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8.6:

์ „์ž์ „๋‹ฌ๊ณ„

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– [๊ฐ•์‚ฌ] ๋ฏธํ† ์ฝ˜๋“œ๋ฆฌ์•„์˜ ๋‚ด๋ง‰์˜์—ฌ๋Ÿฌ ์ฃผ๋ฆ„ ์†์— ๋ฌปํ˜€์žˆ๋Š” ๊ฒƒ์€ ์ „์ž ์ „๋‹ฌ์‚ฌ์Šฌ์˜ ์ˆ˜๋งŽ์€ ๋ณต์ œ๋ฌผ์ž…๋‹ˆ๋‹ค.์ด๊ฒƒ์€ ์—๋„ˆ์ง€ ์ถ”์ถœ์— ํ•„์ˆ˜์ ์ธ์ผ๋ จ์˜ 4 ๊ฐ€์ง€ ๋‹จ๋ฐฑ์งˆ ๋ณตํ•ฉ์ฒด์™€๊ด€๋ จ ์œ ๊ธฐ ๋ถ„์ž์ž…๋‹ˆ๋‹ค.์ „์ž๋Š” ์บ๋ฆฌ์–ด ๋ถ„์ž,๋‹ˆ์ฝ”ํ‹ด ์•„๋ฏธ๋“œ ์•„๋ฐ๋‹Œ ๋”” ๋‰ดํด๋ ˆ์˜คํ‹ฐ๋“œ๋˜๋Š” NADH, ํ”Œ๋ผ ๋นˆ ์•„๋ฐ๋‹Œ ๋”” ๋‰ดํด๋ ˆ์˜คํ‹ฐ๋“œ๋˜๋Š” FADH2๋ฅผ ์‚ฌ์šฉํ•˜์—ฌ ์‚ฌ์Šฌ๋กœ ๋“ค์–ด๊ฐ„๋‹ค.์ด๋Š” ๊ตฌ์—ฐ์‚ฐ ์‚ฌ์ดํด ์ค‘์— ์ƒ์„ฑ๋œ๋‹ค.์‹œ์ž‘ํ•˜๊ธฐ ์œ„ํ•ด, NADH๋Š” 2 ๊ฐœ์˜ ์ „์ž๋ฅผ ๋ณตํ•ฉ์ฒด 1๋กœ์ „๋‹ฌํ•˜์—ฌ NADH๋ฅผ NAD +๋กœ ์‚ฐํ™”์‹œํ‚จ๋‹ค.์ด๋“ค ์ „์ž๋Š” ๋ณด์กฐ ์ธ์ž ํ”Œ๋ผ ๋นˆ ๋ชจ๋…ธ ๋‰ดํด๋ ˆ์˜คํƒ€์ด๋“œ๋˜๋Š” FMN์œผ๋กœ ์ „๋‹ฌ๋œ๋‹ค.๊ทธ ๋‹ค์Œ ์ „์ž๊ฐ€ ์ฒ  – ํ™ฉ ๋‹จ๋ฐฑ์งˆ๋กœ์ „๋‹ฌ๋จ์— ๋”ฐ๋ผ ์‚ฐํ™”๋œ๋‹ค.๊ทธ๋Ÿฐ ๋‹ค์Œ์ด ํด๋Ÿฌ์Šคํ„ฐ๋Š” ์ „์ž๋ฅผ ์บ๋ฆฌ์–ด๋ถ„์ž ์ธ ์œ ๋น„ ํ€ด๋†€ ๋˜๋Š” Q๋กœ ์ „๋‹ฌํ•ฉ๋‹ˆ๋‹ค.์ด ์–‘์ž์ ์€ ์ „์ž๋ฅผ๋ณตํ•ฉ์ฒด 3์— ์ „๋‹ฌํ•˜๋ฉด์„œ ๋‘ ๊ฐœ์˜ ์–‘์„ฑ์ž๋ฅผ ์„ญ์ทจํ•ฉ๋‹ˆ๋‹ค.์—๋„ˆ์ง€ ๋ฐฉ์ถœ์˜ ๊ฒฐ๊ณผ๋กœ4 ๊ฐœ์˜ ์–‘์„ฑ์ž๊ฐ€ ๋ณตํ•ฉ์ฒด 1์„ ํ†ตํ•ด๋ง‰ ๊ฐ„ ๊ณต๊ฐ„์œผ๋กœ ๋Šฅ๋™์ ์œผ๋กœ ํŽŒํ•‘๋˜์–ด ๋‚ด๋ง‰์„ํ†ต๊ณผํ•˜๋Š” ์–‘์„ฑ์ž ๊ธฐ์šธ๊ธฐ๋ฅผ ์ƒ์„ฑํ•ฉ๋‹ˆ๋‹ค.FADH2๋Š” FADH2๋ฅผ FAD +๋กœ ์‚ฐํ™”์‹œํ‚ค๋Š” 2 ๊ฐœ์˜ ์ „์ž๋ฅผ์ง์ ‘ ๋ณตํ•ฉ๋ฌผ 2์— ์ „๋‹ฌํ•ฉ๋‹ˆ๋‹ค.