [(=2.684, )] TJ ET BT 480.668 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 486.089 469.311 Td /F1 9.8 Tf [(=0.016 for direct and )] TJ ET BT 26.250 457.406 Td /F1 9.8 Tf [(indirect pathway MSNs, respectively\) \(Fig. 2B\). In both direct and indirect pathway MSNs, almost all neurons \(6/7 and 9/10 for )] TJ ET BT 26.250 445.501 Td /F1 9.8 Tf [(direct and indirect pathway MSNs, respectively\) showed a decrease in the peak response amplitude although for each group )] TJ ET BT 26.250 433.596 Td /F1 9.8 Tf [(there was considerable variation in the absolute amplitudes of the responses of the neurons. In contrast, in direct pathway )] TJ ET BT 26.250 421.692 Td /F1 9.8 Tf [(MSNs of YAC128 mice, peak amplitude increased significantly \()] TJ ET BT 301.512 421.692 Td /F5 9.8 Tf [(t)] TJ ET BT 304.223 421.692 Td /F1 9.8 Tf [(=4.128, )] TJ ET BT 339.732 421.692 Td /F5 9.8 Tf [(p)] TJ ET BT 345.153 421.692 Td /F1 9.8 Tf [(=0.001; 8/9 neurons\) while in indirect pathway MSNs )] TJ ET BT 26.250 409.787 Td /F1 9.8 Tf [(peak amplitude decreased significantly \()] TJ ET BT 199.654 409.787 Td /F5 9.8 Tf [(t)] TJ ET BT 202.364 409.787 Td /F1 9.8 Tf [(=6.017, )] TJ ET BT 237.874 409.787 Td /F5 9.8 Tf [(p)] TJ ET BT 243.295 409.787 Td /F1 9.8 Tf [(<0.001; 7/7 neurons\) \(Fig. 2B\).)] TJ ET BT 26.250 390.382 Td /F1 9.8 Tf [(Decay time increased significantly for both direct and indirect pathway MSNs in WTs and YAC128 mice \()] TJ ET BT 477.070 390.382 Td /F5 9.8 Tf [(t)] TJ ET BT 479.781 390.382 Td /F1 9.8 Tf [(=2.817, )] TJ ET BT 515.291 390.382 Td /F5 9.8 Tf [(p)] TJ ET BT 520.712 390.382 Td /F1 9.8 Tf [(=0.014;)] TJ ET BT 553.510 390.382 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 378.477 Td /F1 9.8 Tf [(=3.131, )] TJ ET BT 61.760 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 378.477 Td /F1 9.8 Tf [(=0.006 for direct \(7/7 neurons\) and indirect pathway \(11/11 neurons\) MSNs in WTs, respectively; )] TJ ET BT 486.869 378.477 Td /F5 9.8 Tf [(t)] TJ ET BT 489.580 378.477 Td /F1 9.8 Tf [(=4.601, )] TJ ET BT 525.089 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 530.510 378.477 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 563.309 378.477 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 366.573 Td /F1 9.8 Tf [(=5.153, )] TJ ET BT 61.760 366.573 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 366.573 Td /F1 9.8 Tf [(<0.001 for direct \(9/9 neurons\) and indirect pathway \(7/7 neurons\) MSNs in YAC128 mice, respectively\) \(Fig. 2C\). )] TJ ET BT 26.250 354.668 Td /F1 9.8 Tf [(Response area increased significantly in direct pathway MSNs in only YAC128 mice \()] TJ ET BT 394.712 354.668 Td /F5 9.8 Tf [(t)] TJ ET BT 397.423 354.668 Td /F1 9.8 Tf [(=3.143, )] TJ ET BT 432.932 354.668 Td /F5 9.8 Tf [(p)] TJ ET BT 438.353 354.668 Td /F1 9.8 Tf [(=0.007; 9/9 neurons\) \(Fig. 2D\). )] TJ ET BT 26.250 342.763 Td /F1 9.8 Tf [(In contrast, in indirect pathway MSNs response area increased significantly in both WTs and YAC128 mice \()] TJ ET BT 492.261 342.763 Td /F5 9.8 Tf [(t)] TJ ET BT 494.972 342.763 Td /F1 9.8 Tf [(=3.371, )] TJ ET BT 530.481 342.763 Td /F5 9.8 Tf [(p)] TJ ET BT 535.902 342.763 Td /F1 9.8 Tf [(<0.004 )] TJ ET BT 26.250 330.858 Td /F1 9.8 Tf [(for WTs, 11/11 neurons; )] TJ ET BT 133.549 330.858 Td /F5 9.8 Tf [(t)] TJ ET BT 136.259 330.858 Td /F1 9.8 Tf [(=3.352, )] TJ ET BT 171.769 330.858 Td /F5 9.8 Tf [(p)] TJ ET BT 177.190 330.858 Td /F1 9.8 Tf [(=0.004 for YAC128, 7/7 neurons\).)] TJ ET BT 26.250 311.454 Td /F1 9.8 Tf [(Because DL-TBOA affected direct and indirect pathway MSN peak values in opposing ways, the area was normalized by peak )] TJ ET BT 26.250 299.549 Td /F1 9.8 Tf [(amplitude \(\(pA*ms\)/pA\) to verify a difference in the density of extrasynaptic NMDARs between direct and indirect pathway MSNs)] TJ ET 0.267 0.267 0.267 rg BT 26.250 289.151 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 31.069 287.644 Td /F1 9.8 Tf [(. In this way the relative area could be compared without the confounding influence of the peak amplitude. Normalized )] TJ ET BT 26.250 275.739 Td /F1 9.8 Tf [(response area increased significantly in all groups \()] TJ ET BT 247.877 275.739 Td /F5 9.8 Tf [(t)] TJ ET BT 250.588 275.739 Td /F1 9.8 Tf [(=5.090, )] TJ ET BT 286.097 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 291.518 275.739 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 324.317 275.739 Td /F5 9.8 Tf [( t)] TJ ET BT 329.738 275.739 Td /F1 9.8 Tf [(=2.64, )] TJ ET BT 359.827 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 365.248 275.739 Td /F1 9.8 Tf [(=0.017 for direct and indirect pathway MSNs in )] TJ ET BT 26.250 263.835 Td /F1 9.8 Tf [(WTs, respectively; )] TJ ET BT 108.599 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 111.309 263.835 Td /F1 9.8 Tf [(=4.125, )] TJ ET BT 146.819 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 152.240 263.835 Td /F1 9.8 Tf [(=0.001; )] TJ ET BT 187.749 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 190.460 263.835 Td /F1 9.8 Tf [(=6.622, )] TJ ET BT 225.969 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 231.390 263.835 Td /F1 9.8 Tf [(<0.001 for direct and indirect pathway MSNs in YAC128, respectively\) \(Fig. 2E\). )] TJ ET BT 26.250 251.930 Td /F1 9.8 Tf [(In addition, the normalized area increase was significantly greater for indirect pathway MSNs in YAC128 mice compared to WTs )] TJ ET BT 26.250 240.025 Td /F1 9.8 Tf [(\()] TJ ET BT 29.497 240.025 Td /F5 9.8 Tf [(t)] TJ ET BT 32.207 240.025 Td /F1 9.8 Tf [(=3.649, )] TJ ET BT 67.717 240.025 Td /F5 9.8 Tf [(p)] TJ ET BT 73.138 240.025 Td /F1 9.8 Tf [(=0.001\). These findings indicate that both response amplitude and decay are affected by TBOA but in slightly )] TJ ET BT 26.250 228.120 Td /F1 9.8 Tf [(different ways in direct and indirect pathway MSNs in YAC128 mice.)] TJ ET 0.965 0.965 0.965 rg 26.250 -201.011 555.000 419.250 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 218.239 m 581.250 218.239 l 581.250 217.489 l 26.250 217.489 l f q 450.000 0 0 403.500 35.250 -195.011 cm /I14 Do Q q 35.250 -201.011 537.000 0.000 re W n Q Q q 15.000 -201.011 577.500 978.011 re W n 0.965 0.965 0.965 rg 26.250 632.287 555.000 144.713 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 632.287 m 581.250 632.287 l 581.250 633.037 l 26.250 633.037 l f q 35.250 643.537 537.000 133.463 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 1: AMPAR- and NMDAR-mediated currents were significantly reduced in direct and indirect pathway MSNs in )] TJ ET BT 35.250 755.571 Td /F4 9.8 Tf [(YAC128 mice.)] TJ ET BT 35.250 736.201 Td /F4 9.8 Tf [(A)] TJ ET BT 42.290 736.201 Td /F1 9.8 Tf [(: Typical traces of AMPAR- mediated currents in WT and YAC128 direct and indirect pathway MSNs evoked at the 0.10 )] TJ ET BT 35.250 722.465 Td /F1 9.8 Tf [(mA stimulus intensity are shown. )] TJ ET BT 179.921 722.465 Td /F4 9.8 Tf [(B)] TJ ET BT 186.960 722.465 Td /F1 9.8 Tf [(: Input-output functions of evoked AMPAR-mediated currents were significantly )] TJ ET BT 35.250 708.729 Td /F1 9.8 Tf [(decreased in both direct and indirect pathway MSNs of YAC128 mice. )] TJ ET BT 339.255 708.729 Td /F4 9.8 Tf [(C)] TJ ET BT 346.294 708.729 Td /F1 9.8 Tf [(: Typical traces of NMDAR-mediated currents in )] TJ ET BT 35.250 694.993 Td /F1 9.8 Tf [(WT and YAC128 direct and indirect pathway MSNs evoked at 0.10 mA stimulus intensity are shown. )] TJ ET BT 470.383 694.993 Td /F4 9.8 Tf [(D)] TJ ET BT 477.422 694.993 Td /F1 9.8 Tf [(: Input-output )] TJ ET BT 35.250 681.256 Td /F1 9.8 Tf [(functions of evoked NMDAR-mediated currents also were significantly reduced in both direct and indirect pathway MSNs )] TJ ET BT 35.250 667.520 Td /F1 9.8 Tf [(from YAC128 mice. In this and other figures * indicates p<0.05, ** indicates p<0.01, *** indicates p<0.001. See text for )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(details of statistics.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(There were no significant differences in peak AMPAR- or NMDAR-mediated currents between direct or indirect pathway MSNs )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(from either WT or YAC128 mice. Furthermore, there were no differences in NMDA/AMPA ratios among direct and indirect )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(pathway MSNs in WT and YAC128 mice \(0.510.074 vs. 0.440.08 for WTs and YAC128 from direct pathway MSNs, )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(respectively, p=0.542; 0.450.047 vs. 0.560.056 for WTs and YAC128 from indirect pathway MSNs, respectively, p=0.135\).)] TJ ET BT 26.250 560.144 Td /F4 9.8 Tf [(Glutamate transport block revealed alterations in extrasynaptic NMDAR-mediated responses in YAC128 MSNs.)] TJ ET BT 26.250 540.739 Td /F1 9.8 Tf [(We used DL-TBOA \(30 ?M\), a glutamate transport blocker which promotes activation of extrasynaptic NMDARs through )] TJ ET BT 26.250 528.834 Td /F1 9.8 Tf [(glutamate spillover, to assess differences in evoked NMDAR-mediated currents in direct and indirect pathway MSNs. We )] TJ ET BT 26.250 516.930 Td /F1 9.8 Tf [(examined alterations in four indices, peak amplitude, decay time, response area and response area normalized by peak )] TJ ET BT 26.250 505.025 Td /F1 9.8 Tf [(response separately for direct or indirect pathway MSNs \(Fig.2\). An increase in the contribution from extrasynaptic NMDA )] TJ ET BT 26.250 493.120 Td /F1 9.8 Tf [(receptors would be shown by changes in peak amplitude, decay time and response area of the currents. We only tested the )] TJ ET BT 26.250 481.215 Td /F1 9.8 Tf [(effects of TBOA at the stimulus intensity that produced 75% of the maximum response for each neuron. In WTs for both direct )] TJ ET BT 26.250 469.311 Td /F1 9.8 Tf [(and indirect pathway MSNs, peak amplitude decreased significantly in TBOA \()] TJ ET BT 363.298 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 366.008 469.311 Td /F1 9.8 Tf [(=2.177, )] TJ ET BT 401.518 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 406.939 469.311 Td /F1 9.8 Tf [(=0.047; )] TJ ET BT 442.448 469.311 Td /F5 9.8 Tf [(t)] TJ ET BT 445.159 469.311 Td /F1 9.8 Tf [(=2.684, )] TJ ET BT 480.668 469.311 Td /F5 9.8 Tf [(p)] TJ ET BT 486.089 469.311 Td /F1 9.8 Tf [(=0.016 for direct and )] TJ ET BT 26.250 457.406 Td /F1 9.8 Tf [(indirect pathway MSNs, respectively\) \(Fig. 2B\). In both direct and indirect pathway MSNs, almost all neurons \(6/7 and 9/10 for )] TJ ET BT 26.250 445.501 Td /F1 9.8 Tf [(direct and indirect pathway MSNs, respectively\) showed a decrease in the peak response amplitude although for each group )] TJ ET BT 26.250 433.596 Td /F1 9.8 Tf [(there was considerable variation in the absolute amplitudes of the responses of the neurons. In contrast, in direct pathway )] TJ ET BT 26.250 421.692 Td /F1 9.8 Tf [(MSNs of YAC128 mice, peak amplitude increased significantly \()] TJ ET BT 301.512 421.692 Td /F5 9.8 Tf [(t)] TJ ET BT 304.223 421.692 Td /F1 9.8 Tf [(=4.128, )] TJ ET BT 339.732 421.692 Td /F5 9.8 Tf [(p)] TJ ET BT 345.153 421.692 Td /F1 9.8 Tf [(=0.001; 8/9 neurons\) while in indirect pathway MSNs )] TJ ET BT 26.250 409.787 Td /F1 9.8 Tf [(peak amplitude decreased significantly \()] TJ ET BT 199.654 409.787 Td /F5 9.8 Tf [(t)] TJ ET BT 202.364 409.787 Td /F1 9.8 Tf [(=6.017, )] TJ ET BT 237.874 409.787 Td /F5 9.8 Tf [(p)] TJ ET BT 243.295 409.787 Td /F1 9.8 Tf [(<0.001; 7/7 neurons\) \(Fig. 2B\).)] TJ ET BT 26.250 390.382 Td /F1 9.8 Tf [(Decay time increased significantly for both direct and indirect pathway MSNs in WTs and YAC128 mice \()] TJ ET BT 477.070 