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The Impact of Sensory-guided Behavior of Neuronal Processing in the Mouse Barrel Cortex and and a Novel Two-alternative Forced Choice Paradigm in Head-fixed Mice and Rats

机译:小鼠桶状皮质中神经元加工的感官指导行为的影响以及在头固定型小鼠和大鼠中的新型两种替代性强迫选择范例

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摘要

The reconfiguration of neuronal networks due to perceptual learning, and the impact of behavior in a task on online sensory processing are currently critical questions in neuroscience. In the work presented here, these topics were investigated in the barrel cortex of mice performing a sophisticated detection and discrimination task through the lens of two-photon calcium imaging. Layer 2/3 of the primary sensory cortex has been established as an essential locus of adult experience-dependent plasticity. Due to its unique topographical organization, the barrel cortex is an excellent area to investigate modulations of sensory processing in during whisker-dependent behavior. Before recording neuronal activity during learning of a sensory-guided task in project number two, we examined the ability of mice to perform a complex task successfully.\udThus, the primary goal of the first project was to establish if mice can learn complex behavior equally well as rats. In our two-alternative forced choice (2-AFC) paradigm, rodents learned to discriminate simultaneously applied single-whisker vibrotactile stimuli of different repetition rates. Both rats and mice could perform over 400 trials a day, with performance levels reaching over 90% of correct responses. The key finding of this project is that both species show comparable results regarding several behavioral readouts. Namely, their psychometric curves were similar with an average perceptual threshold of 53.0 Hz, and a 50.6- Hz frequency difference with corresponding Weber fractions of 0.58 and 0.56 respectively in rats and mice. Furthermore, their reaction times, the amount of omitted trials, rates of learning, and impulsivity were comparable among the two species. Additionally, we found that whisking before stimulus presentation impaired performance. A fundamental characteristic of our 2-AFC behavioral paradigm paired with whisker tracking is that it allows for a very precise control of the applied vibrotactile stimuli. Since the animal is head-fixed, additional recording and/or stimulating methods can be employed, such as electrophysiology or optogenetics. In the second project, its combination with long-term two-photon imaging will be presented. Furthermore, the task is very suitable for investigating interhemispheric interactions due to the possibility of bilateral stimulation. The main conclusion of the first project is that mice are equally adept as rats in learning complex behavior, such as the 2-AFC detection and discrimination tasks. This result opens up a multitude of possible research avenues by virtue of fusing behavioral measurements with potent recording and/or stimulating methods and taking advantage of the rich array of transgenic lines created in mice.