์ด ์ „์ž๋Š” ๋˜ ๋‹ค๋ฅธ์ฒ  – ํ™ฉ ๋‹จ๋ฐฑ์งˆ๋กœ ์ „๋‹ฌ ๋œ ๋‹ค์Œ ์บ๋ฆฌ์–ดQ๋กœ ์ด๋™ํ•˜๋ฉฐ,์ด Q๋Š” ๋ฏธํ† ์ฝ˜๋“œ๋ฆฌ์•„๊ธฐ์งˆ์—์„œ ์ „์ž๋ฅผ ๋ณตํ•ฉ๋ฌผ 3์œผ๋กœ์šด๋ฐ˜ํ•˜๋ฉด์„œ ๋‘ ๊ฐœ์˜์–‘์„ฑ์ž๋ฅผ ์„ญ์ทจํ•œ๋‹ค.์„ธ ๋ฒˆ์งธ ์ฝคํ”Œ๋ ‰์Šค์—๋Š” Q ์‚ฌ์ดํด์ด๋ผ๊ณ ํ•˜๋Š”์ผ๋ จ์˜ ์ „์ž ์ „๋‹ฌ์ด ์žˆ์Šต๋‹ˆ๋‹ค.๋จผ์ €, ์ „์ž๊ฐ€ Q์—์„œ์ฒ  – ํ™ฉ ๋‹จ๋ฐฑ์งˆ๋กœ ์˜ฎ๊ฒจ์ง„๋‹ค.๊ทธ๋Ÿฐ ๋‹ค์Œ Q์— ์˜ํ•ด ์šด๋ฐ˜ ๋œ ๋‘ ๊ฐœ์˜ ์–‘์„ฑ์ž๊ฐ€๋ง‰๊ฐ„ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ•‘๋ฉ๋‹ˆ๋‹ค.์‹œํ† ํฌ๋กฌ c1์ด๋ผ ๋ถˆ๋ฆฌ๋Š”์ค‘๊ฐ„ ์‹œํ† ํฌ๋กฌ ๋ถ„์ž๋ฅผ ํ†ต๊ณผ ํ•œ ํ›„,์ „์ž๋Š” ์‹œํ† ํฌ๋กฌ C ์ „์ž ์ „๋‹ฌ์ฒด๋ฅผํ†ต๊ณผํ•˜์—ฌ ์ „์ž ์ „๋‹ฌ์ฒด๋ฅผ ๊ฐ์†Œ์‹œํ‚จ๋‹ค.๋‹ค์Œ์œผ๋กœ, Q์— ์˜ํ•ด ์šด๋ฐ˜ ๋œ ๋‘ ๋ฒˆ์งธ ์ „์ž๋Š”์‹œํ† ํฌ๋กฌ b ๋ณตํ•ฉ์ฒด๋กœ ์ „๋‹ฌ ๋œ ๋‹ค์Œ Q ๋ถ„์ž๋กœ์ „๋‹ฌ๋˜์–ด Q ๋ถ„์ž์—์„œ ๋งคํŠธ๋ฆญ์Šค๋กœ๋ถ€ํ„ฐ๋‘ ๊ฐœ์˜ ์–‘์„ฑ์ž๋ฅผ ๊ฒฐํ•ฉํ•ฉ๋‹ˆ๋‹ค.์ด์ œ ๋˜ ๋‹ค๋ฅธ Q ๋ถ„์ž๊ฐ€ ๋ณตํ•ฉ์ฒด 3์„ ๊ฒฐํ•ฉ์‹œํ‚ค๊ณ  ์ฃผ๊ธฐ์˜์ฒซ ๋ฒˆ์งธ ๋ถ€๋ถ„์ด ๋ฐ˜๋ณต๋˜์–ด ๋‘ ๊ฐœ์˜ ์–‘์„ฑ์ž๊ฐ€๋ฉค๋ธŒ๋ ˆ์ธ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ•‘๋˜์–ด Q ์‚ฌ์ดํด ๋‹น์ด 4 ๊ฐœ์˜ ์–‘์„ฑ์ž๊ฐ€ ์ƒ์„ฑ๋ฉ๋‹ˆ๋‹ค.