390.382 Td /F5 9.8 Tf [(t)] TJ ET BT 479.781 390.382 Td /F1 9.8 Tf [(=2.817, )] TJ ET BT 515.291 390.382 Td /F5 9.8 Tf [(p)] TJ ET BT 520.712 390.382 Td /F1 9.8 Tf [(=0.014;)] TJ ET BT 553.510 390.382 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 378.477 Td /F1 9.8 Tf [(=3.131, )] TJ ET BT 61.760 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 378.477 Td /F1 9.8 Tf [(=0.006 for direct \(7/7 neurons\) and indirect pathway \(11/11 neurons\) MSNs in WTs, respectively; )] TJ ET BT 486.869 378.477 Td /F5 9.8 Tf [(t)] TJ ET BT 489.580 378.477 Td /F1 9.8 Tf [(=4.601, )] TJ ET BT 525.089 378.477 Td /F5 9.8 Tf [(p)] TJ ET BT 530.510 378.477 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 563.309 378.477 Td /F5 9.8 Tf [( t)] TJ ET BT 26.250 366.573 Td /F1 9.8 Tf [(=5.153, )] TJ ET BT 61.760 366.573 Td /F5 9.8 Tf [(p)] TJ ET BT 67.180 366.573 Td /F1 9.8 Tf [(<0.001 for direct \(9/9 neurons\) and indirect pathway \(7/7 neurons\) MSNs in YAC128 mice, respectively\) \(Fig. 2C\). )] TJ ET BT 26.250 354.668 Td /F1 9.8 Tf [(Response area increased significantly in direct pathway MSNs in only YAC128 mice \()] TJ ET BT 394.712 354.668 Td /F5 9.8 Tf [(t)] TJ ET BT 397.423 354.668 Td /F1 9.8 Tf [(=3.143, )] TJ ET BT 432.932 354.668 Td /F5 9.8 Tf [(p)] TJ ET BT 438.353 354.668 Td /F1 9.8 Tf [(=0.007; 9/9 neurons\) \(Fig. 2D\). )] TJ ET BT 26.250 342.763 Td /F1 9.8 Tf [(In contrast, in indirect pathway MSNs response area increased significantly in both WTs and YAC128 mice \()] TJ ET BT 492.261 342.763 Td /F5 9.8 Tf [(t)] TJ ET BT 494.972 342.763 Td /F1 9.8 Tf [(=3.371, )] TJ ET BT 530.481 342.763 Td /F5 9.8 Tf [(p)] TJ ET BT 535.902 342.763 Td /F1 9.8 Tf [(<0.004 )] TJ ET BT 26.250 330.858 Td /F1 9.8 Tf [(for WTs, 11/11 neurons; )] TJ ET BT 133.549 330.858 Td /F5 9.8 Tf [(t)] TJ ET BT 136.259 330.858 Td /F1 9.8 Tf [(=3.352, )] TJ ET BT 171.769 330.858 Td /F5 9.8 Tf [(p)] TJ ET BT 177.190 330.858 Td /F1 9.8 Tf [(=0.004 for YAC128, 7/7 neurons\).)] TJ ET BT 26.250 311.454 Td /F1 9.8 Tf [(Because DL-TBOA affected direct and indirect pathway MSN peak values in opposing ways, the area was normalized by peak )] TJ ET BT 26.250 299.549 Td /F1 9.8 Tf [(amplitude \(\(pA*ms\)/pA\) to verify a difference in the density of extrasynaptic NMDARs between direct and indirect pathway MSNs)] TJ ET 0.267 0.267 0.267 rg BT 26.250 289.151 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 31.069 287.644 Td /F1 9.8 Tf [(. In this way the relative area could be compared without the confounding influence of the peak amplitude. Normalized )] TJ ET BT 26.250 275.739 Td /F1 9.8 Tf [(response area increased significantly in all groups \()] TJ ET BT 247.877 275.739 Td /F5 9.8 Tf [(t)] TJ ET BT 250.588 275.739 Td /F1 9.8 Tf [(=5.090, )] TJ ET BT 286.097 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 291.518 275.739 Td /F1 9.8 Tf [(<0.001;)] TJ ET BT 324.317 275.739 Td /F5 9.8 Tf [( t)] TJ ET BT 329.738 275.739 Td /F1 9.8 Tf [(=2.64, )] TJ ET BT 359.827 275.739 Td /F5 9.8 Tf [(p)] TJ ET BT 365.248 275.739 Td /F1 9.8 Tf [(=0.017 for direct and indirect pathway MSNs in )] TJ ET BT 26.250 263.835 Td /F1 9.8 Tf [(WTs, respectively; )] TJ ET BT 108.599 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 111.309 263.835 Td /F1 9.8 Tf [(=4.125, )] TJ ET BT 146.819 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 152.240 263.835 Td /F1 9.8 Tf [(=0.001; )] TJ ET BT 187.749 263.835 Td /F5 9.8 Tf [(t)] TJ ET BT 190.460 263.835 Td /F1 9.8 Tf [(=6.622, )] TJ ET BT 225.969 263.835 Td /F5 9.8 Tf [(p)] TJ ET BT 231.390 263.835 Td /F1 9.8 Tf [(<0.001 for direct and indirect pathway MSNs in YAC128, respectively\) \(Fig. 2E\). )] TJ ET BT 26.250 251.930 Td /F1 9.8 Tf [(In addition, the normalized area increase was significantly greater for indirect pathway MSNs in YAC128 mice compared to WTs )] TJ ET BT 26.250 240.025 Td /F1 9.8 Tf [(\()] TJ ET BT 29.497 240.025 Td /F5 9.8 Tf [(t)] TJ ET BT 32.207 240.025 Td /F1 9.8 Tf [(=3.649, )] TJ ET BT 67.717 240.025 Td /F5 9.8 Tf [(p)] TJ ET BT 73.138 240.025 Td /F1 9.8 Tf [(=0.001\). 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B: Peak amplitude is differentially )] TJ ET BT 35.250 708.729 Td /F1 9.8 Tf [(altered in direct and indirect pathway MSNs by DL-TBOA. Peak amplitude increases in direct pathway MSNs but decreases )] TJ ET BT 35.250 694.993 Td /F1 9.8 Tf [(in indirect pathway MSNs in YAC128 mice. C: In DL-TBOA response decay time increases in direct and indirect pathway )] TJ ET BT 35.250 681.256 Td /F1 9.8 Tf [(MSNs in both WT and YAC128 mice. D: In DL-TBOA, response area increases only in direct pathway MSNs in YAC128 )] TJ ET BT 35.250 667.520 Td /F1 9.8 Tf [(mice and in indirect pathway MSNs in both WT and YAC128 mice. E: In DL-TBOA, normalized area increases in all groups )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(but displays the greatest increase in indirect pathway MSNs in YAC128 mice.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(To more specifically compare the effects of TBOA in direct and indirect pathway MSNs, changes induced by TBOA were )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(converted to percentages \(Fig. 3\). In direct pathway MSNs, TBOA decreased the evoked NMDAR peak current in WTs but )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(increased it in YAC128 mice \()] TJ ET BT 154.667 591.453 Td /F5 9.8 Tf [(t)] TJ ET BT 157.378 591.453 Td /F1 9.8 Tf [(=6.449,)] TJ ET BT 190.177 591.453 Td /F5 9.8 Tf [( p)] TJ ET BT 198.308 591.453 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3A\). In indirect pathway MSNs TBOA decreased the peak in both WT and )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(YAC128 mice, but the reduction was significantly larger in YAC128 D2 MSNs \()] TJ ET BT 364.370 579.549 Td /F5 9.8 Tf [(t)] TJ ET BT 367.081 579.549 Td /F1 9.8 Tf [(=3.323, )] TJ ET BT 402.590 579.549 Td /F5 9.8 Tf [(p)] TJ ET BT 408.011 579.549 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3A\). The difference in )] TJ ET BT 26.250 567.644 Td /F1 9.8 Tf [(peak amplitude between direct and indirect pathway MSNs in YAC128 mice also was significant \()] TJ ET BT 445.120 567.644 Td /F5 9.8 Tf [(t)] TJ ET BT 447.830 567.644 Td /F1 9.8 Tf [(=9.150, )] TJ ET BT 483.340 567.644 Td /F5 9.8 Tf [(p)] TJ ET BT 488.761 567.644 Td /F1 9.8 Tf [(<0.001\). Decay time )] TJ ET BT 26.250 555.739 Td /F1 9.8 Tf [(was significantly elevated in indirect YAC128 MSNs compared to WTs \()] TJ ET BT 334.555 555.739 Td /F5 9.8 Tf [(t)] TJ ET BT 337.265 555.739 Td /F1 9.8 Tf [(=3.601, )] TJ ET BT 372.775 555.739 Td /F5 9.8 Tf [(p)] TJ ET BT 378.196 555.739 Td /F1 9.8 Tf [(=0.001\) \(Fig. 3B\) and also was significantly )] TJ ET BT 26.250 543.834 Td /F1 9.8 Tf [(greater in indirect pathway MSNs compared to direct pathway MSNs in YAC128 mice \()] TJ ET BT 400.123 543.834 Td /F5 9.8 Tf [(t)] TJ ET BT 402.834 543.834 Td /F1 9.8 Tf [(=2.258, )] TJ ET BT 438.344 543.834 Td /F5 9.8 Tf [(p)] TJ ET BT 443.764 543.834 Td /F1 9.8 Tf [(=0.031\). Area was significantly )] TJ ET BT 26.250 531.930 Td /F1 9.8 Tf [(larger in YAC128 direct pathway MSNs compared to area of WTs \()] TJ ET BT 313.973 531.930 Td /F5 9.8 Tf [(t)] TJ ET BT 316.683 531.930 Td /F1 9.8 Tf [(=3.482, )] TJ ET BT 352.193 531.930 Td /F5 9.8 Tf [(p)] TJ ET BT 357.614 531.930 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3C\). The increased area of indirect )] TJ ET BT 26.250 520.025 Td /F1 9.8 Tf [(pathway MSNs approached statistical significance \()] TJ ET BT 248.413 520.025 Td /F5 9.8 Tf [(t)] TJ ET BT 251.124 520.025 Td /F1 9.8 Tf [(=1.829, )] TJ ET BT 286.634 520.025 Td /F5 9.8 Tf [(p)] TJ ET BT 292.055 520.025 Td /F1 9.8 Tf [(=0.077\). The increased area of direct versus indirect pathway )] TJ ET BT 26.250 508.120 Td /F1 9.8 Tf [(MSNs was significantly greater in only the YAC128 mice \()] TJ ET BT 274.953 508.120 Td /F5 9.8 Tf [(t)] TJ ET BT 277.663 508.120 Td /F1 9.8 Tf [(=2.353, )] TJ ET BT 313.173 508.120 Td /F5 9.8 Tf [(p)] TJ ET BT 318.594 508.120 Td /F1 9.8 Tf [(<0.025\). There was a significant increase in relative area in )] TJ ET BT 26.250 496.215 Td /F1 9.8 Tf [(indirect pathway MSNs in YAC128 mice compared to WTs \()] TJ ET BT 284.157 496.215 Td /F5 9.8 Tf [(t)] TJ ET BT 286.868 496.215 Td /F1 9.8 Tf [(=5.012, )] TJ ET BT 322.377 496.215 Td /F5 9.8 Tf [(p)] TJ ET BT 327.798 496.215 Td /F1 9.8 Tf [(<0.001\) as well as a significant increase in indirect )] TJ ET BT 26.250 484.311 Td /F1 9.8 Tf [(pathway MSNs compared to direct pathway MSNs but only in MSNs in YAC128 mice \()] TJ ET BT 398.495 484.311 Td /F5 9.8 Tf [(t)] TJ ET BT 401.206 484.311 Td /F1 9.8 Tf [(=4.158, )] TJ ET BT 436.715 484.311 Td /F5 9.8 Tf [(p)] TJ ET BT 442.136 484.311 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 238.080 555.000 236.350 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 474.430 m 581.250 474.430 l 581.250 473.680 l 26.250 473.680 l f 26.250 238.080 m 581.250 238.080 l 581.250 238.830 l 26.250 238.830 l f q 450.000 0 0 129.000 35.250 335.680 cm /I20 Do Q q 35.250 249.330 537.000 80.350 re W n 0.271 0.267 0.267 rg BT 35.250 320.156 Td /F4 9.8 Tf [(Fig. 3: Differential effects of DL-TBOA on direct and indirect pathway MSNs exemplified by percentage changes.)] TJ ET BT 35.250 300.786 Td /F1 9.8 Tf [(A: Percentage changes in peak response amplitude were differentially altered in direct and indirect pathway MSNs of )] TJ ET BT 35.250 287.050 Td /F1 9.8 Tf [(YAC128 mice. B: Percentage changes in decay times were greater in indirect pathway MSNs from YAC128 mice. C: )] TJ ET BT 35.250 273.313 Td /F1 9.8 Tf [(Percentage change in response area were greatest in direct pathway MSNs of YAC128 mice. D: Normalized area increases )] TJ ET BT 35.250 259.577 Td /F1 9.8 Tf [(were greatest in indirect pathway MSNs of YAC128 mice.)] TJ ET Q BT 26.250 203.858 Td /F4 12.0 Tf [(Discussion)] TJ ET BT 26.250 183.904 Td /F1 9.8 Tf [(The present study demonstrated decreases in evoked excitatory postsynaptic currents mediated by AMPARs and NMDARs in )] TJ ET BT 26.250 171.999 Td /F1 9.8 Tf [(YAC128 direct and indirect pathway MSNs. Furthermore, in contrast to the downregulation of synaptic activity, NMDAR-)] TJ ET BT 26.250 160.095 Td /F1 9.8 Tf [(mediated responses evoked in the presence of TBOA were increased in both direct and indirect pathway MSNs of YAC128 )] TJ ET BT 26.250 148.190 Td /F1 9.8 Tf [(mice, but in slightly different ways. In direct pathway MSNs, response increases were mainly due to higher amplitudes while in )] TJ ET BT 26.250 136.285 Td /F1 9.8 Tf [(indirect pathway MSNs, the increase was due to longer decay times. This outcome indicates differential changes in synaptic )] TJ ET BT 26.250 124.380 Td /F1 9.8 Tf [(versus extrasynaptic NMDA receptors in both direct and indirect pathway MSNs from these older, symptomatic YAC128 mice.)] TJ ET BT 26.250 104.976 Td /F1 9.8 Tf [(We previously reported a progressive disconnection of striatal MSNs from their glutamatergic inputs in various models of HD)] TJ ET 0.267 0.267 0.267 rg BT 561.593 106.483 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 566.412 108.864 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 568.821 106.483 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 573.640 104.976 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 93.071 Td /F1 9.8 Tf [(In 12 month-old YAC128 mice we also showed that the reduction in the spontaneous excitatory postsynaptic current frequency )] TJ ET BT 26.250 81.166 Td /F1 9.8 Tf [(was greater in direct than indirect pathway MSNs but we did