兔子視網膜中方向選擇性節細胞的發育過程與突觸連結

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研究生：	詹雅茜 Chan, Ya-Chien
論文名稱：	兔子視網膜中方向選擇性節細胞的發育過程與突觸連結 The maturation process and synaptic mechanism of the direction selective ganglion cell in the rabbit retina
指導教授：	焦傳金 Chiao, Chuan-Chin
口試委員:	焦傳金張兗君連正章王致恬陳示國
學位類別：	博士 Doctor
系所名稱：	生命科學暨醫學院 - 分子醫學研究所 Institute of Molecular Medicine
論文出版年：	2013
畢業學年度：	101
語文別：	英文
論文頁數：	78
中文關鍵詞：	視網膜節細胞、方向選擇性強度、細胞間隙追蹤劑聯結現象、背景樣式調控、麩胺酸受器輸入訊息、伽馬氨基丁酸受器側向抑制作用
外文關鍵詞：	Retinal ganglion cell, Direction tuning strength, Tracer coupling pattern, Contextual tuning, Glutamatergic input, GABAergic lateral inhibition
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方向選擇性乃視覺系統中極為重要的特性之一，這樣的反應特徵在視網膜中即可透過精密的神經網絡而形成。目前已知在哺乳類動物視網膜中有一類名為ON-OFF方向選擇性節細胞(簡稱為DSGC)，這類細胞對於物體往其偏好方向移動時可產生強烈的反應，而對於物體往其無效(相反)方向移動時卻產生極弱的反應或甚至沒有反應。本論文是由兩篇關於DSGC的研究構成，分別是針對其發育成熟過程及突觸連結機制所進行的深入探討。
先前研究指出，在動物開眼時方向選擇性的神經網絡即已建立完成，然而關於四種DSGC細胞型態(亦即偏好偵測物體在視覺空間中的移動為往上、往下、往前及往後的四個方向)在此發育時期的兩大特徵(亦即偏好方向的分佈情形及其對應於視網膜上垂直與水平軸的程度)卻不清楚。在第一篇研究中，我們使用出生後10~12天(開眼後1~2天)的兔子來觀測DSGC的偏好方向，我們發現DSGC的四種細胞型態尚未明顯區隔且其偏好方向並非完全能對應於視網膜的四大軸。此外，我們亦發現此時期的DSGC其方向選擇性的強度較成兔弱，加上與其偏好方向角度無明顯關聯性，因此推測DSGC在方向選擇性強度與偏好方向角度上的兩個發育過程是獨立進行的。另外，我們亦觀察到原本在成兔中不應具有電突觸連結的DSGC類型卻在此時期有細胞間隙追蹤劑聯結現象，因此推斷方向選擇性的神經網絡在動物開眼時尚未建立完成。總括來說，第一篇研究的結果表示，兔子視網膜中四種DSGC類型在開眼後仍需要經過相當程度的修飾以完成其功能上的發育成熟。
DSGC除了對於物體移動方向具有選擇性外，它的反應亦會受到物體移動時的背景樣式所影響。先前的研究已證實，當視覺刺激的時空樣式在其中央與周圍感受域不同時，DSGC的反應會隨之不同，然而此項特性背後的突觸連結機制尚不清楚。第二篇研究即是要找出DSGC對於背景相位差不同反應的主要原因。我們使用出生後14~25天的兔子進行實驗，並利用細胞上鬆弛膜片鉗與全細胞膜片鉗記錄方式來偵測DSGC對於中央與周圍光柵所造成不同相位差的反應。我們的結果顯示，當中央與周圍光柵為相同相位時，DSGC的動作電位反應會被強烈抑制，而在相反相位時其抑制反應會明顯減弱。更重要的是，我們發現無論光柵移動方向為偏好方向或無效方向，DSGC的突觸前興奮性輸入訊號亦在不同背景相位差時有不同程度的反應。藉由去除DSGC上經由菸鹼酸乙醯膽鹼受器所接收的突觸前輸入訊息，我們亦發現經由麩胺酸受器接收來自於雙極細胞的突觸前興奮性輸入訊息，在不同背景相位差時會接受不同程度的抑制作用。進一步的藥理實驗則指出此抑制作用是來自於雙極細胞軸突末端上的伽馬氨基丁酸A型與C型受器的活化。總括來說，第二篇研究的結果指出背景樣式對於DSGC不同反應的影響，主要是藉由視覺刺激在周圍感受域的側向抑制作用，引發雙極細胞上伽馬氨基丁酸受器的活化，以抑制雙極細胞的突觸前興奮性輸入來調控DSGC對背景相位的反應。
總體來說，本論文對於兔子視網膜中DSGC發育成熟過程及其突觸連結機制的研究，有效的提供了我們探索神經系統發育與運作機制的基礎。而透過對於視網膜神經網絡中的發育成熟過程及其突觸連結機制的了解，本論文亦對發展重建視覺與大腦功能的治療方向有重要貢獻。