์ƒˆ๋กญ๊ฒŒ ๊ฒฐํ•ฉ ๋œ Q ๋ถ„์ž์˜ ๋‘ ๋ฒˆ์งธ ์ „์ž๋Š”์‹œํ† ํฌ๋กฌ b๋กœ ์ด๋™ ํ•œ๋‹ค์Œ ์ด์ „์— ์ „์ž๋ฅผ๋ฐ›์€Q ๋ถ„์ž๋กœ ์ด๋™ํ•ฉ๋‹ˆ๋‹ค.์ด์ œ์ด Q๋Š” 2 ๊ฐœ์˜ ์ „์ž๋ฅผ ๊ฐ€์ง€๋ฉฐ,์ด๊ฒƒ์€ ๋ณตํ•ฉ์ฒด 3์—์„œ ๋ฐฉ์ถœ๋˜์–ด์ƒˆ๋กœ์šด Q ์‚ฌ์ดํด์—์„œ ์ „์ž๋ฅผ ๊ธฐ์ฆ ํ•  ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค.๋งˆ์ง€๋ง‰์œผ๋กœ, ๋ณตํ•ฉ์ฒด 4์— ๋ถ€์ฐฉ ๋œ ์‹œํ† ํฌ๋กฌC ์ „์ž ์บ๋ฆฌ์–ด์™€ 2 ๊ฐœ์˜์ „์ž๊ฐ€ ์‹œํ† ํฌ๋กฌ a3 ๋ถ„์ž์™€๊ตฌ๋ฆฌ ์›์ž๋ฅผ ๊ฐ์†Œ์‹œ์ผœ์‚ฐ์†Œ ๋ถ„์ž๋ฅผ ๊ฒฐํ•ฉ์‹œํ‚ต๋‹ˆ๋‹ค.์‚ฐ์†Œ ๋ถ„์ž๊ฐ€ ์™„์ „ํžˆ ํ™˜์›๋˜๋ฉด,4 ๊ฐœ์˜ ์ˆ˜์†Œ ์ด์˜จ๊ณผ ์Šคํ”Œ๋ฆฟ์ด ์ง‘์–ด ๋“ค๊ณ 2 ๊ฐœ์˜ ๋ฌผ ๋ถ„์ž๋ฅผ ํ˜•์„ฑํ•ฉ๋‹ˆ๋‹ค.์ด ๊ณผ์ •์—์„œ ๋„ค ๊ฐœ์˜์–‘์„ฑ์ž๊ฐ€ ๋ฉค๋ธŒ๋ ˆ์ธ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ•‘๋ฉ๋‹ˆ๋‹ค.๋”ฐ๋ผ์„œ ์ „์ž ์ˆ˜์†ก ์‚ฌ์Šฌ์€ ์–‘์„ฑ์ž๋ฅผ๋ฏธํ† ์ฝ˜๋“œ๋ฆฌ์•„์˜ ๋ง‰๊ฐ„ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ•‘ํ•จ์œผ๋กœ์จ์–‘์„ฑ์ž ๊ธฐ์šธ๊ธฐ๋ฅผ ์ƒ์„ฑํ•œ๋‹ค.๊ทธ๋Ÿฐ ๋‹ค์Œ ์ด๋“ค ์–‘์„ฑ์ž๋Š” ATP ํ•ฉ์„ฑ ํšจ์†Œ๋ฅผํ†ตํ•ด ๋ฏธํ† ์ฝ˜๋“œ๋ฆฌ์•„ ๊ธฐ์งˆ๋กœ ๊ทธ๋ผ๋””์–ธํŠธ๋ฅผ ๋‹ค์‹œ ํ๋ฅด๊ฒŒํ•˜์—ฌํ™”ํ•™ ๋ณ€์ด๋กœ ์•Œ๋ ค์ง„ ๊ณผ์ •์—์„œ ATP๋ฅผ ์ƒ์„ฑํ•ฉ๋‹ˆ๋‹ค.์‚ฐํ™” ๋œ ์ „์ž ์บ๋ฆฌ์–ด๋Š” ๊ตฌ์—ฐ์‚ฐ ์ˆœํ™˜์œผ๋กœ๋˜๋Œ์•„์™€ ๋” ๋งŽ์€ ์ „์ž๋ฅผํฌ์ฐฉ ํ•  ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค.

