The function and connectivity of neocortical layer 7 and long-range inputs to somatosensory cortex.

新皮质第1层和第6层(或第7层)位于皮质地幔的两侧，在新皮质个体发育过程中从第一回路中出现，仍然是新皮质中最神秘的一层。第一层含有少量局部连接的中间神经元，并因其丰富的长程轴突和终末阵列而脱颖而出，这些轴突和终末携带着影响第2/3层和第5层锥体神经元顶束树突的高级加工信息。远程输入对第2/3层和第5层局部中间神经元和潜在锥体神经元胞体的影响在很大程度上是未知的。第七层有各种各样的兴奋性和抑制性神经元，包括一个突出的轴突投射到第一层，也值得注意的是，作为唯一对促进觉醒的神经肽增食欲素做出反应的皮质层。第7层输入和输出的生理机制仍然完全未被研究。在这里，我们研究了第一层和第七层的局部和远程输入的突触生理学。我们将使用我们实验室开发的使用钙整合器Campari的新的电路映射策略，以及有针对性的膜片钳记录，来全面阐明到第七层的输入和输出的连接和生理，以及受第一层影响的神经元的有针对性的修补。为了进一步表征第一层的功能，我们将在体内测量睡眠-觉醒行为，同时用增食欲素影响第七层特定的神经元。从这些实验中获得的数据将深入了解第1层和第7层的功能连接和潜在功能，以及第7层在唤醒和睡眠期间的行为影响。

Neocortical layer 1 and layer 6b (or layer 7), which reside on opposite sides of the cortical mantle and emerge from the first circuits during neocortical ontogeny, remain the most mysterious lamina in the neocortex. Layer 1 contains a sprinkle of locally-connected interneurons, and stands out among cortical layers for its rich array of long-range axons and terminals that carry high-order processed information that influences apical tuft dendrites of pyramidal neurons in layer 2/3 and layer 5. The influence of long-range inputs onto local interneurons and underlying pyramidal neuron somata in layer 2/3 and layer 5 are largely unknown. Layer 7 has a diverse array of excitatory and inhibitory neurons, including a prominent axonal projection into layer 1 and is also noteworthy as the only cortical layer that responds to the wakefulness promoting neuropeptide orexin. The physiology of layer 7 inputs and outputs remain entirely unexplored. Here we examine the synaptic physiology of local and long-range inputs to layer 1 and layer 7. We will employ both a novel circuit mapping strategy developed in our laboratory using the calcium integrator CaMPARI, as well as targeted patch clamp recordings to fully elucidate the connectivity and physiology of inputs and outputs to layer 7 and targeted patching of neurons influenced by layer 1. To further characterize the function of layer 7 we will perform in vivo measurements of sleep-wake behavior while influencing layer 7-specific neurons both optogenetically and pharmacologically with orexin. The data acquired from these experiments will provide a large inroad into the functional connectivity and potential functions of layer 1 and 7 and the behavioral influence of layer 7 during wakefulness and sleep.