Department of Veterinary Sciences, University of Antwerp, Antwerp, Belgium.
Histochem Cell Biol. 2011 Oct;136(4):371-85. doi: 10.1007/s00418-011-0837-8. Epub 2011 Aug 6.
Afferent activities arising from sensory nerve terminals located in lungs and airways are carried almost exclusively by fibres travelling through the vagus nerve. Based on electrophysiological investigations, intrapulmonary airway-related vagal afferent receptors have been classified into three main subtypes, two of which are myelinated and mechanosensitive, i.e., rapidly and slowly adapting receptors. To allow for a full functional identification of the distinct populations of airway receptors, morphological and neurochemical characteristics still need to be determined. Nerve terminals visualised using markers for myelinated vagal afferents seem to be almost uniquely associated with two morphologically well-formed airway receptor end organs, smooth muscle-associated airway receptors (SMARs) and neuroepithelial bodies (NEBs), localised in airway smooth muscle and epithelium, respectively. Due to the lack of a selective marker for SMARs in mice, no further neurochemical coding is available today. NEBs are extensively innervated diffusely spread groups of neuroendocrine cells in the airway epithelium, and are known to receive at least two separate populations of myelinated vagal afferent nerve terminals. So far, however, no evidence has been reported for the expression of channels that may underlie direct sensing and transduction of mechanical stimuli by the receptor terminals in NEBs and SMARs. This study focused on the expression of mechanogated two-pore domain K(+) (K(2P)) channels, TREK-1 and TRAAK, in mouse airways and more particular in the NEB micro-environment and in SMARs by multiple immunostaining. TREK-1 could be detected on smooth muscle cells surrounding intrapulmonary airways and blood vessels, while TRAAK was expressed on myelinated vagal afferents terminating both in SMARs and in the NEB micro-environment. Co-stainings with known markers for subpopulations of myelinated vagal afferents and general neuronal markers revealed that all identified SMARs exhibit TRAAK immunoreactivity, and that at least three subpopulations exist in mouse airways. Also, the intraepithelial terminals of both subpopulations of NEB-associated myelinated vagal sensory nerve fibres were shown to express TRAAK. In conclusion, the present study finally characterised an intrinsically mechanosensitive ion channel, the K(2P) channel TRAAK, on the terminals of identified myelinated vagal nodose airway afferents, organised as SMARs and as components of the innervation of NEBs. These data support the hypothesis that both SMARs and NEBs harbour the morphological counterparts of electrophysiologically identified myelinated vagal airway mechanoreceptors. TRAAK appears to be strongly involved in regulating airway mechanosensing since it was found to be expressed on the terminals of all subpopulations of potential vagal mechanosensors.
位于肺部和气道的感觉神经末梢的传入活动几乎完全由通过迷走神经传递的纤维携带。基于电生理学研究,肺内气道相关的迷走传入受体已被分为三种主要亚型,其中两种是有髓和机械敏感的,即快速和缓慢适应受体。为了充分识别气道受体的不同群体,仍需要确定形态和神经化学特征。使用有髓迷走传入神经末梢标记物可视化的神经末梢似乎几乎仅与两种形态形成良好的气道受体终末器官相关,即平滑肌相关气道受体 (SMARs) 和神经上皮体 (NEBs),分别位于气道平滑肌和上皮中。由于在小鼠中缺乏 SMARs 的选择性标记物,目前尚无进一步的神经化学编码。NEBs 是气道上皮中广泛分布的神经内分泌细胞的弥散群集,已知接收至少两种单独的有髓迷走传入神经末梢群体。然而,迄今为止,尚无证据表明受体末端在 NEB 和 SMAR 中直接感知和转导机械刺激的通道表达。本研究通过多重免疫染色聚焦于机械门控双孔域 K(+) (K(2P)) 通道 TREK-1 和 TRAAK 在小鼠气道中的表达,特别是在 NEB 微环境和 SMARs 中的表达。TREK-1 可在肺内气道和血管周围的平滑肌细胞上检测到,而 TRAAK 在终止于 SMARs 和 NEB 微环境中的有髓迷走传入神经末梢上表达。用有髓迷走传入神经末梢的亚群和一般神经元标记物的已知标记物进行共染色显示,所有鉴定的 SMARs 均表现出 TRAAK 免疫反应性,并且在小鼠气道中至少存在三个亚群。此外,两个 NEB 相关有髓迷走感觉神经纤维亚群的上皮内末端均表达 TRAAK。总之,本研究最终在作为 SMARs 组织的以及作为 NEB 支配的组成部分的已鉴定的有髓迷走结节气道传入神经末梢的末端上对内在机械敏感离子通道,即 K(2P) 通道 TRAAK 进行了特征描述。这些数据支持这样的假设,即 SMARs 和 NEB 都包含电生理上鉴定的有髓迷走气道机械感受器的形态对应物。TRAAK 似乎强烈参与调节气道机械感觉,因为它在所有潜在的迷走机械感受器的亚群的末端上均有表达。