not examine AMPAR and NMDAR responses separately and our )] TJ ET BT 26.250 69.261 Td /F1 9.8 Tf [(results were based on AMPAR-mediated responses)] TJ ET 0.267 0.267 0.267 rg BT 250.042 70.769 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 254.860 69.261 Td /F1 9.8 Tf [(. The reduction in evoked synaptic responses agrees with previous studies )] TJ ET BT 26.250 57.357 Td /F1 9.8 Tf [(showing decreases in glutamate release as well as AMPAR- and NMDAR-mediated currents in symptomatic YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 555.080 58.864 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 559.899 57.357 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 45.452 Td /F1 9.8 Tf [(The reduction in the amplitude of evoked responses indicates that there is a loss of glutamatergic inputs onto MSNs in YAC128 )] TJ ET Q q 15.000 31.166 577.500 745.834 re W n 0.965 0.965 0.965 rg 26.250 632.287 555.000 144.713 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 632.287 m 581.250 632.287 l 581.250 633.037 l 26.250 633.037 l f q 35.250 643.537 537.000 133.463 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 2: DL-TBOA alters peak amplitude, decay, area and relative area of direct and indirect pathway MSNs of WT )] TJ ET BT 35.250 755.571 Td /F4 9.8 Tf [(and YAC128 mice.)] TJ ET BT 35.250 736.201 Td /F1 9.8 Tf [(A: Typical traces of NMDAR-mediated currents in WT and YAC128 direct and indirect pathway MSNs evoked at 75% of the )] TJ ET BT 35.250 722.465 Td /F1 9.8 Tf [(maximal stimulus intensity showing alterations in peak amplitudes and decay times. B: Peak amplitude is differentially )] TJ ET BT 35.250 708.729 Td /F1 9.8 Tf [(altered in direct and indirect pathway MSNs by DL-TBOA. Peak amplitude increases in direct pathway MSNs but decreases )] TJ ET BT 35.250 694.993 Td /F1 9.8 Tf [(in indirect pathway MSNs in YAC128 mice. C: In DL-TBOA response decay time increases in direct and indirect pathway )] TJ ET BT 35.250 681.256 Td /F1 9.8 Tf [(MSNs in both WT and YAC128 mice. D: In DL-TBOA, response area increases only in direct pathway MSNs in YAC128 )] TJ ET BT 35.250 667.520 Td /F1 9.8 Tf [(mice and in indirect pathway MSNs in both WT and YAC128 mice. E: In DL-TBOA, normalized area increases in all groups )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(but displays the greatest increase in indirect pathway MSNs in YAC128 mice.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(To more specifically compare the effects of TBOA in direct and indirect pathway MSNs, changes induced by TBOA were )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(converted to percentages \(Fig. 3\). In direct pathway MSNs, TBOA decreased the evoked NMDAR peak current in WTs but )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(increased it in YAC128 mice \()] TJ ET BT 154.667 591.453 Td /F5 9.8 Tf [(t)] TJ ET BT 157.378 591.453 Td /F1 9.8 Tf [(=6.449,)] TJ ET BT 190.177 591.453 Td /F5 9.8 Tf [( p)] TJ ET BT 198.308 591.453 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3A\). In indirect pathway MSNs TBOA decreased the peak in both WT and )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(YAC128 mice, but the reduction was significantly larger in YAC128 D2 MSNs \()] TJ ET BT 364.370 579.549 Td /F5 9.8 Tf [(t)] TJ ET BT 367.081 579.549 Td /F1 9.8 Tf [(=3.323, )] TJ ET BT 402.590 579.549 Td /F5 9.8 Tf [(p)] TJ ET BT 408.011 579.549 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3A\). The difference in )] TJ ET BT 26.250 567.644 Td /F1 9.8 Tf [(peak amplitude between direct and indirect pathway MSNs in YAC128 mice also was significant \()] TJ ET BT 445.120 567.644 Td /F5 9.8 Tf [(t)] TJ ET BT 447.830 567.644 Td /F1 9.8 Tf [(=9.150, )] TJ ET BT 483.340 567.644 Td /F5 9.8 Tf [(p)] TJ ET BT 488.761 567.644 Td /F1 9.8 Tf [(<0.001\). Decay time )] TJ ET BT 26.250 555.739 Td /F1 9.8 Tf [(was significantly elevated in indirect YAC128 MSNs compared to WTs \()] TJ ET BT 334.555 555.739 Td /F5 9.8 Tf [(t)] TJ ET BT 337.265 555.739 Td /F1 9.8 Tf [(=3.601, )] TJ ET BT 372.775 555.739 Td /F5 9.8 Tf [(p)] TJ ET BT 378.196 555.739 Td /F1 9.8 Tf [(=0.001\) \(Fig. 3B\) and also was significantly )] TJ ET BT 26.250 543.834 Td /F1 9.8 Tf [(greater in indirect pathway MSNs compared to direct pathway MSNs in YAC128 mice \()] TJ ET BT 400.123 543.834 Td /F5 9.8 Tf [(t)] TJ ET BT 402.834 543.834 Td /F1 9.8 Tf [(=2.258, )] TJ ET BT 438.344 543.834 Td /F5 9.8 Tf [(p)] TJ ET BT 443.764 543.834 Td /F1 9.8 Tf [(=0.031\). Area was significantly )] TJ ET BT 26.250 531.930 Td /F1 9.8 Tf [(larger in YAC128 direct pathway MSNs compared to area of WTs \()] TJ ET BT 313.973 531.930 Td /F5 9.8 Tf [(t)] TJ ET BT 316.683 531.930 Td /F1 9.8 Tf [(=3.482, )] TJ ET BT 352.193 531.930 Td /F5 9.8 Tf [(p)] TJ ET BT 357.614 531.930 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3C\). The increased area of indirect )] TJ ET BT 26.250 520.025 Td /F1 9.8 Tf [(pathway MSNs approached statistical significance \()] TJ ET BT 248.413 520.025 Td /F5 9.8 Tf [(t)] TJ ET BT 251.124 520.025 Td /F1 9.8 Tf [(=1.829, )] TJ ET BT 286.634 520.025 Td /F5 9.8 Tf [(p)] TJ ET BT 292.055 520.025 Td /F1 9.8 Tf [(=0.077\). The increased area of direct versus indirect pathway )] TJ ET BT 26.250 508.120 Td /F1 9.8 Tf [(MSNs was significantly greater in only the YAC128 mice \()] TJ ET BT 274.953 508.120 Td /F5 9.8 Tf [(t)] TJ ET BT 277.663 508.120 Td /F1 9.8 Tf [(=2.353, )] TJ ET BT 313.173 508.120 Td /F5 9.8 Tf [(p)] TJ ET BT 318.594 508.120 Td /F1 9.8 Tf [(<0.025\). There was a significant increase in relative area in )] TJ ET BT 26.250 496.215 Td /F1 9.8 Tf [(indirect pathway MSNs in YAC128 mice compared to WTs \()] TJ ET BT 284.157 496.215 Td /F5 9.8 Tf [(t)] TJ ET BT 286.868 496.215 Td /F1 9.8 Tf [(=5.012, )] TJ ET BT 322.377 496.215 Td /F5 9.8 Tf [(p)] TJ ET BT 327.798 496.215 Td /F1 9.8 Tf [(<0.001\) as well as a significant increase in indirect )] TJ ET BT 26.250 484.311 Td /F1 9.8 Tf [(pathway MSNs compared to direct pathway MSNs but only in MSNs in YAC128 mice \()] TJ ET BT 398.495 484.311 Td /F5 9.8 Tf [(t)] TJ ET BT 401.206 484.311 Td /F1 9.8 Tf [(=4.158, )] TJ ET BT 436.715 484.311 Td /F5 9.8 Tf [(p)] TJ ET BT 442.136 484.311 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 238.080 555.000 236.350 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 474.430 m 581.250 474.430 l 581.250 473.680 l 26.250 473.680 l f 26.250 238.080 m 581.250 238.080 l 581.250 238.830 l 26.250 238.830 l f q 450.000 0 0 129.000 35.250 335.680 cm /I22 Do Q q 35.250 249.330 537.000 80.350 re W n 0.271 0.267 0.267 rg BT 35.250 320.156 Td /F4 9.8 Tf [(Fig. 3: Differential effects of DL-TBOA on direct and indirect pathway MSNs exemplified by percentage changes.)] TJ ET BT 35.250 300.786 Td /F1 9.8 Tf [(A: Percentage changes in peak response amplitude were differentially altered in direct and indirect pathway MSNs of )] TJ ET BT 35.250 287.050 Td /F1 9.8 Tf [(YAC128 mice. B: Percentage changes in decay times were greater in indirect pathway MSNs from YAC128 mice. C: )] TJ ET BT 35.250 273.313 Td /F1 9.8 Tf [(Percentage change in response area were greatest in direct pathway MSNs of YAC128 mice. D: Normalized area increases )] TJ ET BT 35.250 259.577 Td /F1 9.8 Tf [(were greatest in indirect pathway MSNs of YAC128 mice.)] TJ ET Q BT 26.250 203.858 Td /F4 12.0 Tf [(Discussion)] TJ ET BT 26.250 183.904 Td /F1 9.8 Tf [(The present study demonstrated decreases in evoked excitatory postsynaptic currents mediated by AMPARs and NMDARs in )] TJ ET BT 26.250 171.999 Td /F1 9.8 Tf [(YAC128 direct and indirect pathway MSNs. Furthermore, in contrast to the downregulation of synaptic activity, NMDAR-)] TJ ET BT 26.250 160.095 Td /F1 9.8 Tf [(mediated responses evoked in the presence of TBOA were increased in both direct and indirect pathway MSNs of YAC128 )] TJ ET BT 26.250 148.190 Td /F1 9.8 Tf [(mice, but in slightly different ways. In direct pathway MSNs, response increases were mainly due to higher amplitudes while in )] TJ ET BT 26.250 136.285 Td /F1 9.8 Tf [(indirect pathway MSNs, the increase was due to longer decay times. This outcome indicates differential changes in synaptic )] TJ ET BT 26.250 124.380 Td /F1 9.8 Tf [(versus extrasynaptic NMDA receptors in both direct and indirect pathway MSNs from these older, symptomatic YAC128 mice.)] TJ ET BT 26.250 104.976 Td /F1 9.8 Tf [(We previously reported a progressive disconnection of striatal MSNs from their glutamatergic inputs in various models of HD)] TJ ET 0.267 0.267 0.267 rg BT 561.593 106.483 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 566.412 108.864 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 568.821 106.483 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 573.640 104.976 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 93.071 Td /F1 9.8 Tf [(In 12 month-old YAC128 mice we also showed that the reduction in the spontaneous excitatory postsynaptic current frequency )] TJ ET BT 26.250 81.166 Td /F1 9.8 Tf [(was greater in direct than indirect pathway MSNs but we did not examine AMPAR and NMDAR responses separately and our )] TJ ET BT 26.250 69.261 Td /F1 9.8 Tf [(results were based on AMPAR-mediated responses)] TJ ET 0.267 0.267 0.267 rg BT 250.042 70.769 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 254.860 69.261 Td /F1 9.8 Tf [(. The reduction in evoked synaptic responses agrees with previous studies )] TJ ET BT 26.250 57.357 Td /F1 9.8 Tf [(showing decreases in glutamate release as well as AMPAR- and NMDAR-mediated currents in symptomatic YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 555.080 58.864 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 559.899 57.357 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 45.452 Td /F1 9.8 Tf [(The reduction in the amplitude of evoked responses indicates that there is a loss of glutamatergic inputs onto MSNs in YAC128 )] TJ ET Q q 15.000 31.166 577.500 745.834 re W n 0.965 0.965 0.965 rg 26.250 632.287 555.000 144.713 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 632.287 m 581.250 632.287 l 581.250 633.037 l 26.250 633.037 l f q 35.250 643.537 537.000 133.463 re W n 0.271 0.267 0.267 rg BT 35.250 767.476 Td /F4 9.8 Tf [(Fig. 2: DL-TBOA alters peak amplitude, decay, area and relative area of direct and indirect pathway MSNs of WT )] TJ ET BT 35.250 755.571 Td /F4 9.8 Tf [(and YAC128 mice.)] TJ ET BT 35.250 736.201 Td /F1 9.8 Tf [(A: Typical traces of NMDAR-mediated currents in WT and YAC128 direct and indirect pathway MSNs evoked at 75% of the )] TJ ET BT 35.250 722.465 Td /F1 9.8 Tf [(maximal stimulus intensity showing alterations in peak amplitudes and decay times. B: Peak amplitude is differentially )] TJ ET BT 35.250 708.729 Td /F1 9.8 Tf [(altered in direct and indirect pathway MSNs by DL-TBOA. Peak amplitude increases in direct pathway MSNs but decreases )] TJ ET BT 35.250 694.993 Td /F1 9.8 Tf [(in indirect pathway MSNs in YAC128 mice. C: In DL-TBOA response decay time increases in direct and indirect pathway )] TJ ET BT 35.250 681.256 Td /F1 9.8 Tf [(MSNs in both WT and YAC128 mice. D: In DL-TBOA, response area increases only in direct pathway MSNs in YAC128 )] TJ ET BT 35.250 667.520 Td /F1 9.8 Tf [(mice and in indirect pathway MSNs in both WT and YAC128 mice. E: In DL-TBOA, normalized area increases in all groups )] TJ ET BT 35.250 653.784 Td /F1 9.8 Tf [(but displays the greatest increase in indirect pathway MSNs in YAC128 mice.)] TJ ET Q BT 26.250 615.263 Td /F1 9.8 Tf [(To more specifically compare the effects of TBOA in direct and indirect pathway MSNs, changes