8.6:

์ „์ž์ „๋‹ฌ๊ณ„

์„ธํฌํ˜ธํก(cellular respiration)์˜ ๋งˆ์ง€๋ง‰ ๋‹จ๊ณ„๋Š” ์‚ฐํ™”์  ์ธ์‚ฐํ™”(oxidative phosphorylation)์ธ๋ฐ, ์ด๋Š” (1) ์ „์ž์ „๋‹ฌ๊ณ„(electron transport chain)์™€ (2) ํ™”ํ•™์‚ผํˆฌ(chemiosmosis)๋กœ ๊ตฌ์„ฑ๋ฉ๋‹ˆ๋‹ค.

์ „์ž์ „๋‹ฌ๊ณ„๋Š” ์ง„ํ•ต์„ธํฌ(eukaryotic cell)์˜ ๋ฏธํ† ์ฝ˜๋“œ๋ฆฌ์•„ ๋‚ด๋ง‰๊ณผ ์›ํ•ต์„ธํฌ(prokaryotic cell)์˜ ์›ํ˜•์งˆ๋ง‰(plasma membrane)์— ์žˆ๋Š” ๋‹จ๋ฐฑ์งˆ๊ณผ ๊ธฐํƒ€ ์œ ๊ธฐ ๋ถ„์ž๋กœ ๊ตฌ์„ฑ๋˜์–ด ์žˆ์Šต๋‹ˆ๋‹ค. ์ „์ž์ „๋‹ฌ๊ณ„๋Š” ๋‘ ๊ฐ€์ง€ ์ฃผ์š” ๊ธฐ๋Šฅ์„ ํ•ฉ๋‹ˆ๋‹ค. ํ•˜๋‚˜๋Š” ํ™”ํ•™์‚ผํˆฌ ์ค‘ ATP๋ฅผ ์ƒ์„ฑํ•˜๋Š” ๋ฐ ์ด์šฉ๋˜๋Š” ์–‘์„ฑ์ž ๊ตฌ๋ฐฐ(proton gradient)๋ฅผ ์ƒ์„ฑํ•˜๋Š” ๊ฒƒ์ด๊ณ , ๋‹ค๋ฅธ ํ•˜๋‚˜๋Š” ํ•ด๋‹น๊ณผ์ •(glycolysis)๊ณผ ์‹œํŠธ๋ฅด์‚ฐํšŒ๋กœ(citric acid cycle)์— ์‚ฌ์šฉ๋˜๋Š” ์ „์ž์šด๋ฐ˜์ฒด(electron carrier; ์˜ˆ: NAD+์™€ FAD)๋ฅผ ์ƒ์„ฑํ•˜๋Š” ๊ฒƒ์ž…๋‹ˆ๋‹ค.

์ผ๋ฐ˜์ ์œผ๋กœ ์ „์ž์ „๋‹ฌ๊ณ„์˜ ๋ถ„์ž๋Š” ๋„ค ๊ฐœ์˜ ๋ณตํ•ฉ์ฒด(I, II, III, IV)๋กœ ๊ตฌ์„ฑ๋ฉ๋‹ˆ๋‹ค. ๋ถ„์ž๋Š” ์—ฌ๋Ÿฌ ๋ฒˆ์˜ ์‚ฐํ™”ํ™˜์›๋ฐ˜์‘(redox reaction)์„ ํ†ตํ•ด ์„œ๋กœ์—๊ฒŒ ์ „์ž๋ฅผ ์ „๋‹ฌํ•˜๋Š”๋ฐ, ์ด๋•Œ ์‚ฐํ™”ํ™˜์›๋ฐ˜์‘์€ ์ „์ž์ „๋‹ฌ๊ณ„๋ฅผ ํ†ตํ•ด ์ „์ž๋ฅผ ๋†’์€ ์—๋„ˆ์ง€ ๋ ˆ๋ฒจ์—์„œ ๋‚ฎ์€ ์—๋„ˆ์ง€ ๋ ˆ๋ฒจ๋กœ ์˜ฎ๊น๋‹ˆ๋‹ค. ์ด๋Ÿฐ ๋ฐ˜์‘์€ ๋ณตํ•ฉ์ฒด๊ฐ€ ๋‚ด๋ง‰์„ ๊ฐ€๋กœ์งˆ๋Ÿฌ(์ฆ‰ ๊ธฐ์งˆ(matrix)์—์„œ ๋ง‰๊ฐ„๊ณต๊ฐ„(intermembrane space)์œผ๋กœ) H+๋ฅผ ํŽŒํ”„์งˆํ•˜๋Š” ๋ฐ ์‚ฌ์šฉํ•˜๋Š” ์—๋„ˆ์ง€๋ฅผ ๋ฐฉ์ถœํ•ฉ๋‹ˆ๋‹ค. ๋”ฐ๋ผ์„œ ์ด๋Š” ๋‚ด๋ง‰ ์ „์ฒด์— ์–‘์„ฑ์ž ๊ตฌ๋ฐฐ๋ฅผ ํ˜•์„ฑํ•ฉ๋‹ˆ๋‹ค.

