토끼 망막에서 칼빈딘 면역반응성 ON 원뿔두극세포의 속얼기층 OFF 아층판에서의 리본연접
Ribbon Synapses Formed at the Axon of the Calbindin-Immunoreactive ON Cone Bipolar Cell in OFF Sublamina of the Inner Plexiform Layer in the Rabbit Retina
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초록
내인성광수용신경절세포(intrinsically photosensitive retinal ganglion cell)를 비롯한 일부 망막 신경원은속얼기층(inner plexiform layer)의 OFF 아층판(sublayer)에 가지돌기가 분포함에도 불구하고 역설적 ON 빛반응을 보인다. 이러한 기전을 이해하고자, 토끼 망막에서 이러한 신경원과 리본연접(ribbon synapse)을 OFF 아층판에서 형성할 것으로 생각되는 칼빈딘 면역반응성 ON 원뿔두극세포(calbindin-immunoreactive ON cone bipolar cell) 축삭이 속얼기층의 OFF 아층판에서 실제로 연접을 이루어 ON 신호를 전달할 수 있는지를 조사하였다. 칼빈딘 항체를 이용한 면역전자현미경법(immunoelectron microscopy)을 수행하여 표본을 만들고 전자현미경을 통해 망막 속얼기층의 OFF 아층판을 통과하는 칼빈딘 ON 원뿔두극세포의 축삭에서 리본연접을 관찰하였다. OFF 아층판에서의 리본연접은 주로 속얼기층의 가장 바깥쪽인 속핵층(inner nuclear layer)과 속얼기층 경계부에서 주로 발견되었고, 연접후성분은 한동이(monad)였다. 또한 이러한 리본연접에는 하나 이상의 연접리본(synaptic ribbon)이 동원되는 경향이 있었다. 이러한 양상들은 ON 아층판인 속얼기층의 b 아층판(sublamina b)에 종지하는 ON 원뿔두극세포의 축삭종말이 형성하는 통상적인 리본연접(conventional ribbon synapse)과는 다른 형태로, ON 원뿔두극세포가 OFF 아층판인 속얼기층 a 아층판에서 이루는 축삭리본연접(axonal ribbon synapse)의 특징으로 생각된다. 본 연구결과는 망막의 신경회로 연구에 있어서 속얼기층에서 연접의 종류를 파악하는데 유용하게 활용될 것으로 생각된다.
Abstract
Some retinal neurons, including intrinsically photosensitive retinal ganglion cells have their dendrites stratified in sublamina a of the inner plexiform (IPL), the OFF sublayer, but paradoxically show light-driven ON electrophysiological responses. In order to understand the mechanism on this paradoxical response, by using immunoelectron microscopy with a specific antibody against calbindin, we examined the synaptic connections of the calbindin-immunoreactive ON cone bipolar cell of the rabbit retina, which is thought to make the ribbon synapse in sublamina a of the IPL. The ribbon synapses in sublamina a by calbindin-immunoreactive ON cone bipolar cells were mainly found at the border between the inner nuclear layer and the IPL. Interestingly, the output targets at these ribbon synapses turned out as monads, and multiple synaptic ribbons were engaged in each synapse. These findings were different from those at the conventional ribbon synapse formed by calbindin-immunoreactive ON cone bipolar axon terminals. Thus, these findings may be the characteristics of the calbindin-immunoreactive ON cone bipolar ribbon synapse in sublamina a and can be used to classify the synapse in the retinal circuit research.
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