induced by TBOA were )] TJ ET BT 26.250 603.358 Td /F1 9.8 Tf [(converted to percentages \(Fig. 3\). In direct pathway MSNs, TBOA decreased the evoked NMDAR peak current in WTs but )] TJ ET BT 26.250 591.453 Td /F1 9.8 Tf [(increased it in YAC128 mice \()] TJ ET BT 154.667 591.453 Td /F5 9.8 Tf [(t)] TJ ET BT 157.378 591.453 Td /F1 9.8 Tf [(=6.449,)] TJ ET BT 190.177 591.453 Td /F5 9.8 Tf [( p)] TJ ET BT 198.308 591.453 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3A\). In indirect pathway MSNs TBOA decreased the peak in both WT and )] TJ ET BT 26.250 579.549 Td /F1 9.8 Tf [(YAC128 mice, but the reduction was significantly larger in YAC128 D2 MSNs \()] TJ ET BT 364.370 579.549 Td /F5 9.8 Tf [(t)] TJ ET BT 367.081 579.549 Td /F1 9.8 Tf [(=3.323, )] TJ ET BT 402.590 579.549 Td /F5 9.8 Tf [(p)] TJ ET BT 408.011 579.549 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3A\). The difference in )] TJ ET BT 26.250 567.644 Td /F1 9.8 Tf [(peak amplitude between direct and indirect pathway MSNs in YAC128 mice also was significant \()] TJ ET BT 445.120 567.644 Td /F5 9.8 Tf [(t)] TJ ET BT 447.830 567.644 Td /F1 9.8 Tf [(=9.150, )] TJ ET BT 483.340 567.644 Td /F5 9.8 Tf [(p)] TJ ET BT 488.761 567.644 Td /F1 9.8 Tf [(<0.001\). Decay time )] TJ ET BT 26.250 555.739 Td /F1 9.8 Tf [(was significantly elevated in indirect YAC128 MSNs compared to WTs \()] TJ ET BT 334.555 555.739 Td /F5 9.8 Tf [(t)] TJ ET BT 337.265 555.739 Td /F1 9.8 Tf [(=3.601, )] TJ ET BT 372.775 555.739 Td /F5 9.8 Tf [(p)] TJ ET BT 378.196 555.739 Td /F1 9.8 Tf [(=0.001\) \(Fig. 3B\) and also was significantly )] TJ ET BT 26.250 543.834 Td /F1 9.8 Tf [(greater in indirect pathway MSNs compared to direct pathway MSNs in YAC128 mice \()] TJ ET BT 400.123 543.834 Td /F5 9.8 Tf [(t)] TJ ET BT 402.834 543.834 Td /F1 9.8 Tf [(=2.258, )] TJ ET BT 438.344 543.834 Td /F5 9.8 Tf [(p)] TJ ET BT 443.764 543.834 Td /F1 9.8 Tf [(=0.031\). Area was significantly )] TJ ET BT 26.250 531.930 Td /F1 9.8 Tf [(larger in YAC128 direct pathway MSNs compared to area of WTs \()] TJ ET BT 313.973 531.930 Td /F5 9.8 Tf [(t)] TJ ET BT 316.683 531.930 Td /F1 9.8 Tf [(=3.482, )] TJ ET BT 352.193 531.930 Td /F5 9.8 Tf [(p)] TJ ET BT 357.614 531.930 Td /F1 9.8 Tf [(=0.002\) \(Fig. 3C\). The increased area of indirect )] TJ ET BT 26.250 520.025 Td /F1 9.8 Tf [(pathway MSNs approached statistical significance \()] TJ ET BT 248.413 520.025 Td /F5 9.8 Tf [(t)] TJ ET BT 251.124 520.025 Td /F1 9.8 Tf [(=1.829, )] TJ ET BT 286.634 520.025 Td /F5 9.8 Tf [(p)] TJ ET BT 292.055 520.025 Td /F1 9.8 Tf [(=0.077\). The increased area of direct versus indirect pathway )] TJ ET BT 26.250 508.120 Td /F1 9.8 Tf [(MSNs was significantly greater in only the YAC128 mice \()] TJ ET BT 274.953 508.120 Td /F5 9.8 Tf [(t)] TJ ET BT 277.663 508.120 Td /F1 9.8 Tf [(=2.353, )] TJ ET BT 313.173 508.120 Td /F5 9.8 Tf [(p)] TJ ET BT 318.594 508.120 Td /F1 9.8 Tf [(<0.025\). There was a significant increase in relative area in )] TJ ET BT 26.250 496.215 Td /F1 9.8 Tf [(indirect pathway MSNs in YAC128 mice compared to WTs \()] TJ ET BT 284.157 496.215 Td /F5 9.8 Tf [(t)] TJ ET BT 286.868 496.215 Td /F1 9.8 Tf [(=5.012, )] TJ ET BT 322.377 496.215 Td /F5 9.8 Tf [(p)] TJ ET BT 327.798 496.215 Td /F1 9.8 Tf [(<0.001\) as well as a significant increase in indirect )] TJ ET BT 26.250 484.311 Td /F1 9.8 Tf [(pathway MSNs compared to direct pathway MSNs but only in MSNs in YAC128 mice \()] TJ ET BT 398.495 484.311 Td /F5 9.8 Tf [(t)] TJ ET BT 401.206 484.311 Td /F1 9.8 Tf [(=4.158, )] TJ ET BT 436.715 484.311 Td /F5 9.8 Tf [(p)] TJ ET BT 442.136 484.311 Td /F1 9.8 Tf [(<0.001\) \(Fig. 3D\).)] TJ ET 0.965 0.965 0.965 rg 26.250 238.080 555.000 236.350 re f 0.267 0.267 0.267 rg 0.267 0.267 0.267 RG 26.250 474.430 m 581.250 474.430 l 581.250 473.680 l 26.250 473.680 l f 26.250 238.080 m 581.250 238.080 l 581.250 238.830 l 26.250 238.830 l f q 450.000 0 0 129.000 35.250 335.680 cm /I24 Do Q q 35.250 249.330 537.000 80.350 re W n 0.271 0.267 0.267 rg BT 35.250 320.156 Td /F4 9.8 Tf [(Fig. 3: Differential effects of DL-TBOA on direct and indirect pathway MSNs exemplified by percentage changes.)] TJ ET BT 35.250 300.786 Td /F1 9.8 Tf [(A: Percentage changes in peak response amplitude were differentially altered in direct and indirect pathway MSNs of )] TJ ET BT 35.250 287.050 Td /F1 9.8 Tf [(YAC128 mice. B: Percentage changes in decay times were greater in indirect pathway MSNs from YAC128 mice. C: )] TJ ET BT 35.250 273.313 Td /F1 9.8 Tf [(Percentage change in response area were greatest in direct pathway MSNs of YAC128 mice. D: Normalized area increases )] TJ ET BT 35.250 259.577 Td /F1 9.8 Tf [(were greatest in indirect pathway MSNs of YAC128 mice.)] TJ ET Q BT 26.250 203.858 Td /F4 12.0 Tf [(Discussion)] TJ ET BT 26.250 183.904 Td /F1 9.8 Tf [(The present study demonstrated decreases in evoked excitatory postsynaptic currents mediated by AMPARs and NMDARs in )] TJ ET BT 26.250 171.999 Td /F1 9.8 Tf [(YAC128 direct and indirect pathway MSNs. Furthermore, in contrast to the downregulation of synaptic activity, NMDAR-)] TJ ET BT 26.250 160.095 Td /F1 9.8 Tf [(mediated responses evoked in the presence of TBOA were increased in both direct and indirect pathway MSNs of YAC128 )] TJ ET BT 26.250 148.190 Td /F1 9.8 Tf [(mice, but in slightly different ways. In direct pathway MSNs, response increases were mainly due to higher amplitudes while in )] TJ ET BT 26.250 136.285 Td /F1 9.8 Tf [(indirect pathway MSNs, the increase was due to longer decay times. This outcome indicates differential changes in synaptic )] TJ ET BT 26.250 124.380 Td /F1 9.8 Tf [(versus extrasynaptic NMDA receptors in both direct and indirect pathway MSNs from these older, symptomatic YAC128 mice.)] TJ ET BT 26.250 104.976 Td /F1 9.8 Tf [(We previously reported a progressive disconnection of striatal MSNs from their glutamatergic inputs in various models of HD)] TJ ET 0.267 0.267 0.267 rg BT 561.593 106.483 Td /F4 8.7 Tf [(7)] TJ ET 0.271 0.267 0.267 rg BT 566.412 108.864 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 568.821 106.483 Td /F4 8.7 Tf [(8)] TJ ET 0.271 0.267 0.267 rg BT 573.640 104.976 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 93.071 Td /F1 9.8 Tf [(In 12 month-old YAC128 mice we also showed that the reduction in the spontaneous excitatory postsynaptic current frequency )] TJ ET BT 26.250 81.166 Td /F1 9.8 Tf [(was greater in direct than indirect pathway MSNs but we did not examine AMPAR and NMDAR responses separately and our )] TJ ET BT 26.250 69.261 Td /F1 9.8 Tf [(results were based on AMPAR-mediated responses)] TJ ET 0.267 0.267 0.267 rg BT 250.042 70.769 Td /F4 8.7 Tf [(4)] TJ ET 0.271 0.267 0.267 rg BT 254.860 69.261 Td /F1 9.8 Tf [(. The reduction in evoked synaptic responses agrees with previous studies )] TJ ET BT 26.250 57.357 Td /F1 9.8 Tf [(showing decreases in glutamate release as well as AMPAR- and NMDAR-mediated currents in symptomatic YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 555.080 58.864 Td /F4 8.7 Tf [(9)] TJ ET 0.271 0.267 0.267 rg BT 559.899 57.357 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 45.452 Td /F1 9.8 Tf [(The reduction in the amplitude of evoked responses indicates that there is a loss of glutamatergic inputs onto MSNs in YAC128 )] TJ ET Q q 450.000 0 0 129.000 35.250 335.680 cm /I26 Do Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(5)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Huntington Disease)] TJ ET Q endstream endobj 219 0 obj << /Type /Annot /Subtype /Link /A 220 0 R /Border [0 0 0] /H /I /Rect [ 35.2500 335.6797 485.2500 464.6797 ] >> endobj 220 0 obj << /Type /Action /S /URI /URI (https://currents.plos.org/hd/files/2014/03/Figure3.png) >> endobj 221 0 obj << /Type /XObject /Subtype /Image /Width 600 /Height 172 /Filter /FlateDecode /DecodeParms << /Predictor 15 /Colors 1 /Columns 600 /BitsPerComponent 8>> /ColorSpace /DeviceGray /BitsPerComponent 8 /Length 9629>> stream xdj?H$I&I&$LL2333dff&LLd$I5u]㺮ityv{_=;|sWp)^j ̳swyK̵P MHmtqy;q1S_ͻΎ˿! 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@{EYl;`y6Kӄ-emj,*ДR[m4TkӢXVت^ uE*g'iDjOQQpytNim X }.K*B*PU*M:S=i<6/GD!iIC74Q,TJ%_6M QOE#rڙ?~녟D^E)fFƔGFOSG?U?>?:4՞q"\ `GzX^*;}jA_2j4{ˢBbv26[ 5Y]X&lX aybBtLi R{)P+/ܺ6:ZnxsS~:܇4Ὡ7 'f]@EӁ"vaSMcz[D'%Ói- #L6X][N<a&/ub:1I`_U+penWm?%"ѵpa" iv GpmpRA~2kK;{v5T{('udB> R+P^Qhڷ%B`>8KQG"6HAP!*M Owb (P#ힰ.ƐRu\;]Nvl lӚ @p$2ԏH9Wӟ_/zV^:fԱz_mbvvzw:V"j & iš3L|M7}D/|_?:H_<#m=.ԃ=G4WlYe' (U –IqL:VQC/|a=zrm0OĴӂq+۽d:&Lh>=B0m<5bQ62ՂܙC:bMl Y.YZik{ˢJg]8+b*4T-SHIJ43J'nKZ+evsMҽ`g[p62D,m|Qp" R6\aW.͛h tq3J5@5=Xt:t됹]2 IY \OQ2#\*h`JqEhNMGX'qG& )P2I9v'J}P&]s%-FA]Na񲂮>YqJU5aܚ_ tvB씵PvELa hXrО| Kawp$ SSߓ]R2BHюlڦqTʔ}qSG齟(Ol:H$\PB4qe:VSWĚ2pch%BITlba!=bZp(-U/ ;bѳI7E隐dv,pݟ3Vr&bqY#dE\>YmvXSMF 5l++B9Vxu6b|Yx-L^jxTu1'|^loĊ|[!\pmw'Zlc/+zw}Z'F*uƢvwSIOnFsaE,ߛ:A7xmĜpx}eL.!v䥆 3c8 %qɣpCw^-݋p1-n]#Ƅ=#oNlq;{w ,۾ۈ9iGH ,ۄs]lbEHy^^^xKsE8L"$|Oomn/mrLq!yky6Flr=.KT!yyhn4Ó84J̖kr&rb0Tmĸش<_l?Y*i]|I,y<+BiDOsn4p,#Ңi,p(3ҹ /3ZhgN<݋X<_i2%1wwUVr,"$,I@N{ 'p!-Z1!CƱ.E}|^lkl~>ye[Ê> endobj 258 0 obj << /Length 29527 >> stream 0.271 0.267 0.267 rg q 15.000 33.768 577.500 743.232 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(mice at this age for MSNs originating both pathways in HD. The loss of glutamatergic inputs may be due to pre- and )] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(postsynaptic mechanisms. In YAC128 mice a reduction in spine density has been observed)] TJ ET 0.267 0.267 0.267 rg BT 420.218 757.079 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 429.856 755.571 Td /F1 9.8 Tf [(. While the reduction in spine )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(density could be caused by pre- and post-synaptic mechanisms, recent studies demonstrate a critical role of postsynaptic )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(NMDA receptors containing GluN3A subunits in HD spine loss)] TJ ET 0.267 0.267 0.267 rg BT 294.463 733.269 Td /F4 8.7 Tf [(11)] TJ ET 0.271 0.267 0.267 rg BT 304.100 731.762 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 712.357 Td /F1 9.8 Tf [(In contrast to the reduction in evoked synaptic currents, NMDA responses evoked in the presence of TBOA were increased. )] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(Previous studies demonstrated an increase in the extrasynaptic NMDAR-mediated responses in young YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 531.281 701.960 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 536.099 700.452 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(However, differences between direct and indirect pathway MSNs were not examined. The present study not only confirms that )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(this increase in extrasynaptic NMDAR-mediated responses occurs in old YAC128 mice, but also demonstrates that it occurs in )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(both direct and indirect pathway MSNs though it appears to be mechanistically due to differing underlying effetcs. Direct )] TJ ET BT 26.250 652.833 Td /F1 9.8 Tf [(pathway MSNs show a larger increase in peak amplitude while indirect pathway MSNs display a larger increase in normalized )] TJ ET BT 26.250 640.929 Td /F1 9.8 Tf [(area due to a longer decay time. This latter effect could suggest that MSNs of the indirect pathway are more sensitive to )] TJ ET BT 26.250 629.024 Td /F1 9.8 Tf [(changes in glutamate transport, thus contributing to their increased vulnerability in HD.)] TJ ET BT 26.250 609.619 Td /F1 9.8 Tf [(NMDARs are composed by GluN2A and GluN2B subunits. GluN2A