NADH์™€ FADH2๋Š” ์ด์ „ ์„ธํฌํ˜ธํก ๋‹จ๊ณ„์—์„œ ์ƒ์„ฑ๋˜์–ด ์ด๋ฏธ ํ™˜์›๋œ ์ƒํƒœ์˜ ์ „์ž์šด๋ฐ˜์ฒด์ž…๋‹ˆ๋‹ค. NADH๋Š” ๋ฐฉ์ถœ๋œ ์—๋„ˆ์ง€๋ฅผ ์‚ฌ์šฉํ•˜์—ฌ ์–‘์„ฑ์ž๋ฅผ ๋ง‰๊ฐ„ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ”„์งˆํ•˜๋Š” ๋ณตํ•ฉ์ฒดI์— ์ง์ ‘ ์ „์ž๋ฅผ ์ „๋‹ฌํ•  ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค. FADH2๋„ ์ „์ž๋ฅผ ๋ณตํ•ฉ์ฒดII์— ์ „๋‹ฌํ•˜์ง€๋งŒ, ์ด ๋ณตํ•ฉ์ฒด๋Š” ๋ณตํ•ฉ์ฒด ์ค‘ ์œ ์ผํ•˜๊ฒŒ ์–‘์„ฑ์ž๋ฅผ ๋ง‰๊ฐ„ ๊ณต๊ฐ„์œผ๋กœ ํŽŒํ”„์งˆํ•˜์ง€ ์•Š์Šต๋‹ˆ๋‹ค. ๋”ฐ๋ผ์„œ FADH2๋Š” NADH๋ณด๋‹ค ์–‘์„ฑ์ž ๊ตฌ๋ฐฐ์— ๋œ ๊ธฐ์—ฌํ•ฉ๋‹ˆ๋‹ค. ์ดํ›„ NADH์™€ FADH2๋Š” ๊ฐ๊ฐ ์ „์ž์šด๋ฐ˜์ฒด์ธ NAD+์™€ FAD๋กœ ๋ณ€ํ™˜๋ฉ๋‹ˆ๋‹ค.

NADH์™€ FADH2 ๋ชจ๋‘ ์ „์ž๋ฅผ ๋ณตํ•ฉ์ฒดIII์— ์ „๋‹ฌํ•˜๋Š” ์ด๋™ ์ „์ž์šด๋ฐ˜์ฒด(mobile electron carrier)์ธ ์œ ๋น„ํ€ด๋…ผ(ubiquinone)์— ์ „์ž๋ฅผ ์ „๋‹ฌํ•ฉ๋‹ˆ๋‹ค. ๊ฑฐ๊ธฐ์„œ๋ถ€ํ„ฐ ์ „์ž๋Š” ์ด๋™ ์ „์ž์šด๋ฐ˜์ฒด ์‹œํ† ํฌ๋กฌc (cytochrome c, ์ค„์—ฌ์„œ cyt c)๋กœ ์ „๋‹ฌ๋ฉ๋‹ˆ๋‹ค. cyt c๋Š” ์ „์ž๋ฅผ ๋ณตํ•ฉ์ฒดIV์— ๋ณด๋‚ด๊ณ , ๋ณตํ•ฉ์ฒดIV๋Š” ์ „์ž๋ฅผ O2์— ์ „๋‹ฌํ•ฉ๋‹ˆ๋‹ค. ๊ทธ๋Ÿฌ๋ฉด ์‚ฐ์†Œ๋Š” ๋ถ„ํ•ดํ•˜์—ฌ 2๊ฐœ์˜ ์‚ฐ์†Œ ์›์ž๋ฅผ ํ˜•์„ฑํ•˜๊ณ , ๊ฐ๊ฐ 2๊ฐœ์˜ ์–‘์„ฑ์ž๋ฅผ ์ˆ˜์šฉํ•˜์—ฌ ๋ฌผ์„ ํ˜•์„ฑํ•ฉ๋‹ˆ๋‹ค.

Suggested Reading

  1. Reece, J.B.et al. Campbell Biology. 10th ed. Pearson, London, UK (2014).
  2. Guo, R., Zong, S., Wu, M., Gu, J., Yang, M. Architecture of Human Mitochondrial Respiratory Megacomplex I2III2IV2. Cell. 170 (6), 1247-1257 (2017).
  3. Clark, M. A., Douglas, M., Choi, J. Section 7.4: Oxidative Phosphorylation. In Biology 2e. OpenStax. Houston, TX (2018).