鲫鱼视网膜水平细胞谷氨酸受体对γ-氨基丁酸转运体的调控
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摘要
水平细胞是视网膜内核层的抑制性中间神经元,接受来自光感受器细胞的谷氨酸能输入,同时通过GABA转运体释放GABA,参与水平细胞对光感受器细胞的反馈调节,及双极细胞和神经节细胞的中心外周拮抗的感受野的形成。谷氨酸可以激活水平细胞膜上的包括NMDA受体和AMPA受体在内的谷氨酸受体。已有报道证明激活鳐鱼视网膜水平细胞的AMPA受体可以抑制GABA转运体电流。我们应用全细胞膜片钳技术,在鲫鱼视网膜上也发现了类似的现象,激活鲫鱼视网膜水平细胞的AMPA受体也可以抑制GABA转运体电流。由于NMDA受体是最近在鲫鱼水平细胞上发现的,其功能尚不清楚。我们因此对激活NMDA受体是否能调节GABA转运体电流进行了研究。实验结果显示,激活水平细胞的NMDA受体可以抑制GABA转运体电流。NMDA受体的阻断剂AP-5(100μM)可以消除这个效应。说明这种效应是通过激活NMDA受体来实现的。当使用无钙的Ringer’s液灌流的时候,这种NMDA对GABA转运体的抑制就会被解除。当胞内钙库的钙释放和回收被20μM ryanodine + 2μM thapsigargin灌流3 min所阻断后,NMDA对GABA转运体的抑制作用也会被减弱。在全细胞模式膜片钳形成的时候,电极内液中10 mM的BAPTA经5 min扩散到细胞内后,这种NMDA的抑制效应也被削弱。以上结果说明激活NMDA受体可以通过影响胞内钙过程来调节GABA转运体的活动。进一步地,基于本实验室以前的工作,用数学方程构建了一个含有内质网膜和细胞膜的细胞模型来定量地分析NMDA引起的Ca2+动力学及GABA转运体电流的变化。模型结果提示,胞内钙库活动参与NMDA受体对GABA转运体活动的调节。
除了鲫鱼,在大鼠和人的视网膜水平细胞上也存在NMDA受体,但是水平细胞上NMDA受体的生理功能尚不清楚。本文在鲫鱼视网膜水平细胞上发现激活NMDA受体可以抑制GABA转运体的活动。在生理条件下,视网膜光感受器细胞在暗中持续释放谷氨酸,水平细胞上的钙通透的AMPA受体和NMDA受体均会被激活,胞外钙离子就会通过这两种受体进入胞内,发挥抑制GABA转运体的作用。相对于AMPA受体的快速失敏,NMDA受体的失敏很慢。提示生理条件下NMDA受体在对GABA转运体的抑制性调控可能发挥更重要的作用。
Horizontal cells are interneurons in the outer retina, which receive glutamate input from photoreceptors, and release GABA via GABA transporters. Horizontal cells are responsible for the formation of the antagonistic surround receptive field property of the retinal bipolar cells and ganglion cells. Glutamate can activate glutamate receptors on horizontal cells, including AMPA receptors and NMDA receptors. It has been reported that activation of AMPA receptors inhibits GABA transporter currents on skate retinal horizontal cells. We have observed similar inhibition on carp horizontal cells using patch-clamp technique. Recently, it was reported that functional NMDA receptors are expressed in carp retinal horizontal cells, but the relevant functions of NMDA receptors in the horizontal cells are not fully understood. We consequently explored whether activation of NMDA receptors on horizontal cells could also alter the activity of GABA transport in these cells. Our results show that activation of NMDA receptors inhibited GABA transporter currents on carp retinal horizontal cells. Application of AP5 (100μM) eliminated the inhibition, which indicates NMDA receptors are necessary. When the extracellular Ca2+ was removed from the Ringer's solution, the NMDA inhibitory effect on the GABA transporter current was eliminated. The suppression effect could be attenuated when the Ca2+ release and Ca2+ uptake of intracellular Ca2+ store were blocked after the cell had been pre-incubated with 20μM ryanodine plus 2μM thapsigargin for 3 min. After achieving the whole-cell configuration, the retinal horizontal cell was loaded with 10 mM BAPTA by diffusion from the patch pipette for 5 min, and then the NMDA modulation of GABA transporters was suppressed. These results suggest that the activation of NMDA receptors inhibits GABA transporter-mediated current by affecting Ca2+ processes in the retinal horizontal cells. Furthermore, a model containing endoplasmic reticulum (ER) membrane processes and plasma membrane processes was constructed for quantitative analyses of the NMDA-triggered Ca2+ dynamics and its modulation of GABA transporters. The model results suggest that the intracellular Ca2+ store is involved in the NMDA modulation of GABA transporters.