subunits are thought to be restricted mainly to the synaptic )] TJ ET BT 26.250 597.714 Td /F1 9.8 Tf [(site)] TJ ET 0.267 0.267 0.267 rg BT 41.421 599.222 Td /F4 8.7 Tf [(12)] TJ ET 0.271 0.267 0.267 rg BT 51.058 597.714 Td /F1 9.8 Tf [(, while the GluN2B subunits are located on synaptic and extrasynaptic sites)] TJ ET 0.267 0.267 0.267 rg BT 375.665 599.222 Td /F4 8.7 Tf [(13)] TJ ET 0.271 0.267 0.267 rg BT 385.302 597.714 Td /F1 9.8 Tf [(. It has been suggested that the activation of )] TJ ET BT 26.250 585.810 Td /F1 9.8 Tf [(the synaptic NMDARs is neuroprotective, while activation of extrasynaptic NMDA receptors stimulates a pro-apoptotic cellular )] TJ ET BT 26.250 573.905 Td /F1 9.8 Tf [(pathway)] TJ ET 0.267 0.267 0.267 rg BT 62.559 575.412 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 72.196 573.905 Td /F1 9.8 Tf [(. Alterations in NMDAR subunit trafficking is thought to contribute to increased density of GluN2B subunits at )] TJ ET BT 26.250 562.000 Td /F1 9.8 Tf [(extrasynaptic sites in HD)] TJ ET 0.267 0.267 0.267 rg BT 133.529 563.507 Td /F4 8.7 Tf [(15)] TJ ET 0.271 0.267 0.267 rg BT 143.167 565.888 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 145.576 563.507 Td /F4 8.7 Tf [(16)] TJ ET 0.271 0.267 0.267 rg BT 155.213 562.000 Td /F1 9.8 Tf [(. In a recent study employing co-culture of cortical and striatal MSNs from YAC128 mice, a larger )] TJ ET BT 26.250 550.095 Td /F1 9.8 Tf [(extrasynaptic NMDAR-mediated current occurred in association with an increase in GluN2B subunit expression at the cellular )] TJ ET BT 26.250 538.191 Td /F1 9.8 Tf [(surface compared with the co-culture obtained from WT mice)] TJ ET 0.267 0.267 0.267 rg BT 289.588 539.698 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 294.406 542.079 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 296.816 539.698 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 306.453 538.191 Td /F1 9.8 Tf [(. In addition, low-dose memantine, a blocker of GluN2B-)] TJ ET BT 26.250 526.286 Td /F1 9.8 Tf [(containing NMDARs at extrasynaptic sites, ameliorates neuropathological and behavioral manifestations of HD in YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 575.711 527.793 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 26.250 518.269 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 28.659 515.888 Td /F4 8.7 Tf [(18)] TJ ET 0.271 0.267 0.267 rg BT 38.297 514.381 Td /F1 9.8 Tf [(. In co-cultured YAC128 MSNs, low doses of memantine and ifenprodil attenuated extrasynaptic NMDAR-mediated neuronal )] TJ ET BT 26.250 502.476 Td /F1 9.8 Tf [(apoptosis)] TJ ET 0.267 0.267 0.267 rg BT 67.980 503.984 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 77.617 502.476 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 483.072 Td /F1 9.8 Tf [(In conclusion, the present results showed that the glutamatergic inputs to MSNs are decreased in 16-month old YAC128 mice. )] TJ ET BT 26.250 471.167 Td /F1 9.8 Tf [(This study also demonstrates that in late stage HD, NMDA responses evoked in both direct and indirect pathway MSNs in the )] TJ ET BT 26.250 459.262 Td /F1 9.8 Tf [(presence of TBOA are significantly increased in YAC128 mice and the parameters for the increase are different. Direct pathway )] TJ ET BT 26.250 447.357 Td /F1 9.8 Tf [(MSNs in YAC128 mice display increases in peak amplitude while indirect pathway MSNs in YAC128 mice display increases in )] TJ ET BT 26.250 435.453 Td /F1 9.8 Tf [(normalized area due to significantly slower decay times. These results suggest that alterations in glutamate reuptake have )] TJ ET BT 26.250 423.548 Td /F1 9.8 Tf [(different effects on indirect compared with direct pathway MSNs and highlight previous studies showing that MSNs of both )] TJ ET BT 26.250 411.643 Td /F1 9.8 Tf [(pathways are differentially affected in mouse models of HD)] TJ ET 0.267 0.267 0.267 rg BT 280.394 413.150 Td /F4 8.7 Tf [(19)] TJ ET 0.271 0.267 0.267 rg BT 290.031 415.531 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 292.440 413.150 Td /F4 8.7 Tf [(20)] TJ ET 0.271 0.267 0.267 rg BT 302.078 411.643 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 375.041 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 355.086 Td /F1 9.8 Tf [(The authors thank Donna Crandall for help with the illustrations.)] TJ ET BT 26.250 325.984 Td /F4 12.0 Tf [(References)] TJ ET BT 26.250 298.530 Td /F1 9.8 Tf [(1.)] TJ ET BT 38.132 298.530 Td /F1 9.8 Tf [(G. Bates, P.S. Harper, L. Jones \(Eds.\), Huntington's disease, Oxford University Press, Oxford ; New York, 2002, xvi, 558 p.)] TJ ET BT 26.250 279.125 Td /F1 9.8 Tf [(2.)] TJ ET BT 38.132 279.125 Td /F1 9.8 Tf [(Slow EJ, van Raamsdonk J, Rogers D, Coleman SH, Graham RK, Deng Y, Oh R, Bissada N, Hossain SM, Yang YZ, Li XJ, )] TJ ET BT 26.250 267.220 Td /F1 9.8 Tf [(Simpson EM, Gutekunst CA, Leavitt BR, Hayden MR. Selective striatal neuronal loss in a YAC128 mouse model of Huntington )] TJ ET BT 26.250 255.315 Td /F1 9.8 Tf [(disease. Hum Mol Genet. 2003 Jul 1;12\(13\):1555-67. PubMed PMID:12812983.)] TJ ET BT 26.250 235.911 Td /F1 9.8 Tf [(3.)] TJ ET BT 38.132 235.911 Td /F1 9.8 Tf [(Ferrante RJ. Mouse models of Huntington's disease and methodological considerations for therapeutic trials. Biochim )] TJ ET BT 26.250 224.006 Td /F1 9.8 Tf [(Biophys Acta. 2009 Jun;1792\(6\):506-20. PubMed PMID:19362590.)] TJ ET BT 26.250 204.601 Td /F1 9.8 Tf [(4.)] TJ ET BT 38.132 204.601 Td /F1 9.8 Tf [(Andr VM, Cepeda C, Fisher YE, Huynh M, Bardakjian N, Singh S, Yang XW, Levine MS. Differential electrophysiological )] TJ ET BT 26.250 192.696 Td /F1 9.8 Tf [(changes in striatal output neurons in Huntington's disease. J Neurosci. 2011 Jan 26;31\(4\):1170-82. PubMed PMID:21273402.)] TJ ET BT 26.250 173.292 Td /F1 9.8 Tf [(5.)] TJ ET BT 38.132 173.292 Td /F1 9.8 Tf [(Andr VM, Fisher YE, Levine MS. Altered Balance of Activity in the Striatal Direct and Indirect Pathways in Mouse Models of )] TJ ET BT 26.250 161.387 Td /F1 9.8 Tf [(Huntington's Disease. Front Syst Neurosci. 2011;5:46. PubMed PMID:21720523.)] TJ ET BT 26.250 141.982 Td /F1 9.8 Tf [(6.)] TJ ET BT 38.132 141.982 Td /F1 9.8 Tf [(Milnerwood AJ, Gladding CM, Pouladi MA, Kaufman AM, Hines RM, Boyd JD, Ko RW, Vasuta OC, Graham RK, Hayden MR, )] TJ ET BT 26.250 130.077 Td /F1 9.8 Tf [(Murphy TH, Raymond LA. Early increase in extrasynaptic NMDA receptor signaling and expression contributes to phenotype )] TJ ET BT 26.250 118.173 Td /F1 9.8 Tf [(onset in Huntington's disease mice. Neuron. 2010 Jan 28;65\(2\):178-90. PubMed PMID:20152125.)] TJ ET BT 26.250 98.768 Td /F1 9.8 Tf [(7.)] TJ ET BT 38.132 98.768 Td /F1 9.8 Tf [(Cepeda C, Hurst RS, Calvert CR, Hernndez-Echeagaray E, Nguyen OK, Jocoy E, Christian LJ, Ariano MA, Levine MS. )] TJ ET BT 26.250 86.863 Td /F1 9.8 Tf [(Transient and progressive electrophysiological alterations in the corticostriatal pathway in a mouse model of Huntington's )] TJ ET BT 26.250 74.958 Td /F1 9.8 Tf [(disease. J Neurosci. 2003 Feb 1;23\(3\):961-9. PubMed PMID:12574425.)] TJ ET BT 26.250 55.554 Td /F1 9.8 Tf [(8.)] TJ ET BT 38.132 55.554 Td /F1 9.8 Tf [(Cummings DM, Cepeda C, Levine MS. Alterations in striatal synaptic transmission are consistent across genetic mouse )] TJ ET BT 26.250 43.649 Td /F1 9.8 Tf [(models of Huntington's disease. ASN Neuro. 2010 Jun 18;2\(3\):e00036. PubMed PMID:20585470.)] TJ ET Q q 15.000 33.768 577.500 743.232 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(mice at this age for MSNs originating both pathways in HD. The loss of glutamatergic inputs may be due to pre- and )] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(postsynaptic mechanisms. In YAC128 mice a reduction in spine density has been observed)] TJ ET 0.267 0.267 0.267 rg BT 420.218 757.079 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 429.856 755.571 Td /F1 9.8 Tf [(. While the reduction in spine )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(density could be caused by pre- and post-synaptic mechanisms, recent studies demonstrate a critical role of postsynaptic )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(NMDA receptors containing GluN3A subunits in HD spine loss)] TJ ET 0.267 0.267 0.267 rg BT 294.463 733.269 Td /F4 8.7 Tf [(11)] TJ ET 0.271 0.267 0.267 rg BT 304.100 731.762 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 712.357 Td /F1 9.8 Tf [(In contrast to the reduction in evoked synaptic currents, NMDA responses evoked in the presence of TBOA were increased. )] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(Previous studies demonstrated an increase in the extrasynaptic NMDAR-mediated responses in young YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 531.281 701.960 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 536.099 700.452 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(However, differences between direct and indirect pathway MSNs were not examined. The present study not only confirms that )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(this increase in extrasynaptic NMDAR-mediated responses occurs in old YAC128 mice, but also demonstrates that it occurs in )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(both direct and indirect pathway MSNs though it appears to be mechanistically due to differing underlying effetcs. Direct )] TJ ET BT 26.250 652.833 Td /F1 9.8 Tf [(pathway MSNs show a larger increase in peak amplitude while indirect pathway MSNs display a larger increase in normalized )] TJ ET BT 26.250 640.929 Td /F1 9.8 Tf [(area due to a longer decay time. This latter effect could suggest that MSNs of the indirect pathway are more sensitive to )] TJ ET BT 26.250 629.024 Td /F1 9.8 Tf [(changes in glutamate transport, thus contributing to their increased vulnerability in HD.)] TJ ET BT 26.250 609.619 Td /F1 9.8 Tf [(NMDARs are composed by GluN2A and GluN2B subunits. GluN2A subunits are thought to be restricted mainly to the synaptic )] TJ ET BT 26.250 597.714 Td /F1 9.8 Tf [(site)] TJ ET 0.267 0.267 0.267 rg BT 41.421 599.222 Td /F4 8.7 Tf [(12)] TJ ET 0.271 0.267 0.267 rg BT 51.058 597.714 Td /F1 9.8 Tf [(, while the GluN2B subunits are located on synaptic and extrasynaptic sites)] TJ ET 0.267 0.267 0.267 rg BT 375.665 599.222 Td /F4 8.7 Tf [(13)] TJ ET 0.271 0.267 0.267 rg BT 385.302 597.714 Td /F1 9.8 Tf [(. It has been suggested that the activation of )] TJ ET BT 26.250 585.810 Td /F1 9.8 Tf [(the synaptic NMDARs is neuroprotective, while activation of extrasynaptic NMDA receptors stimulates a pro-apoptotic cellular )] TJ ET BT 26.250 573.905 Td /F1 9.8 Tf [(pathway)] TJ ET 0.267 0.267 0.267 rg BT 62.559 575.412 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 72.196 573.905 Td /F1 9.8 Tf [(. Alterations in NMDAR subunit trafficking is thought to contribute to increased density of GluN2B subunits at )] TJ ET BT 26.250 562.000 Td /F1 9.8 Tf [(extrasynaptic sites in HD)] TJ ET 0.267 0.267 0.267 rg BT 133.529 563.507 Td /F4 8.7 Tf [(15)] TJ ET 0.271 0.267 0.267 rg BT 143.167 565.888 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 145.576 563.507 Td /F4 8.7 Tf [(16)] TJ ET 0.271 0.267 0.267 rg BT 155.213 562.000 Td /F1 9.8 Tf [(. In a recent study employing co-culture of cortical and striatal MSNs from YAC128 mice, a larger )] TJ ET BT 26.250 550.095 Td /F1 9.8 Tf [(extrasynaptic NMDAR-mediated current occurred in association with an increase in GluN2B subunit expression at the cellular )] TJ ET BT 26.250 538.191 Td /F1 9.8 Tf [(surface compared with the co-culture obtained from WT mice)] TJ ET 0.267 0.267 0.267 rg BT 289.588 539.698 