Direction selectivity is an important feature throughout the visual system, and this arises from within the intricate neural network of the retina. The ON-OFF direction selective ganglion cells (DSGCs) in the mammalian retina respond vigorously to an object moving in their preferred direction but show little or no response at all to movement in their null (the opposite) direction. In this dissertation, there are two independent studies of the DSGCs, one examining their maturation process and the other investigating their synaptic mechanism.
Although it has been known that the basic neural circuit of direction selectivity is established at around the time of eye opening, it is less known if the four DSGC subtypes (i.e., those responsible for the detection of motion in the superior, inferior, anterior, and posterior directions of the visual field) can be unambiguously distinguished and their preferred directions are aligned with four canonical axes at this developmental stage. By examining the preferred directions of DSGCs in P10-12 rabbit retinas and characterizing their distribution pattern, in the first study, we have shown that the preferred directions of DSGCs at eye opening are not distinctly segregated but rather are diffusely distributed along the four canonical axes. Furthermore, the fact that the direction tuning strength of DSGCs at P10-12 is weaker than that in adults, and this was found not to be correlated with their preferred directions, suggests that the maturations of direction selectivity and preferred direction are independent processes. In addition, we also found that the subtypes of DSGCs, which do not display tracer coupling pattern in the adult, show extensive coupling at P10-12. Taken together, the first study supports that the significant refinement after eye opening is required for the development of the four functional DSGC subtypes in the rabbit retina.
In addition to the prominent trigger feature of direction selectivity, it is also known that the spiking response of DSGCs is context dependent (i.e., the cell responds differently depending on the spatiotemporal relationship of visual stimuli between the receptive field center and its surround), though the underlying synaptic mechanism is not fully understood. The second study was to identify the key components of the contextual phase-tuning of DSGCs. By using loose on-cell and whole-cell patch clamp recording, effects of the phase difference between of the center and surround moving gratings on the responses of the DSGC were investigated in P14-25 rabbit retinas. Consistent with the previous study, we have shown that spike responses of DSGCs to the center drifting grating are strongly suppressed by the surround grating when the two gratings are moving in-phase, but are only minimally suppressed when the grating are moving out-of-phase. Importantly, the excitatory inputs to the DSGC are also contextually tuned, regardless whether the gratings move in the preferred or the null direction. By removing the nicotinic cholinergic input, we have shown that the glutamatergic input from bipolar cells is already contextually tuned. Further experiments showed that the tuning is mediated by GABAergic inputs through the activation of both GABAA and GABAC receptors. Taken together, the second study suggests that contextual effect of DSGCs is mediated predominantly by the tuning of the excitatory inputs from bipolar cells via GABAergic lateral inhibition from the receptive field surround.
In conclusion, by elucidating the maturation process and synaptic mechanism of DSGCs in the rabbit retina, this dissertation sheds light on how intricate neural circuits develop and function in the mammalian retina. Understanding the activity-dependent refinement and synaptic connection of information processing in the retina will potentially illuminate the key steps in developing therapeutic strategies aimed at restoring vision and brain function.

