생쥐 망막에서 콜린성 무축삭세포의 분포 양상 및 연접회로에 대한 연구

Morphology and Synaptic Connectivity of Cholinergic Amacrine Cells in the Mouse Retina

  • 강화선 (가톨릭대학교 의과대학 세포의학.생물학과) ;
  • 천명훈 (가톨릭대학교 의과대학 해부학교실)
  • Kang, Wha-Sun (Catholic University of Korea, College of Medicine, Department of Natural Science) ;
  • Chun, Myung-Hoon (Catholic University of Korea, College of Medicine, Department of Anatomy)
  • 발행 : 2004.12.01

초록

생쥐의 망막에서 아세틸콜린의 합성효소인 choline acetyltransferase (ChAT)에 대한 항체를 이용한 면역세포화학법으로 콜린성 무축삭세포를 동정하고 분포양상을 조사하였다. 콜린성 무축삭세포는 망막에서 세포체의 위치에 따라서 두 종류로 구분된다. 즉 속핵층에 세포체가 위치하고 세포돌기는 속얼기층의 a 아층판에 위치하는 세포와 세포체가 신경절세포층에 위치하면서 세포돌기는 b 아층판에 위치하는 세포이다. GABA 항체를 이용한 이중 면역염색 결과 모든 콜린성 무축삭세포가 GABA에 대한 염색반응성을 나타내었다. 전자현미경 관찰 결과 콜린성 무축삭세포의 연접회로가 속얼기층에서 관찰되었다. 콜린성 무축삭세포는 두극세포와 가장 많은 수입연접을 형성하고 있었으며, 이 외에 콜린성 무축삭세포와 염색되지 않은 무축삭세포와의 연접도 관찰되었다. 콜린성 무축삭세포의 수출연접의 주요 대상은 신경절세포로 속얼기층의 b 아층판에서 더 빈번하게 관찰되었다. 이러한 연구결과로 생쥐 망막의 콜린성 무축삭세포도 다른 종류의 포유류와 매우 유사한 특징을 가지고 있으며, 방향선택성 신경절세포로 신호가 전달되는 과정에서 중요한 역할을 할 것으로 생각된다.

We investigated the morphology, distribution and synaptic connectivity of cholinergic neurons in the mouse retina by immunocytochemistry, using antisera against choline acetyltransferase (ChAT). ChAT-immunoreactive amacrine cells fall into two groups according to the localization of their somas in the retina: one is situated in the inner nuclear layer (INL), near the border of the inner plexiform layer (IPL), and the other is displaced in the ganglion cell layer (GCL). The dendrites of amacrine cells from the INL ramify in sublamina a and that of the displaced amacrine cells ramify in sublamina b of the IPL. Double labeling with an antisera against ChAT and r-aminobutyric acid (GABA) demonstrated that these labeled cells formed a subpopulation of GABAergic amacrine cells. The synaptic connectivity of ChAT-immunoreactive amacrine cells was identified in the IPL by electron microscopy. The most frequent synaptic input of ChAT-labeled amacrine cells was from bipolar cells in both sublaminae a and b of the IPL, followed by labeled amacrine cells and unlabeled amacrine cells. Their primary output targets were onto ganglion cells in both sublaminae a and b and output onto ganglion cells was more frequently observed in sublamina b of the IPL. Our results suggest that cholinergic amacrine cells in the mouse retina are very similar to their counter parts in other mammals, and they can attribute a major role in the pathway feeding into directionally selective ganglion cells.