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 294.406 542.079 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 296.816 539.698 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 306.453 538.191 Td /F1 9.8 Tf [(. In addition, low-dose memantine, a blocker of GluN2B-)] TJ ET BT 26.250 526.286 Td /F1 9.8 Tf [(containing NMDARs at extrasynaptic sites, ameliorates neuropathological and behavioral manifestations of HD in YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 575.711 527.793 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 26.250 518.269 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 28.659 515.888 Td /F4 8.7 Tf [(18)] TJ ET 0.271 0.267 0.267 rg BT 38.297 514.381 Td /F1 9.8 Tf [(. In co-cultured YAC128 MSNs, low doses of memantine and ifenprodil attenuated extrasynaptic NMDAR-mediated neuronal )] TJ ET BT 26.250 502.476 Td /F1 9.8 Tf [(apoptosis)] TJ ET 0.267 0.267 0.267 rg BT 67.980 503.984 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 77.617 502.476 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 483.072 Td /F1 9.8 Tf [(In conclusion, the present results showed that the glutamatergic inputs to MSNs are decreased in 16-month old YAC128 mice. )] TJ ET BT 26.250 471.167 Td /F1 9.8 Tf [(This study also demonstrates that in late stage HD, NMDA responses evoked in both direct and indirect pathway MSNs in the )] TJ ET BT 26.250 459.262 Td /F1 9.8 Tf [(presence of TBOA are significantly increased in YAC128 mice and the parameters for the increase are different. Direct pathway )] TJ ET BT 26.250 447.357 Td /F1 9.8 Tf [(MSNs in YAC128 mice display increases in peak amplitude while indirect pathway MSNs in YAC128 mice display increases in )] TJ ET BT 26.250 435.453 Td /F1 9.8 Tf [(normalized area due to significantly slower decay times. These results suggest that alterations in glutamate reuptake have )] TJ ET BT 26.250 423.548 Td /F1 9.8 Tf [(different effects on indirect compared with direct pathway MSNs and highlight previous studies showing that MSNs of both )] TJ ET BT 26.250 411.643 Td /F1 9.8 Tf [(pathways are differentially affected in mouse models of HD)] TJ ET 0.267 0.267 0.267 rg BT 280.394 413.150 Td /F4 8.7 Tf [(19)] TJ ET 0.271 0.267 0.267 rg BT 290.031 415.531 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 292.440 413.150 Td /F4 8.7 Tf [(20)] TJ ET 0.271 0.267 0.267 rg BT 302.078 411.643 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 375.041 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 355.086 Td /F1 9.8 Tf [(The authors thank Donna Crandall for help with the illustrations.)] TJ ET BT 26.250 325.984 Td /F4 12.0 Tf [(References)] TJ ET BT 26.250 298.530 Td /F1 9.8 Tf [(1.)] TJ ET BT 38.132 298.530 Td /F1 9.8 Tf [(G. Bates, P.S. Harper, L. Jones \(Eds.\), Huntington's disease, Oxford University Press, Oxford ; New York, 2002, xvi, 558 p.)] TJ ET BT 26.250 279.125 Td /F1 9.8 Tf [(2.)] TJ ET BT 38.132 279.125 Td /F1 9.8 Tf [(Slow EJ, van Raamsdonk J, Rogers D, Coleman SH, Graham RK, Deng Y, Oh R, Bissada N, Hossain SM, Yang YZ, Li XJ, )] TJ ET BT 26.250 267.220 Td /F1 9.8 Tf [(Simpson EM, Gutekunst CA, Leavitt BR, Hayden MR. Selective striatal neuronal loss in a YAC128 mouse model of Huntington )] TJ ET BT 26.250 255.315 Td /F1 9.8 Tf [(disease. Hum Mol Genet. 2003 Jul 1;12\(13\):1555-67. PubMed PMID:12812983.)] TJ ET BT 26.250 235.911 Td /F1 9.8 Tf [(3.)] TJ ET BT 38.132 235.911 Td /F1 9.8 Tf [(Ferrante RJ. Mouse models of Huntington's disease and methodological considerations for therapeutic trials. Biochim )] TJ ET BT 26.250 224.006 Td /F1 9.8 Tf [(Biophys Acta. 2009 Jun;1792\(6\):506-20. PubMed PMID:19362590.)] TJ ET BT 26.250 204.601 Td /F1 9.8 Tf [(4.)] TJ ET BT 38.132 204.601 Td /F1 9.8 Tf [(Andr VM, Cepeda C, Fisher YE, Huynh M, Bardakjian N, Singh S, Yang XW, Levine MS. Differential electrophysiological )] TJ ET BT 26.250 192.696 Td /F1 9.8 Tf [(changes in striatal output neurons in Huntington's disease. J Neurosci. 2011 Jan 26;31\(4\):1170-82. PubMed PMID:21273402.)] TJ ET BT 26.250 173.292 Td /F1 9.8 Tf [(5.)] TJ ET BT 38.132 173.292 Td /F1 9.8 Tf [(Andr VM, Fisher YE, Levine MS. Altered Balance of Activity in the Striatal Direct and Indirect Pathways in Mouse Models of )] TJ ET BT 26.250 161.387 Td /F1 9.8 Tf [(Huntington's Disease. Front Syst Neurosci. 2011;5:46. PubMed PMID:21720523.)] TJ ET BT 26.250 141.982 Td /F1 9.8 Tf [(6.)] TJ ET BT 38.132 141.982 Td /F1 9.8 Tf [(Milnerwood AJ, Gladding CM, Pouladi MA, Kaufman AM, Hines RM, Boyd JD, Ko RW, Vasuta OC, Graham RK, Hayden MR, )] TJ ET BT 26.250 130.077 Td /F1 9.8 Tf [(Murphy TH, Raymond LA. Early increase in extrasynaptic NMDA receptor signaling and expression contributes to phenotype )] TJ ET BT 26.250 118.173 Td /F1 9.8 Tf [(onset in Huntington's disease mice. Neuron. 2010 Jan 28;65\(2\):178-90. PubMed PMID:20152125.)] TJ ET BT 26.250 98.768 Td /F1 9.8 Tf [(7.)] TJ ET BT 38.132 98.768 Td /F1 9.8 Tf [(Cepeda C, Hurst RS, Calvert CR, Hernndez-Echeagaray E, Nguyen OK, Jocoy E, Christian LJ, Ariano MA, Levine MS. )] TJ ET BT 26.250 86.863 Td /F1 9.8 Tf [(Transient and progressive electrophysiological alterations in the corticostriatal pathway in a mouse model of Huntington's )] TJ ET BT 26.250 74.958 Td /F1 9.8 Tf [(disease. J Neurosci. 2003 Feb 1;23\(3\):961-9. PubMed PMID:12574425.)] TJ ET BT 26.250 55.554 Td /F1 9.8 Tf [(8.)] TJ ET BT 38.132 55.554 Td /F1 9.8 Tf [(Cummings DM, Cepeda C, Levine MS. Alterations in striatal synaptic transmission are consistent across genetic mouse )] TJ ET BT 26.250 43.649 Td /F1 9.8 Tf [(models of Huntington's disease. ASN Neuro. 2010 Jun 18;2\(3\):e00036. PubMed PMID:20585470.)] TJ ET Q q 15.000 33.768 577.500 743.232 re W n 0.271 0.267 0.267 rg BT 26.250 767.476 Td /F1 9.8 Tf [(mice at this age for MSNs originating both pathways in HD. The loss of glutamatergic inputs may be due to pre- and )] TJ ET BT 26.250 755.571 Td /F1 9.8 Tf [(postsynaptic mechanisms. In YAC128 mice a reduction in spine density has been observed)] TJ ET 0.267 0.267 0.267 rg BT 420.218 757.079 Td /F4 8.7 Tf [(10)] TJ ET 0.271 0.267 0.267 rg BT 429.856 755.571 Td /F1 9.8 Tf [(. While the reduction in spine )] TJ ET BT 26.250 743.667 Td /F1 9.8 Tf [(density could be caused by pre- and post-synaptic mechanisms, recent studies demonstrate a critical role of postsynaptic )] TJ ET BT 26.250 731.762 Td /F1 9.8 Tf [(NMDA receptors containing GluN3A subunits in HD spine loss)] TJ ET 0.267 0.267 0.267 rg BT 294.463 733.269 Td /F4 8.7 Tf [(11)] TJ ET 0.271 0.267 0.267 rg BT 304.100 731.762 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 712.357 Td /F1 9.8 Tf [(In contrast to the reduction in evoked synaptic currents, NMDA responses evoked in the presence of TBOA were increased. )] TJ ET BT 26.250 700.452 Td /F1 9.8 Tf [(Previous studies demonstrated an increase in the extrasynaptic NMDAR-mediated responses in young YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 531.281 701.960 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 536.099 700.452 Td /F1 9.8 Tf [(. )] TJ ET BT 26.250 688.548 Td /F1 9.8 Tf [(However, differences between direct and indirect pathway MSNs were not examined. The present study not only confirms that )] TJ ET BT 26.250 676.643 Td /F1 9.8 Tf [(this increase in extrasynaptic NMDAR-mediated responses occurs in old YAC128 mice, but also demonstrates that it occurs in )] TJ ET BT 26.250 664.738 Td /F1 9.8 Tf [(both direct and indirect pathway MSNs though it appears to be mechanistically due to differing underlying effetcs. Direct )] TJ ET BT 26.250 652.833 Td /F1 9.8 Tf [(pathway MSNs show a larger increase in peak amplitude while indirect pathway MSNs display a larger increase in normalized )] TJ ET BT 26.250 640.929 Td /F1 9.8 Tf [(area due to a longer decay time. This latter effect could suggest that MSNs of the indirect pathway are more sensitive to )] TJ ET BT 26.250 629.024 Td /F1 9.8 Tf [(changes in glutamate transport, thus contributing to their increased vulnerability in HD.)] TJ ET BT 26.250 609.619 Td /F1 9.8 Tf [(NMDARs are composed by GluN2A and GluN2B subunits. GluN2A subunits are thought to be restricted mainly to the synaptic )] TJ ET BT 26.250 597.714 Td /F1 9.8 Tf [(site)] TJ ET 0.267 0.267 0.267 rg BT 41.421 599.222 Td /F4 8.7 Tf [(12)] TJ ET 0.271 0.267 0.267 rg BT 51.058 597.714 Td /F1 9.8 Tf [(, while the GluN2B subunits are located on synaptic and extrasynaptic sites)] TJ ET 0.267 0.267 0.267 rg BT 375.665 599.222 Td /F4 8.7 Tf [(13)] TJ ET 0.271 0.267 0.267 rg BT 385.302 597.714 Td /F1 9.8 Tf [(. It has been suggested that the activation of )] TJ ET BT 26.250 585.810 Td /F1 9.8 Tf [(the synaptic NMDARs is neuroprotective, while activation of extrasynaptic NMDA receptors stimulates a pro-apoptotic cellular )] TJ ET BT 26.250 573.905 Td /F1 9.8 Tf [(pathway)] TJ ET 0.267 0.267 0.267 rg BT 62.559 575.412 Td /F4 8.7 Tf [(14)] TJ ET 0.271 0.267 0.267 rg BT 72.196 573.905 Td /F1 9.8 Tf [(. Alterations in NMDAR subunit trafficking is thought to contribute to increased density of GluN2B subunits at )] TJ ET BT 26.250 562.000 Td /F1 9.8 Tf [(extrasynaptic sites in HD)] TJ ET 0.267 0.267 0.267 rg BT 133.529 563.507 Td /F4 8.7 Tf [(15)] TJ ET 0.271 0.267 0.267 rg BT 143.167 565.888 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 145.576 563.507 Td /F4 8.7 Tf [(16)] TJ ET 0.271 0.267 0.267 rg BT 155.213 562.000 Td /F1 9.8 Tf [(. In a recent study employing co-culture of cortical and striatal MSNs from YAC128 mice, a larger )] TJ ET BT 26.250 550.095 Td /F1 9.8 Tf [(extrasynaptic NMDAR-mediated current occurred in association with an increase in GluN2B subunit expression at the cellular )] TJ ET BT 26.250 538.191 Td /F1 9.8 Tf [(surface compared with the co-culture obtained from WT mice)] TJ ET 0.267 0.267 0.267 rg BT 289.588 539.698 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 294.406 542.079 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 296.816 539.698 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 306.453 538.191 Td /F1 9.8 Tf [(. In addition, low-dose memantine, a blocker of GluN2B-)] TJ ET BT 26.250 526.286 Td /F1 9.8 Tf [(containing NMDARs at extrasynaptic sites, ameliorates neuropathological and behavioral manifestations of HD in YAC128 mice)] TJ ET 0.267 0.267 0.267 rg BT 575.711 527.793 Td /F4 8.7 Tf [(6)] TJ ET 0.271 0.267 0.267 rg BT 26.250 518.269 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 28.659 515.888 Td /F4 8.7 Tf [(18)] TJ ET 0.271 0.267 0.267 rg BT 38.297 514.381 Td /F1 9.8 Tf [(. In co-cultured YAC128 MSNs, low doses of memantine and ifenprodil attenuated extrasynaptic NMDAR-mediated neuronal )] TJ ET BT 26.250 502.476 Td /F1 9.8 Tf [(apoptosis)] TJ ET 0.267 0.267 0.267 rg BT 67.980 503.984 Td /F4 8.7 Tf [(17)] TJ ET 0.271 0.267 0.267 rg BT 77.617 502.476 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 483.072 Td /F1 9.8 Tf [(In conclusion, the present results showed that the glutamatergic inputs to MSNs are decreased in 16-month old YAC128 mice. )] TJ ET BT 26.250 471.167 Td /F1 9.8 Tf [(This study also demonstrates that in late stage HD, NMDA responses evoked in both direct and indirect pathway MSNs in the )] TJ ET BT 26.250 459.262 Td /F1 9.8 Tf [(presence of TBOA are significantly increased in YAC128 mice and the parameters for the increase are different. Direct pathway )] TJ ET BT 26.250 447.357 Td /F1 9.8 Tf [(MSNs in YAC128 mice display increases in peak amplitude while indirect pathway MSNs in YAC128 mice display increases in )] TJ ET BT 26.250 435.453 Td /F1 9.8 Tf [(normalized area due to significantly slower decay times. These results suggest that alterations in glutamate reuptake have )] TJ ET BT 26.250 423.548 Td /F1 9.8 Tf [(different effects on indirect compared with direct pathway MSNs and highlight previous studies showing that MSNs of both )] TJ ET BT 26.250 411.643 Td /F1 9.8 Tf [(pathways are differentially affected in mouse models of HD)] TJ ET 0.267 0.267 0.267 rg BT 280.394 413.150 Td /F4 8.7 Tf [(19)] TJ ET 0.271 0.267 0.267 rg BT 290.031 415.531 Td /F1 8.7 Tf [(,)] TJ ET 0.267 0.267 0.267 rg BT 292.440 413.150 Td /F4 8.7 Tf [(20)] TJ ET 0.271 0.267 0.267 rg BT 302.078 411.643 Td /F1 9.8 Tf [(.)] TJ ET BT 26.250 375.041 Td /F4 12.0 Tf [(Acknowledgements)] TJ ET BT 26.250 355.086 Td /F1 9.8 Tf [(The authors thank Donna Crandall for help with the illustrations.)] TJ ET BT 26.250 325.984 Td /F4 12.0 Tf [(References)] TJ ET BT 26.250 298.530 Td /F1 9.8 Tf [(1.)] TJ ET BT 38.132 298.530 Td /F1 9.8 Tf [(G. Bates, P.S. Harper, L. Jones \(Eds.