致謝    i
Abstract    ii
中文摘要    v
Abbreviations    vii
Chapter 1: Introduction    1
1.1    The retina    1
1.2    Direction selectivity in the retina    2
1.2.1    The functional diversity of the DSGCs    2
1.2.2    The development of the DSGCs    3
1.3    Contextual selectivity in the retina    4
1.3.1    Contextual tuning of the DSGCs    4
1.3.2    The synaptic mechanism of the DSGCs    5
1.4    Specific aims    6
Chapter 2: Materials and Methods    8
2.1    Ethical approval    8
2.2    Retina preparation    8
2.3    Light Stimuli    10
2.4    Extracellular recording    12
2.5    Intracellular dye injection    13
2.6    Image acquisition    14
2.7    On-cell loose patch clamp recording    14
2.8    Whole-cell patch clamp recording    15
2.9    Pharmacology    15
2.10    Data analysis    16
Chapter 3: Results    19
3.1    The maturation process of the direction selective ganglion cell in the rabbit retina                19
3.1.1    The distribution of the preferred directions of the DSGCs at around the time of eye opening cannot be distinctly segregated into four groups    19
3.1.2    The preferred directions of DSGCs are diffusely distributed along the four canonical axes immediately after eye opening    21
3.1.3    The distribution of the preferred directions of the DSGCS at around the time of eye opening does not correlate with their direction tuning strength    23
3.1.4    The preferred directions of DSGCs displaying tracer coupling patterns immediately after eye opening do not conform to one canonical direction    24
3.1.5    Summary    25
3.2    The synaptic mechanism of the direction selective ganglion cell in the rabbit retina    26
3.2.1    Contextual phase tuning of DSGCs is mediated predominantly by the modulation of the excitatory inputs    26
3.2.2    The excitatory inputs during null-direction stimulus movement are also contextually tuned    28
3.2.3    Contextual phase tuning of the glutamatergic input requires GABAergic inhibition    29
3.2.4    Contextual phase tuning of DSGCs involves a dynamic center-surround antagonistic mechanism    31
3.2.5    Summary    32
Chapter 4: Discussion    34
4.1    The development of the DSGCs continues after eye opening    34
4.1.1    The four subtypes of DSGCs continue to refine after eye opening    34
4.1.2    Refinement of preferred directions of DSGCs but their subtype induction is activity dependent    36
4.1.3    Implications of the DSGC preferred direction distribution at eye opening    38
4.2    The synaptic inputs of the DSGCs are contextually tuned    40
4.2.1    Contextual phase tuning of DSGCs reside at the level of bipolar cells    40
4.2.2    Contextual phase tuning of glutamatergic inputs of DSGCs is shaped by lateral inhibition through activations of GABAA and GABAC receptors    42
4.2.3    Motion enhances contextual phase tuning through a dynamic spatiotemporal antagonism of the receptive field center and surround    43
4.3    Conclusion    44
References    46
Tables    54
Table 1    The statistics for the preferred directions of DSGCs along each of four canonical axes in adult and postnatal rabbits and mice    54
Table 2    A comparison of the tuning strengths of DSGCs in adult and postnatal rabbits        55
Figures    56
Figure 1    The diagram of the vertebrate retina    56
Figure 2    The preferred direction of a DSGC recorded at around the time of eye opening does not belong to one of four canonical directions    57
Figure 3    The distribution of preferred directions of DSGCs at around the time of eye opening cannot be separated into distinct clusters    59
Figure 4    The preferred directions of 2-4 DSGCs recorded on the same retina at around the time of eye opening are not perfectly aligned along the canonical axes    61
Figure 5    The preferred directions of DSGCs on the same retina at around the time of eye opening are not aligned orthogonally or in opposite directions    62
Figure 6    The preferred directions of the DSGCs at around the time of eye opening are diffusely distributed across the four canonical axes    63
Figure 7    The preferred direction distributions of the DSGCs at around the time of eye opening are independent of the tuning strength of their direction selectivity    65
Figure 8    The DSGCs with prominent tracer coupling patterns at around the time of eye opening do not have the same preferred directions    67
Figure 9    The nearest neighbor analysis of tracer coupled DSGCs shown in Figure 8    69
Figure 10    Confocal images of DSGCs at different focal planes shown in Figure 8B                70
Figure 11    Contextual phase tuning of DSGCs is mediated predominantly by the tuning of the excitatory inputs    71
Figure 12    The excitatory inputs during null-direction stimulus movement are also contextually tuned    73
Figure 13    Contextual phase tuning of the glutamatergic input involves a GABAA and GABAC receptor-mediated pathway    74
Figure 14    Contextual phase tuning of DSGCs can be explained by a dynamic center surround antagonistic mechanism at the level of bipolar cells    76
Figure 15    Working model of contextual phase tuning of the DSGC    78

                                

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