Hanson Elizabeth, Danbolt Niels Christian, Dulla Chris G
Department of Neuroscience, Tufts University School of Medicine, 136 Harrison Avenue, Boston, MA 02111, USA; Neuroscience Program, Tufts Sackler School of Biomedical Sciences, 136 Harrison Avenue, Boston, MA 02111, USA.
Department of Molecular Medicine, Institute of Basic Medical Sciences, University of Oslo, P.O. Box 1105 Blindern, N-0317 Oslo, Norway.
Neurobiol Dis. 2016 May;89:157-68. doi: 10.1016/j.nbd.2016.02.012. Epub 2016 Feb 10.
Developmental cortical malformations (DCMs) are linked with severe epilepsy and are caused by both genetic and environmental insults. DCMs include several neurological diseases, such as focal cortical dysplasia, polymicrogyria, schizencephaly, and others. Human studies have implicated astrocyte reactivity and dysfunction in the pathophysiology of DCMs, but their specific role is unknown. As astrocytes powerfully regulate glutamate neurotransmission, and glutamate levels are known to be increased in human epileptic foci, understanding the role of astrocytes in the pathological sequelae of DCMs is extremely important. Additionally, recent studies examining astrocyte glutamate uptake in DCMs have reported conflicting results, adding confusion to the field. In this study we utilized the freeze lesion (FL) model of DCM, which is known to induce reactive astrocytosis and cause significant changes in astrocyte morphology, proliferation, and distribution. Using whole-cell patch clamp recording from astrocytes, we recorded both UV-uncaging and synaptically evoked glutamate transporter currents (TCs), widely accepted assays of functional glutamate transport by astrocytes. With this approach, we set out to test the hypothesis that astrocyte membrane properties and glutamate transport were disrupted in this model of DCM. Though we found that the developmental maturation of astrocyte membrane resistance was disrupted by FL, glutamate uptake by individual astrocytes was robust throughout FL development. Interestingly, using an immunolabeling approach, we observed spatial and developmental differences in excitatory amino acid transporter (EAAT) expression in FL cortex. Spatially specific differences in EAAT2 (GLT-1) and EAAT1 (GLAST) expression suggest that the relative contribution of each EAAT to astrocytic glutamate uptake may be altered in FL cortex. Lastly, we carefully analyzed the amplitudes and onset times of both synaptically- and UV uncaging-evoked TCs. We found that in the FL cortex, synaptically-evoked, but not UV uncaging-evoked TCs, were larger in amplitude. Additionally, we found that the amount of electrical stimulation required to evoke a synaptic TC was significantly reduced in the FL cortex. Both of these findings are consistent with increased excitatory input to the FL cortex, but not with changes in how individual astrocytes remove glutamate. Taken together, our results demonstrate that the maturation of astrocyte membrane resistance, local distribution of glutamate transporters, and glutamatergic input to the cortex are altered in the FL model, but that single-cell astrocytic glutamate uptake is robust.
发育性皮质畸形(DCMs)与严重癫痫相关,由遗传和环境因素共同导致。DCMs包括多种神经系统疾病,如局灶性皮质发育不良、多小脑回、脑裂畸形等。人体研究表明星形胶质细胞反应性和功能障碍与DCMs的病理生理过程有关,但其具体作用尚不清楚。由于星形胶质细胞对谷氨酸神经传递有强大的调节作用,且已知人类癫痫病灶中的谷氨酸水平会升高,因此了解星形胶质细胞在DCMs病理后遗症中的作用极为重要。此外,最近关于DCMs中星形胶质细胞谷氨酸摄取的研究结果相互矛盾,给该领域带来了困惑。在本研究中,我们利用了DCM的冷冻损伤(FL)模型,该模型已知会诱导反应性星形胶质细胞增生,并导致星形胶质细胞形态、增殖和分布发生显著变化。通过对星形胶质细胞进行全细胞膜片钳记录,我们记录了紫外线光解和突触诱发的谷氨酸转运体电流(TCs),这是被广泛接受的星形胶质细胞功能性谷氨酸转运检测方法。通过这种方法,我们旨在检验以下假设:在这个DCM模型中,星形胶质细胞膜特性和谷氨酸转运受到破坏。尽管我们发现FL破坏了星形胶质细胞膜电阻的发育成熟,但在FL发育过程中,单个星形胶质细胞的谷氨酸摄取能力很强。有趣的是,通过免疫标记方法,我们观察到FL皮质中兴奋性氨基酸转运体(EAAT)表达存在空间和发育差异。EAAT2(GLT-1)和EAAT1(GLAST)表达的空间特异性差异表明,在FL皮质中,每种EAAT对星形胶质细胞谷氨酸摄取的相对贡献可能会发生改变。最后,我们仔细分析了突触诱发和紫外线光解诱发的TCs的幅度和起始时间。我们发现,在FL皮质中,突触诱发而非紫外线光解诱发的TCs幅度更大。此外,我们发现诱发突触TC所需的电刺激量在FL皮质中显著减少。这两个发现均与FL皮质兴奋性输入增加一致,但与单个星形胶质细胞清除谷氨酸的方式变化无关。综上所述,我们的结果表明,在FL模型中,星形胶质细胞膜电阻的成熟、谷氨酸转运体的局部分布以及皮质的谷氨酸能输入均发生了改变,但单个星形胶质细胞的谷氨酸摄取能力很强。