\), Huntington's disease, Oxford University Press, Oxford ; New York, 2002, xvi, 558 p.)] TJ ET BT 26.250 279.125 Td /F1 9.8 Tf [(2.)] TJ ET BT 38.132 279.125 Td /F1 9.8 Tf [(Slow EJ, van Raamsdonk J, Rogers D, Coleman SH, Graham RK, Deng Y, Oh R, Bissada N, Hossain SM, Yang YZ, Li XJ, )] TJ ET BT 26.250 267.220 Td /F1 9.8 Tf [(Simpson EM, Gutekunst CA, Leavitt BR, Hayden MR. Selective striatal neuronal loss in a YAC128 mouse model of Huntington )] TJ ET BT 26.250 255.315 Td /F1 9.8 Tf [(disease. Hum Mol Genet. 2003 Jul 1;12\(13\):1555-67. PubMed PMID:12812983.)] TJ ET BT 26.250 235.911 Td /F1 9.8 Tf [(3.)] TJ ET BT 38.132 235.911 Td /F1 9.8 Tf [(Ferrante RJ. Mouse models of Huntington's disease and methodological considerations for therapeutic trials. Biochim )] TJ ET BT 26.250 224.006 Td /F1 9.8 Tf [(Biophys Acta. 2009 Jun;1792\(6\):506-20. PubMed PMID:19362590.)] TJ ET BT 26.250 204.601 Td /F1 9.8 Tf [(4.)] TJ ET BT 38.132 204.601 Td /F1 9.8 Tf [(Andr VM, Cepeda C, Fisher YE, Huynh M, Bardakjian N, Singh S, Yang XW, Levine MS. Differential electrophysiological )] TJ ET BT 26.250 192.696 Td /F1 9.8 Tf [(changes in striatal output neurons in Huntington's disease. J Neurosci. 2011 Jan 26;31\(4\):1170-82. PubMed PMID:21273402.)] TJ ET BT 26.250 173.292 Td /F1 9.8 Tf [(5.)] TJ ET BT 38.132 173.292 Td /F1 9.8 Tf [(Andr VM, Fisher YE, Levine MS. Altered Balance of Activity in the Striatal Direct and Indirect Pathways in Mouse Models of )] TJ ET BT 26.250 161.387 Td /F1 9.8 Tf [(Huntington's Disease. Front Syst Neurosci. 2011;5:46. PubMed PMID:21720523.)] TJ ET BT 26.250 141.982 Td /F1 9.8 Tf [(6.)] TJ ET BT 38.132 141.982 Td /F1 9.8 Tf [(Milnerwood AJ, Gladding CM, Pouladi MA, Kaufman AM, Hines RM, Boyd JD, Ko RW, Vasuta OC, Graham RK, Hayden MR, )] TJ ET BT 26.250 130.077 Td /F1 9.8 Tf [(Murphy TH, Raymond LA. Early increase in extrasynaptic NMDA receptor signaling and expression contributes to phenotype )] TJ ET BT 26.250 118.173 Td /F1 9.8 Tf [(onset in Huntington's disease mice. Neuron. 2010 Jan 28;65\(2\):178-90. PubMed PMID:20152125.)] TJ ET BT 26.250 98.768 Td /F1 9.8 Tf [(7.)] TJ ET BT 38.132 98.768 Td /F1 9.8 Tf [(Cepeda C, Hurst RS, Calvert CR, Hernndez-Echeagaray E, Nguyen OK, Jocoy E, Christian LJ, Ariano MA, Levine MS. )] TJ ET BT 26.250 86.863 Td /F1 9.8 Tf [(Transient and progressive electrophysiological alterations in the corticostriatal pathway in a mouse model of Huntington's )] TJ ET BT 26.250 74.958 Td /F1 9.8 Tf [(disease. J Neurosci. 2003 Feb 1;23\(3\):961-9. PubMed PMID:12574425.)] TJ ET BT 26.250 55.554 Td /F1 9.8 Tf [(8.)] TJ ET BT 38.132 55.554 Td /F1 9.8 Tf [(Cummings DM, Cepeda C, Levine MS. Alterations in striatal synaptic transmission are consistent across genetic mouse )] TJ ET BT 26.250 43.649 Td /F1 9.8 Tf [(models of Huntington's disease. ASN Neuro. 2010 Jun 18;2\(3\):e00036. PubMed PMID:20585470.)] TJ ET Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(6)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Huntington Disease)] TJ ET Q endstream endobj 259 0 obj << /Type /Annot /Subtype /Link /A 260 0 R /Border [0 0 0] /H /I /Rect [ 420.2183 756.2769 429.8556 765.0952 ] >> endobj 260 0 obj << /Type /Action >> endobj 261 0 obj << /Type /Annot /Subtype /Link /A 262 0 R /Border [0 0 0] /H /I /Rect [ 294.4628 732.4674 304.1001 741.2858 ] >> endobj 262 0 obj << /Type /Action >> endobj 263 0 obj << /Type /Annot /Subtype /Link /A 264 0 R /Border [0 0 0] /H /I /Rect [ 531.2805 701.1579 536.0992 709.9762 ] >> endobj 264 0 obj << /Type /Action >> endobj 265 0 obj << /Type /Annot /Subtype /Link /A 266 0 R /Border [0 0 0] /H /I /Rect [ 41.4210 598.4199 51.0583 607.2382 ] >> endobj 266 0 obj << /Type /Action >> endobj 267 0 obj << /Type /Annot /Subtype /Link /A 268 0 R /Border [0 0 0] /H /I /Rect [ 375.6651 598.4199 385.3024 607.2382 ] >> endobj 268 0 obj 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C, Joyce JA, Carroll JB, Leavitt BR, Hayden MR, Levine MS, Bamford )] TJ ET BT 26.250 748.071 Td /F1 9.8 Tf [(NS. Age-dependent alterations of corticostriatal activity in the YAC128 mouse model of Huntington disease. J Neurosci. 2009 )] TJ ET BT 26.250 736.167 Td /F1 9.8 Tf [(Feb 25;29\(8\):2414-27. PubMed PMID:19244517.)] TJ ET BT 26.250 716.762 Td /F1 9.8 Tf [(10.)] TJ ET BT 43.553 716.762 Td /F1 9.8 Tf [(Graham RK, Pouladi MA, Joshi P, Lu G, Deng Y, Wu NP, Figueroa BE, Metzler M, Andr VM, Slow EJ, Raymond L, )] TJ ET BT 26.250 704.857 Td /F1 9.8 Tf [(Friedlander R, Levine MS, Leavitt BR, Hayden MR. Differential susceptibility to excitotoxic stress in YAC128 mouse models of )] TJ ET BT 26.250 692.952 Td /F1 9.8 Tf [(Huntington disease between initiation and progression of disease. J Neurosci. 2009 Feb 18;29\(7\):2193-204. PubMed )] TJ ET BT 26.250 681.048 Td /F1 9.8 Tf [(PMID:19228972.)] TJ ET BT 26.250 661.643 Td /F1 9.8 Tf [(11.)] TJ ET BT 43.553 661.643 Td /F1 9.8 Tf [(Marco S, Giralt A, Petrovic MM, Pouladi MA, Martnez-Turrillas R, Martnez-Hernndez J, Kaltenbach LS, Torres-Peraza J, )] TJ ET BT 26.250 649.738 Td /F1 9.8 Tf [(Graham RK, Watanabe M, Lujn R, Nakanishi N, Lipton SA, Lo DC, Hayden MR, Alberch J, Wesseling JF, Prez-Otao I. )] TJ ET BT 26.250 637.833 Td /F1 9.8 Tf [(Suppressing aberrant GluN3A expression rescues synaptic and behavioral impairments in Huntington's disease models. Nat )] TJ ET BT 26.250 625.929 Td /F1 9.8 Tf [(Med. 2013 Aug;19\(8\):1030-8. PubMed PMID:23852340.)] TJ ET BT 26.250 606.524 Td /F1 9.8 Tf [(12.)] TJ ET BT 43.553 606.524 Td /F1 9.8 Tf [(Thomas CG, Miller AJ, Westbrook GL. Synaptic and extrasynaptic NMDA receptor NR2 subunits in cultured hippocampal )] TJ ET BT 26.250 594.619 Td /F1 9.8 Tf [(neurons. J Neurophysiol. 2006 Mar;95\(3\):1727-34. PubMed PMID:16319212.)] TJ ET BT 26.250 575.214 Td /F1 9.8 Tf [(13.)] TJ ET BT 43.553 575.214 Td /F1 9.8 Tf [(Cull-Candy S, Brickley S, Farrant M. NMDA receptor subunits: diversity, development and disease. Curr Opin Neurobiol. )] TJ ET BT 26.250 563.310 Td /F1 9.8 Tf [(2001 Jun;11\(3\):327-35. PubMed PMID:11399431.)] TJ ET BT 26.250 543.905 Td /F1 9.8 Tf [(14.)] TJ ET BT 43.553 543.905 Td /F1 9.8 Tf [(Hardingham GE, Bading H. Synaptic versus extrasynaptic NMDA receptor signalling: implications for neurodegenerative )] TJ ET BT 26.250 532.000 Td /F1 9.8 Tf [(disorders. Nat Rev Neurosci. 2010 Oct;11\(10\):682-96. PubMed PMID:20842175.)] TJ ET BT 26.250 512.595 Td /F1 9.8 Tf [(15.)] TJ ET BT 43.553 512.595 Td /F1 9.8 Tf [(Fan MM, Raymond LA. N-methyl-D-aspartate \(NMDA\) receptor function and excitotoxicity in Huntington's disease. Prog )] TJ ET BT 26.250 500.691 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):272-93. PubMed PMID:17188796.)] TJ ET BT 26.250 481.286 Td /F1 9.8 Tf [(16.)] TJ ET BT 43.553 481.286 Td /F1 9.8 Tf [(Levine MS, Cepeda C, Andr VM. Location, location, location: contrasting roles of synaptic and extrasynaptic NMDA )] TJ ET BT 26.250 469.381 Td /F1 9.8 Tf [(receptors in Huntington's disease. Neuron. 2010 Jan 28;65\(2\):145-7. PubMed PMID:20152121.)] TJ ET BT 26.250 449.976 Td /F1 9.8 Tf [(17.)] TJ ET BT 43.553 449.976 Td /F1 9.8 Tf [(Milnerwood AJ, Kaufman AM, Sepers MD, Gladding CM, Zhang L, Wang L, Fan J, Coquinco A, Qiao JY, Lee H, Wang YT, )] TJ ET BT 26.250 438.072 Td /F1 9.8 Tf [(Cynader M, Raymond LA. Mitigation of augmented extrasynaptic NMDAR signaling and apoptosis in cortico-striatal co-cultures )] TJ ET BT 26.250 426.167 Td /F1 9.8 Tf [(from Huntington's disease mice. Neurobiol Dis. 2012 Oct;48\(1\):40-51. PubMed PMID:22668780.)] TJ ET BT 26.250 406.762 Td /F1 9.8 Tf [(18.)] TJ ET BT 43.553 406.762 Td /F1 9.8 Tf [(Okamoto S, Pouladi MA, Talantova M, Yao D, Xia P, Ehrnhoefer DE, Zaidi R, Clemente A, Kaul M, Graham RK, Zhang D, )] TJ ET BT 26.250 394.857 Td /F1 9.8 Tf [(Vincent Chen HS, Tong G, Hayden MR, Lipton SA. Balance between synaptic versus extrasynaptic NMDA receptor activity )] TJ ET BT 26.250 382.953 Td /F1 9.8 Tf [(influences inclusions and neurotoxicity of mutant huntingtin. Nat Med. 2009 Dec;15\(12\):1407-13. PubMed PMID:19915593.)] TJ ET BT 26.250 363.548 Td /F1 9.8 Tf [(19.)] TJ ET BT 43.553 363.548 Td /F1 9.8 Tf [(Albin RL, Reiner A, Anderson KD, Dure LS 4th, Handelin B, Balfour R, Whetsell WO Jr, Penney JB, Young AB. Preferential )] TJ ET BT 26.250 351.643 Td /F1 9.8 Tf [(loss of striato-external pallidal projection neurons in presymptomatic Huntington's disease. Ann Neurol. 1992 Apr;31\(4\):425-30. )] TJ ET BT 26.250 339.738 Td /F1 9.8 Tf [(PubMed PMID:1375014.)] TJ ET BT 26.250 320.334 Td /F1 9.8 Tf [(20.)] TJ ET BT 43.553 320.334 Td /F1 9.8 Tf [(Reiner A, Albin RL, Anderson KD, D'Amato CJ, Penney JB, Young AB. Differential loss of striatal projection neurons in )] TJ ET BT 26.250 308.429 Td /F1 9.8 Tf [(Huntington disease. Proc Natl Acad Sci U S A. 1988 Aug;85\(15\):5733-7. PubMed PMID:2456581.)] TJ ET BT 26.250 289.024 Td /F1 9.8 Tf [(21.)] TJ ET BT 43.553 289.024 Td /F1 9.8 Tf [(Cepeda C, Wu N, Andr VM, Cummings DM, Levine MS. The corticostriatal pathway in Huntington's disease. Prog )] TJ ET BT 26.250 277.119 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):253-71. PubMed PMID:17169479.)] TJ ET BT 26.250 257.715 Td /F1 9.8 Tf [(22.)] TJ ET BT 43.553 257.715 Td /F1 9.8 Tf [(Jabaudon D, Shimamoto K, Yasuda-Kamatani Y, Scanziani M, Ghwiler BH, Gerber U. Inhibition of uptake unmasks rapid )] TJ ET BT 26.250 245.810 Td /F1 9.8 Tf [(extracellular turnover of glutamate of nonvesicular origin. Proc Natl Acad Sci U S A. 1999 Jul 20;96\(15\):8733-8. PubMed )] TJ ET BT 26.250 233.905 Td /F1 9.8 Tf [(PMID:10411944.)] TJ ET Q q 15.000 216.524 577.500 560.476 re W n 0.271 0.267 0.267 rg BT 26.250 759.976 Td /F1 9.8 Tf [(9.)] TJ ET BT 38.132 759.976 Td /F1 9.8 Tf [(Joshi PR, Wu NP, Andr VM, Cummings DM, Cepeda C, Joyce JA, Carroll JB, Leavitt BR, Hayden MR, Levine MS, Bamford )] TJ ET BT 26.250 748.071 Td /F1 9.8 Tf [(NS. Age-dependent alterations of corticostriatal activity in the YAC128 mouse model of Huntington disease. J Neurosci. 2009 )] TJ ET BT 26.250 736.167 Td /F1 9.8 Tf [(Feb 25;29\(8\):2414-27. PubMed PMID:19244517.)] TJ ET BT 26.250 716.762 Td /F1 9.8 Tf [(10.)] TJ ET BT 43.553 716.762 Td /F1 9.8 Tf [(Graham RK, Pouladi MA, Joshi P, Lu G, Deng Y, Wu NP, Figueroa BE, Metzler M, Andr VM, Slow EJ, Raymond L, )] TJ ET BT 26.250 704.857 Td /F1 9.8 Tf [(Friedlander R, Levine MS, Leavitt BR, Hayden MR. Differential susceptibility to excitotoxic stress in YAC128 mouse models of )] TJ ET BT 26.250 692.952 Td /F1 9.8 Tf [(Huntington disease between initiation and progression of disease. J Neurosci. 