키워드

참고문헌

  1. Amthor FR, Grzywacz NM: Directional selectivity in vertebrate retinal ganglion cells, Rev Oculomot Res 5 : 79 100,1993
  2. Ariel M, Adolph AR: Neurotransmitter inputs to directionally sensitive turtle retinal ganglion cells, J Neurophysiol 54 : 1123 1143,1985 https://doi.org/10.1152/jn.1985.54.5.1123
  3. Brandon C: Cholinergic neurons in the rabbit retina: dendritic branching and ultrastructural connectivity, Brain Res 426: 119 130, 1987 https://doi.org/10.1016/0006-8993(87)90431-8
  4. Brecha N, Johnson D, Peichel, Wassle H: Cholinergic amacrine cells of the rabbit retina contain glutamate decarboxylase and r aminobutyrate immunoreactivity. Proceedings of NationalAcad Sci U,S A.85 : 6187 6191,1988 https://doi.org/10.1073/pnas.85.16.6187
  5. Burgi PY, Grzywacz NM: Possible roles of spontaneous waves and dendritic growth for retinal receptive field development. Neural comput 9 : 533 553,1997 https://doi.org/10.1162/neco.1997.9.3.533
  6. Conley MD, Fitzpatrick D, Lachica EA: Laminar asymmetry in the distribution of choline acetyltransferase immunoreactive neurons in the retina of the tree shrew (Tupaia belangeri), Brain Res 399 : 332 338,1986 https://doi.org/10.1016/0006-8993(86)91524-6
  7. Euler T, Wassle H: Immunocytochemical identification of cone bipolar cells in the rat retina, J Comp Neurol 36 : 461 478,1995
  8. Famiglietti EV: 'Starburst' amacrine cells and cholinergic neurons: mirror symmetric ON and OFF amacrine cells of rabbit retina, Brain Res 261 : 138 144,1983 https://doi.org/10.1016/0006-8993(83)91293-3
  9. Famiglietti EV, Tumosa N: Immunocytochemical staining of cholinergic amacrine cells in rabbit retina, Brain Res 413(2) :398 403,1987 https://doi.org/10.1016/0006-8993(87)91037-7
  10. Famiglietti EV: Synaptic organization of starburst amacrine cells in rabbit retina: analysis of serial thin sections by electron microscopy and graphic reconstruction, J Comp NeuroI 309(1) : 40 70,1991 https://doi.org/10.1002/cne.903090105
  11. Freed MA: GABAergic circuits in the mammalian retina,Prog Brain Res 90 : 107 131, 1992 https://doi.org/10.1016/S0079-6123(08)63611-0
  12. Guiloff GD, Kolb H: Neurons immunoreactive to choline acetyltransferase in the turtle retina. Vision Research 32 : 2023 2030,1992 https://doi.org/10.1016/0042-6989(92)90063-O
  13. Grzywacz NM, Tootle JS, Amthor FR: Is the input to a GABAergic or cholinergic synapse the sole asymmetry in rabbit's retinal directional selectivity? Vis Neurosci 14 : 39 54,1997 https://doi.org/10.1017/S0952523800008749
  14. He SG, Masland RH: Changes in the responses of directionally selective ganglion cells in the rabbit retina induced by laser ablation of starburst amacrine cells, Invest Ophthalmol Vis Sci 38 : S949, 1997
  15. Kim I B, Oh S J, Na W K, Lee M Y, Chun M H: Distribution pattern and synaptic circuitry of cholinergic neurons in the rat retina. Korean J Anat 31 : 136 150, 1998
  16. Kolb H: The inner plexiform layer in the retina of the cat: electron microscopic observation. J Neurocytol 8 ; 295 329,1979 https://doi.org/10.1007/BF01236124
  17. MacNeil MA, Masland RH: Extreme diversity among amacrine cells implications for function. Neuron 20(5) : 971 982, 1998 https://doi.org/10.1016/S0896-6273(00)80478-X
  18. Marc RE, Liu W: Fundamental GABAergic amacrine cell circuitries in the retina: nested feedback, concatenated inhibition, and axosomatic synapses, J Comp Neurol 2000 425 (4) : 560 582, 2000
  19. Mariani AP, Hersh LB: Synaptic organization of cholinergic amacrine cells in the rhesus monkey retina, J Comp Neurol 267 : 269 280,1988 https://doi.org/10.1002/cne.902670209
  20. Masland RH, Tauchi M. A: Possible amacrine cell substrate for the detection of stimulus motion, Neurosci Res Suppl 12:S185 199,1985
  21. McGuire BA, Stevens J, Stering P: Microcircuitry of bipolar cells in cat retina, J Neurosci 4 : 2920 2938, 1984
  22. McGuire BA, Stevens J, Stering P: Microcircuitry of beta ganglion cells in cat retina, J Neurosci 6 : 907 918, 1986
  23. Mitrofanis J, Stone J: Distribution of cholinergic amacrine cells in the retinas of normally pigmented and hypopigmented strains of rat and cat, Vis Neurosci 1:367 376, 1988 https://doi.org/10.1017/S0952523800004132
  24. Neal MJ, Cunningham JR, Hutson PH, Semark JE: Calcium dependent release of Acetylcholine and gamma aminobutyric acid from the rabbit retina, Neurochem Int 20 : 43 53,1992 https://doi.org/10.1016/0197-0186(92)90124-A
  25. Pourcho RG, Osman K: Cytochemical identification of cholinergic amacrine cells in the cat retina, J Comp Neuro1 247 : 497 504,1986 https://doi.org/10.1002/cne.902470409
  26. Rodieck RW, Marshak DM: Spatial density and distribution of choline acetyltransferase immunoreactive cells in human, macaque, and baboon retinas, J Comp Neurol 321 : 46 64, 1992 https://doi.org/10.1002/cne.903210106
  27. Sandmann D, Englemann R, Peichl L: Starburst cholinergic amacrine cells in the tree shrew retina, J Comp Neurol 389 : 161 176,1997 https://doi.org/10.1002/(SICI)1096-9861(19971208)389:1<161::AID-CNE12>3.0.CO;2-O
  28. Schmidt M, Wassle H, Hunphrey M: Number and distribution of putative cholinergic neurons in the cat retina, Neurosci Lett 59 : 235 240,1985 https://doi.org/10.1016/0304-3940(85)90137-5
  29. Vaney DI: Many diverse types of retinalneurons show tracer coupling when injected with biocytin or Neurobiotin. Neurosci Lett. 29; 125(2): 187 90,1991 https://doi.org/10.1016/0304-3940(91)90024-N
  30. Voigt T: Cholinergic amacrine cells in the rat retina. J. Comp. Neurol. 351 : 374 384,1986 https://doi.org/10.1002/cne.903510305
  31. Werblin FS, Maguire A, Lukasiewicz PD, Eliasof S, Wu SM: Neural interactions mediating detection of motion in the retina of the tiger salamander, Vis Neurosci 1 : 317 329, 1988 https://doi.org/10.1017/S0952523800001978
  32. Wu SM, Maple BR: Amino acid neurotransmitters in the retina: a functional overview, Vision Res 38(10): 1371 1384,1988 https://doi.org/10.1016/S0042-6989(97)00296-4
  33. Zhang J, Jung CS, Slaughter MM: Serial inhibitory synapses in retina, Vis Neurosci 14(3) : 553 563, 1997 https://doi.org/10.1017/S0952523800012219
  34. Zhang J, Wu SM: Immunocytochemical analysis of cholinergic amacrine cells in the tiger salamander retina, Neuro Report 12(7): 1371 1375, 2001
  35. Zucker CL, Ehinger B, Grzywacz NM: GABA receptors are localized to starburst amacrine and ganglion cells in the rabbit retina, Soc Neurosci Abstr 24 : 136, 1998