It is reported that NMDA receptors are expressed in retinal horizontal cells of carp, rat and human, but physiological function of the NMDA receptors is not clear. In present study, our results show that activation of NMDA receptors in carp retinal horizontal cells inhibit GABA transporter current. Under physiological conditions, photoreceptors release glutamate continually in the darkness. AMPA receptors and NMDA receptors on the retinal horizontal cells are activated, and extracellular calcium influx can occur through these glutamate receptors. Since AMPA receptors display very rapid and significant desensitization, and NMDA receptors do not desensitize as quickly or as fully, NMDA receptors might play a significant role in the regulation of GABA transporter activities under physiological conditions.
除了鲫鱼,在大鼠和人的视网膜水平细胞上也存在NMDA受体,但是水平细胞上NMDA受体的生理功能尚不清楚。本文在鲫鱼视网膜水平细胞上发现激活NMDA受体可以抑制GABA转运体的活动。在生理条件下,视网膜光感受器细胞在暗中持续释放谷氨酸,水平细胞上的钙通透的AMPA受体和NMDA受体均会被激活,胞外钙离子就会通过这两种受体进入胞内,发挥抑制GABA转运体的作用。相对于AMPA受体的快速失敏,NMDA受体的失敏很慢。提示生理条件下NMDA受体在对GABA转运体的抑制性调控可能发挥更重要的作用。
Horizontal cells are interneurons in the outer retina, which receive glutamate input from photoreceptors, and release GABA via GABA transporters. Horizontal cells are responsible for the formation of the antagonistic surround receptive field property of the retinal bipolar cells and ganglion cells. Glutamate can activate glutamate receptors on horizontal cells, including AMPA receptors and NMDA receptors. It has been reported that activation of AMPA receptors inhibits GABA transporter currents on skate retinal horizontal cells. We have observed similar inhibition on carp horizontal cells using patch-clamp technique. Recently, it was reported that functional NMDA receptors are expressed in carp retinal horizontal cells, but the relevant functions of NMDA receptors in the horizontal cells are not fully understood. We consequently explored whether activation of NMDA receptors on horizontal cells could also alter the activity of GABA transport in these cells. Our results show that activation of NMDA receptors inhibited GABA transporter currents on carp retinal horizontal cells. Application of AP5 (100μM) eliminated the inhibition, which indicates NMDA receptors are necessary. When the extracellular Ca2+ was removed from the Ringer's solution, the NMDA inhibitory effect on the GABA transporter current was eliminated. The suppression effect could be attenuated when the Ca2+ release and Ca2+ uptake of intracellular Ca2+ store were blocked after the cell had been pre-incubated with 20μM ryanodine plus 2μM thapsigargin for 3 min. After achieving the whole-cell configuration, the retinal horizontal cell was loaded with 10 mM BAPTA by diffusion from the patch pipette for 5 min, and then the NMDA modulation of GABA transporters was suppressed. These results suggest that the activation of NMDA receptors inhibits GABA transporter-mediated current by affecting Ca2+ processes in the retinal horizontal cells. Furthermore, a model containing endoplasmic reticulum (ER) membrane processes and plasma membrane processes was constructed for quantitative analyses of the NMDA-triggered Ca2+ dynamics and its modulation of GABA transporters. The model results suggest that the intracellular Ca2+ store is involved in the NMDA modulation of GABA transporters.
It is reported that NMDA receptors are expressed in retinal horizontal cells of carp, rat and human, but physiological function of the NMDA receptors is not clear. In present study, our results show that activation of NMDA receptors in carp retinal horizontal cells inhibit GABA transporter current. Under physiological conditions, photoreceptors release glutamate continually in the darkness. AMPA receptors and NMDA receptors on the retinal horizontal cells are activated, and extracellular calcium influx can occur through these glutamate receptors. Since AMPA receptors display very rapid and significant desensitization, and NMDA receptors do not desensitize as quickly or as fully, NMDA receptors might play a significant role in the regulation of GABA transporter activities under physiological conditions.
引文
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[14] Kamermans M, Fahrenfort I, Schultz K, Janssen-Bienhold U, Sjoerdsma T, and Weiler R. Hemichannel-mediated inhibition in the outer retina. Science, 2001; 292(5519): 1178-80.
[15] Mangel SC. Analysis of the horizontal cell contribution to the receptive field surround of ganglion cells in the rabbit retina. J Physiol, 1991; 442: 211-34.
[16] Yang XL. Characterization of receptors for glutamate and GABA in retinal neurons. Prog Neurobiol, 2004; 73(2): 127-50.
[17] Shen Y, Zhang M, Jin Y, and Yang XL. Functional N-methyl-D-aspartate receptors are expressed in cone-driven horizontal cells in carp retina. Neurosignals, 2006; 15(4): 174-9.
[18] Borden LA. GABA transporter heterogeneity: pharmacology and cellularlocalization. Neurochem Int, 1996; 29(4): 335-56.
[19] Nelson N. The family of Na+/Cl- neurotransmitter transporters. J Neurochem, 1998; 71(5): 1785-803.
[20] Clark JA. Analysis of the transmembrane topology and membrane assembly of the GAT-1 gamma-aminobutyric acid transporter. J Biol Chem, 1997; 272(23): 14695-704.
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