2009 Feb 18;29\(7\):2193-204. PubMed )] TJ ET BT 26.250 681.048 Td /F1 9.8 Tf [(PMID:19228972.)] TJ ET BT 26.250 661.643 Td /F1 9.8 Tf [(11.)] TJ ET BT 43.553 661.643 Td /F1 9.8 Tf [(Marco S, Giralt A, Petrovic MM, Pouladi MA, Martnez-Turrillas R, Martnez-Hernndez J, Kaltenbach LS, Torres-Peraza J, )] TJ ET BT 26.250 649.738 Td /F1 9.8 Tf [(Graham RK, Watanabe M, Lujn R, Nakanishi N, Lipton SA, Lo DC, Hayden MR, Alberch J, Wesseling JF, Prez-Otao I. )] TJ ET BT 26.250 637.833 Td /F1 9.8 Tf [(Suppressing aberrant GluN3A expression rescues synaptic and behavioral impairments in Huntington's disease models. Nat )] TJ ET BT 26.250 625.929 Td /F1 9.8 Tf [(Med. 2013 Aug;19\(8\):1030-8. PubMed PMID:23852340.)] TJ ET BT 26.250 606.524 Td /F1 9.8 Tf [(12.)] TJ ET BT 43.553 606.524 Td /F1 9.8 Tf [(Thomas CG, Miller AJ, Westbrook GL. Synaptic and extrasynaptic NMDA receptor NR2 subunits in cultured hippocampal )] TJ ET BT 26.250 594.619 Td /F1 9.8 Tf [(neurons. J Neurophysiol. 2006 Mar;95\(3\):1727-34. PubMed PMID:16319212.)] TJ ET BT 26.250 575.214 Td /F1 9.8 Tf [(13.)] TJ ET BT 43.553 575.214 Td /F1 9.8 Tf [(Cull-Candy S, Brickley S, Farrant M. NMDA receptor subunits: diversity, development and disease. Curr Opin Neurobiol. )] TJ ET BT 26.250 563.310 Td /F1 9.8 Tf [(2001 Jun;11\(3\):327-35. PubMed PMID:11399431.)] TJ ET BT 26.250 543.905 Td /F1 9.8 Tf [(14.)] TJ ET BT 43.553 543.905 Td /F1 9.8 Tf [(Hardingham GE, Bading H. Synaptic versus extrasynaptic NMDA receptor signalling: implications for neurodegenerative )] TJ ET BT 26.250 532.000 Td /F1 9.8 Tf [(disorders. Nat Rev Neurosci. 2010 Oct;11\(10\):682-96. PubMed PMID:20842175.)] TJ ET BT 26.250 512.595 Td /F1 9.8 Tf [(15.)] TJ ET BT 43.553 512.595 Td /F1 9.8 Tf [(Fan MM, Raymond LA. N-methyl-D-aspartate \(NMDA\) receptor function and excitotoxicity in Huntington's disease. Prog )] TJ ET BT 26.250 500.691 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):272-93. PubMed PMID:17188796.)] TJ ET BT 26.250 481.286 Td /F1 9.8 Tf [(16.)] TJ ET BT 43.553 481.286 Td /F1 9.8 Tf [(Levine MS, Cepeda C, Andr VM. Location, location, location: contrasting roles of synaptic and extrasynaptic NMDA )] TJ ET BT 26.250 469.381 Td /F1 9.8 Tf [(receptors in Huntington's disease. Neuron. 2010 Jan 28;65\(2\):145-7. PubMed PMID:20152121.)] TJ ET BT 26.250 449.976 Td /F1 9.8 Tf [(17.)] TJ ET BT 43.553 449.976 Td /F1 9.8 Tf [(Milnerwood AJ, Kaufman AM, Sepers MD, Gladding CM, Zhang L, Wang L, Fan J, Coquinco A, Qiao JY, Lee H, Wang YT, )] TJ ET BT 26.250 438.072 Td /F1 9.8 Tf [(Cynader M, Raymond LA. Mitigation of augmented extrasynaptic NMDAR signaling and apoptosis in cortico-striatal co-cultures )] TJ ET BT 26.250 426.167 Td /F1 9.8 Tf [(from Huntington's disease mice. Neurobiol Dis. 2012 Oct;48\(1\):40-51. PubMed PMID:22668780.)] TJ ET BT 26.250 406.762 Td /F1 9.8 Tf [(18.)] TJ ET BT 43.553 406.762 Td /F1 9.8 Tf [(Okamoto S, Pouladi MA, Talantova M, Yao D, Xia P, Ehrnhoefer DE, Zaidi R, Clemente A, Kaul M, Graham RK, Zhang D, )] TJ ET BT 26.250 394.857 Td /F1 9.8 Tf [(Vincent Chen HS, Tong G, Hayden MR, Lipton SA. Balance between synaptic versus extrasynaptic NMDA receptor activity )] TJ ET BT 26.250 382.953 Td /F1 9.8 Tf [(influences inclusions and neurotoxicity of mutant huntingtin. Nat Med. 2009 Dec;15\(12\):1407-13. PubMed PMID:19915593.)] TJ ET BT 26.250 363.548 Td /F1 9.8 Tf [(19.)] TJ ET BT 43.553 363.548 Td /F1 9.8 Tf [(Albin RL, Reiner A, Anderson KD, Dure LS 4th, Handelin B, Balfour R, Whetsell WO Jr, Penney JB, Young AB. Preferential )] TJ ET BT 26.250 351.643 Td /F1 9.8 Tf [(loss of striato-external pallidal projection neurons in presymptomatic Huntington's disease. Ann Neurol. 1992 Apr;31\(4\):425-30. )] TJ ET BT 26.250 339.738 Td /F1 9.8 Tf [(PubMed PMID:1375014.)] TJ ET BT 26.250 320.334 Td /F1 9.8 Tf [(20.)] TJ ET BT 43.553 320.334 Td /F1 9.8 Tf [(Reiner A, Albin RL, Anderson KD, D'Amato CJ, Penney JB, Young AB. Differential loss of striatal projection neurons in )] TJ ET BT 26.250 308.429 Td /F1 9.8 Tf [(Huntington disease. Proc Natl Acad Sci U S A. 1988 Aug;85\(15\):5733-7. PubMed PMID:2456581.)] TJ ET BT 26.250 289.024 Td /F1 9.8 Tf [(21.)] TJ ET BT 43.553 289.024 Td /F1 9.8 Tf [(Cepeda C, Wu N, Andr VM, Cummings DM, Levine MS. The corticostriatal pathway in Huntington's disease. Prog )] TJ ET BT 26.250 277.119 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):253-71. PubMed PMID:17169479.)] TJ ET BT 26.250 257.715 Td /F1 9.8 Tf [(22.)] TJ ET BT 43.553 257.715 Td /F1 9.8 Tf [(Jabaudon D, Shimamoto K, Yasuda-Kamatani Y, Scanziani M, Ghwiler BH, Gerber U. Inhibition of uptake unmasks rapid )] TJ ET BT 26.250 245.810 Td /F1 9.8 Tf [(extracellular turnover of glutamate of nonvesicular origin. Proc Natl Acad Sci U S A. 1999 Jul 20;96\(15\):8733-8. PubMed )] TJ ET BT 26.250 233.905 Td /F1 9.8 Tf [(PMID:10411944.)] TJ ET Q q 15.000 216.524 577.500 560.476 re W n 0.271 0.267 0.267 rg BT 26.250 759.976 Td /F1 9.8 Tf [(9.)] TJ ET BT 38.132 759.976 Td /F1 9.8 Tf [(Joshi PR, Wu NP, Andr VM, Cummings DM, Cepeda C, Joyce JA, Carroll JB, Leavitt BR, Hayden MR, Levine MS, Bamford )] TJ ET BT 26.250 748.071 Td /F1 9.8 Tf [(NS. Age-dependent alterations of corticostriatal activity in the YAC128 mouse model of Huntington disease. J Neurosci. 2009 )] TJ ET BT 26.250 736.167 Td /F1 9.8 Tf [(Feb 25;29\(8\):2414-27. PubMed PMID:19244517.)] TJ ET BT 26.250 716.762 Td /F1 9.8 Tf [(10.)] TJ ET BT 43.553 716.762 Td /F1 9.8 Tf [(Graham RK, Pouladi MA, Joshi P, Lu G, Deng Y, Wu NP, Figueroa BE, Metzler M, Andr VM, Slow EJ, Raymond L, )] TJ ET BT 26.250 704.857 Td /F1 9.8 Tf [(Friedlander R, Levine MS, Leavitt BR, Hayden MR. Differential susceptibility to excitotoxic stress in YAC128 mouse models of )] TJ ET BT 26.250 692.952 Td /F1 9.8 Tf [(Huntington disease between initiation and progression of disease. J Neurosci. 2009 Feb 18;29\(7\):2193-204. PubMed )] TJ ET BT 26.250 681.048 Td /F1 9.8 Tf [(PMID:19228972.)] TJ ET BT 26.250 661.643 Td /F1 9.8 Tf [(11.)] TJ ET BT 43.553 661.643 Td /F1 9.8 Tf [(Marco S, Giralt A, Petrovic MM, Pouladi MA, Martnez-Turrillas R, Martnez-Hernndez J, Kaltenbach LS, Torres-Peraza J, )] TJ ET BT 26.250 649.738 Td /F1 9.8 Tf [(Graham RK, Watanabe M, Lujn R, Nakanishi N, Lipton SA, Lo DC, Hayden MR, Alberch J, Wesseling JF, Prez-Otao I. )] TJ ET BT 26.250 637.833 Td /F1 9.8 Tf [(Suppressing aberrant GluN3A expression rescues synaptic and behavioral impairments in Huntington's disease models. Nat )] TJ ET BT 26.250 625.929 Td /F1 9.8 Tf [(Med. 2013 Aug;19\(8\):1030-8. PubMed PMID:23852340.)] TJ ET BT 26.250 606.524 Td /F1 9.8 Tf [(12.)] TJ ET BT 43.553 606.524 Td /F1 9.8 Tf [(Thomas CG, Miller AJ, Westbrook GL. Synaptic and extrasynaptic NMDA receptor NR2 subunits in cultured hippocampal )] TJ ET BT 26.250 594.619 Td /F1 9.8 Tf [(neurons. J Neurophysiol. 2006 Mar;95\(3\):1727-34. PubMed PMID:16319212.)] TJ ET BT 26.250 575.214 Td /F1 9.8 Tf [(13.)] TJ ET BT 43.553 575.214 Td /F1 9.8 Tf [(Cull-Candy S, Brickley S, Farrant M. NMDA receptor subunits: diversity, development and disease. Curr Opin Neurobiol. )] TJ ET BT 26.250 563.310 Td /F1 9.8 Tf [(2001 Jun;11\(3\):327-35. PubMed PMID:11399431.)] TJ ET BT 26.250 543.905 Td /F1 9.8 Tf [(14.)] TJ ET BT 43.553 543.905 Td /F1 9.8 Tf [(Hardingham GE, Bading H. Synaptic versus extrasynaptic NMDA receptor signalling: implications for neurodegenerative )] TJ ET BT 26.250 532.000 Td /F1 9.8 Tf [(disorders. Nat Rev Neurosci. 2010 Oct;11\(10\):682-96. PubMed PMID:20842175.)] TJ ET BT 26.250 512.595 Td /F1 9.8 Tf [(15.)] TJ ET BT 43.553 512.595 Td /F1 9.8 Tf [(Fan MM, Raymond LA. N-methyl-D-aspartate \(NMDA\) receptor function and excitotoxicity in Huntington's disease. Prog )] TJ ET BT 26.250 500.691 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):272-93. PubMed PMID:17188796.)] TJ ET BT 26.250 481.286 Td /F1 9.8 Tf [(16.)] TJ ET BT 43.553 481.286 Td /F1 9.8 Tf [(Levine MS, Cepeda C, Andr VM. Location, location, location: contrasting roles of synaptic and extrasynaptic NMDA )] TJ ET BT 26.250 469.381 Td /F1 9.8 Tf [(receptors in Huntington's disease. Neuron. 2010 Jan 28;65\(2\):145-7. PubMed PMID:20152121.)] TJ ET BT 26.250 449.976 Td /F1 9.8 Tf [(17.)] TJ ET BT 43.553 449.976 Td /F1 9.8 Tf [(Milnerwood AJ, Kaufman AM, Sepers MD, Gladding CM, Zhang L, Wang L, Fan J, Coquinco A, Qiao JY, Lee H, Wang YT, )] TJ ET BT 26.250 438.072 Td /F1 9.8 Tf [(Cynader M, Raymond LA. Mitigation of augmented extrasynaptic NMDAR signaling and apoptosis in cortico-striatal co-cultures )] TJ ET BT 26.250 426.167 Td /F1 9.8 Tf [(from Huntington's disease mice. Neurobiol Dis. 2012 Oct;48\(1\):40-51. PubMed PMID:22668780.)] TJ ET BT 26.250 406.762 Td /F1 9.8 Tf [(18.)] TJ ET BT 43.553 406.762 Td /F1 9.8 Tf [(Okamoto S, Pouladi MA, Talantova M, Yao D, Xia P, Ehrnhoefer DE, Zaidi R, Clemente A, Kaul M, Graham RK, Zhang D, )] TJ ET BT 26.250 394.857 Td /F1 9.8 Tf [(Vincent Chen HS, Tong G, Hayden MR, Lipton SA. Balance between synaptic versus extrasynaptic NMDA receptor activity )] TJ ET BT 26.250 382.953 Td /F1 9.8 Tf [(influences inclusions and neurotoxicity of mutant huntingtin. Nat Med. 2009 Dec;15\(12\):1407-13. PubMed PMID:19915593.)] TJ ET BT 26.250 363.548 Td /F1 9.8 Tf [(19.)] TJ ET BT 43.553 363.548 Td /F1 9.8 Tf [(Albin RL, Reiner A, Anderson KD, Dure LS 4th, Handelin B, Balfour R, Whetsell WO Jr, Penney JB, Young AB. Preferential )] TJ ET BT 26.250 351.643 Td /F1 9.8 Tf [(loss of striato-external pallidal projection neurons in presymptomatic Huntington's disease. Ann Neurol. 1992 Apr;31\(4\):425-30. )] TJ ET BT 26.250 339.738 Td /F1 9.8 Tf [(PubMed PMID:1375014.)] TJ ET BT 26.250 320.334 Td /F1 9.8 Tf [(20.)] TJ ET BT 43.553 320.334 Td /F1 9.8 Tf [(Reiner A, Albin RL, Anderson KD, D'Amato CJ, Penney JB, Young AB. Differential loss of striatal projection neurons in )] TJ ET BT 26.250 308.429 Td /F1 9.8 Tf [(Huntington disease. Proc Natl Acad Sci U S A. 1988 Aug;85\(15\):5733-7. PubMed PMID:2456581.)] TJ ET BT 26.250 289.024 Td /F1 9.8 Tf [(21.)] TJ ET BT 43.553 289.024 Td /F1 9.8 Tf [(Cepeda C, Wu N, Andr VM, Cummings DM, Levine MS. The corticostriatal pathway in Huntington's disease. Prog )] TJ ET BT 26.250 277.119 Td /F1 9.8 Tf [(Neurobiol. 2007 Apr;81\(5-6\):253-71. PubMed PMID:17169479.)] TJ ET BT 26.250 257.715 Td /F1 9.8 Tf [(22.)] TJ ET BT 43.553 257.715 Td /F1 9.8 Tf [(Jabaudon D, Shimamoto K, Yasuda-Kamatani Y, Scanziani M, Ghwiler BH, Gerber U. Inhibition of uptake unmasks rapid )] TJ ET BT 26.250 245.810 Td /F1 9.8 Tf [(extracellular turnover of glutamate of nonvesicular origin. Proc Natl Acad Sci U S A. 1999 Jul 20;96\(15\):8733-8. PubMed )] TJ ET BT 26.250 233.905 Td /F1 9.8 Tf [(PMID:10411944.)] TJ ET Q q 0.000 0.000 0.000 rg BT 291.710 19.825 Td /F1 11.0 Tf [(7)] TJ ET BT 25.000 19.825 Td /F1 11.0 Tf [(PLOS Currents Huntington Disease)] TJ ET Q endstream endobj xref 0 351 0000000000 65535 f 0000000008 00000 n 0000000073 00000 n 0000000119 00000 n 0000000708 00000 n 0000000745 00000 n 0000001054 00000 n 0000001542 00000 n 0000027991 00000 n 0000028098 00000 n 0000028206 00000 n 0000028317 00000 n 0000028430 00000 n 0000028818 00000 n 0000033299 00000 n 0000033426 00000 n 0000033664 00000 n 0000033791 00000 n 0000034029 00000 n 0000034156 00000 n 0000034394 00000 n 0000034521 00000 n 0000034628 00000 n 0000034754 00000 n 0000034853 00000 n 0000034980 00000 n 0000035081 00000 n 0000035209 00000 n 0000035305 00000 n 0000035433 00000 n 0000035532 00000 n 0000035660 00000 n 0000035769 00000 n 0000035897 00000 n 0000035997 00000 n 0000036125 00000 n 0000036223 00000 n 0000036351 00000 n 0000036600 00000 n 0000036728 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