\udIn the second project, we investigated plasticity effects due to behavior in a sensory-guided task, along with the impact of attentional engagement on sensory processing in the barrel cortex. To achieve this goal, our 2-AFC task was combined with long-term two-photon calcium imaging in layer 2/3 in the somatosensory cortex. This layer is a major site of adult cortical plasticity. However, our understanding of it is limited, since most studies focused on the granular and infragranular layers. Our data revealed strong plasticity effects due to perceptual whisker-dependent learning. During the baseline recording sessions, mice were passively exposed to sensory stimuli. Once the animals engaged in the operant conditioning sensory-guided task, the stimuli gained behavioral relevance by becoming associated with a water reward. Hence, layer 2/3 neurons display flexible modulation depending on the behavioral context, showing a more complex role for the primary sensory cortex than solely a passive relay of sensory information. Plasticity effects were equally reflected in comparing anesthesia recordings from before and after learning. A robust trend in sensory representation is the increase of evoked calcium transients with increasing repetition rates of the vibrotactile stimulus, also shown in this thesis. However, this monotonic increment was disrupted after the learning took place, which exclusively involved the highest and lowest frequencies. The middle frequency, never applied during the task, received fewer calcium transients than the lowest frequency, thus eliminating the distinctive property of vibrotactile frequency representation. Furthermore, we report the excitatory effect on the cortical representation of somatosensory stimuli during active behavior in the task, as opposed to passive stimulus exposure during unattended trials. Our bilateral 2-AFC paradigm allowed for the comparison of contralateral and ipsilateral responses, which was previously unknown in layer 2/3. We report that contralateral calcium responses are substantially larger than ipsilateral ones, contrary to the infragranular layers, where they are similar in size. To summarize the second project, behavior in a sensory-guided paradigm shapes both online sensory processing and long-term plasticity effects in layer 2/3 of the barrel cortex.
机译:由于知觉学习导致的神经元网络的重新配置以及任务中的行为对在线感官处理的影响目前是神经科学中的关键问题。在此处介绍的工作中,通过双光子钙成像镜头在执行复杂检测和辨别任务的小鼠的桶状皮质中对这些主题进行了研究。主要感觉皮层的2/3层已被确定为成年人依赖于经验的可塑性的重要场所。由于其独特的地形组织,桶状皮质是研究晶须依赖行为期间感觉过程调节的极好区域。在记录第二个项目中的感官指导任务期间的神经元活动之前,我们检查了小鼠成功完成复杂任务的能力。\ ud因此,第一个项目的主要目标是确定小鼠是否可以平等地学习复杂行为以及老鼠。在我们的两种选择的强制选择(2-AFC)范式中,啮齿动物学会了区分不同重复率的同时应用的单晶须触觉刺激。大鼠和小鼠每天都可以进行400多次试验,其水平达到正确反应的90%以上。该项目的主要发现是,两个物种在几种行为读数方面均显示出可比的结果。即,它们的心理测度曲线相似,在大鼠和小鼠中平均感知阈值为53.0 Hz,频率差为50.6- Hz,相应的韦伯分数分别为0.58和0.56。此外,它们的反应时间,省略试验的次数,学习率和冲动性在两个物种之间是可比的。此外,我们发现刺激刺激之前的搅拌会损害性能。我们的2-AFC行为范式与晶须跟踪配合使用的基本特征是,它可以非常精确地控制所施加的触觉刺激。由于动物是头部固定的,因此可以采用其他记录和/或刺激方法,例如电生理学或光遗传学。在第二个项目中,将介绍其与长期双光子成像的组合。此外,由于有双边刺激的可能性,该任务非常适合研究半球之间的相互作用。第一个项目的主要结论是,小鼠在学习复杂行为(例如2-AFC检测和区分任务)方面与大鼠一样熟练。通过将行为测量与有效的记录和/或刺激方法相融合,并利用在小鼠中产生的丰富的转基因品系,该结果开辟了许多可能的研究途径。\ ud在第二个项目中,我们研究了由于感官指导任务中的行为,以及注意参与对桶状皮质中感官处理的影响。为了实现此目标,我们将2-AFC任务与体感皮质中第2/3层的长期双光子钙成像相结合。该层是成人皮质可塑性的主要部位。但是,由于大多数研究都集中在颗粒层和颗粒下层,因此我们对它的理解是有限的。我们的数据显示,由于感知晶须依赖学习,因此具有很强的可塑性效应。在基线记录期间,小鼠被动地受到感觉刺激。一旦动物从事了操作性调节感觉指导任务,刺激就会通过与水奖励相关联而获得行为相关性。因此,第2/3层神经元根据行为环境显示灵活的调制方式,与仅被动传递感觉信息相比,对主要感觉皮质的作用更为复杂。在比较学习前后的麻醉记录时,可塑性效果同样得到反映。感觉表征的一个强劲趋势是诱发的钙瞬变随着触觉刺激重复率的增加而增加,这也证明了这一点。但是,这种单调增量在学习发生后就被打乱了,它只涉及最高和最低频率。在任务执行过程中从未使用过的中频接收到的钙瞬变少于最低频率,因此消除了触觉频率表示法的独特特性。此外,我们报告了在任务的主动行为期间,对躯体感觉刺激的皮质表示的兴奋作用,与无人值守试验期间的被动刺激暴露相反。我们的双边2-AFC范式允许对侧和同侧反应的比较,这在第2/3层中是未知的。我们报告说,对侧钙反应比同侧钙反应大得多,这与颗粒大小相似的颗粒下层相反。总结第二个项目,在感觉引导范式下的行为会影响在线感觉处理以及桶皮质的第2/3层中的长期可塑性影响。

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    Skreb, Vida;

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  • 年度 2016
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