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36.6:

์—ด๊ณผ ๋ƒ‰๊ธฐ ์ŠคํŠธ๋ ˆ์Šค์— ๋Œ€ํ•œ ๋ฐ˜์‘

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Responses to Heat and Cold Stress

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๊ทน๋‹จ์ ์ธ ์ฃผ์œ„ ์˜จ๋„๋Š” ์‹๋ฌผ์˜์‹ ์ง„๋Œ€์‚ฌ์— ์˜ํ–ฅ์„ ๋ฏธ์นœ๋‹ค์ง€๋‚˜์นœ ์—ด์€ ํšจ์†Œ์™€ ๋‹ค๋ฅธ ๋‹จ๋ฐฑ์งˆ์„ ๋ณ€์„ฑ์‹œํ‚ค๊ณ ๊ทน๋‹จ์ ์ธ ์ถ”์œ„๋Š” ์„ธํฌ๋‚ด ์ˆ˜๋ถ„์„ ์–ผ๊ฒŒ ํ•œ๋‹ค์‹๋ฌผ์€ ์–ด๋–ป๊ฒŒ ์—ด๊ณผ ์ถ”์œ„ ์ŠคํŠธ๋ ˆ์Šค์— ๋ฐ˜์‘ํ• ๊นŒ?์ฃผ์œ„ ์˜จ๋„๊ฐ€ ๋†’์œผ๋ฉด์‹๋ฌผ์€ ์ง€๋‚˜์นœ ๋ฌผ ์†์‹ค์„ ํ”ผํ•˜๋ ค๊ณ ๋‚ฎ์—๋Š” ๊ธฐ๊ณต์„ ๋‹ซ์•„๋ฒ„๋ฆฌ๋Š”๋ฐ์ข…์ข… ๊ทธ๋กœ ์ธํ•ด CO2 ํก์ˆ˜์™€ ๊ด‘ํ•ฉ์„ฑ์„ ๋ชปํ•˜๊ฒŒ ๋œ๋‹ค์—ด ์ŠคํŠธ๋ ˆ์Šค๋Š” ๋˜ํ•œ ์‹๋ฌผ ์„ธํฌ๊ฐ€๋งŽ์€ ์–‘์˜ ํŠน์ˆ˜ ๋‹จ๋ฐฑ์งˆ์„ ํ•ฉ์„ฑํ•˜๊ฒŒ ๋งŒ๋“œ๋Š”๋ฐ๋ฐ”๋กœ ์—ด์ถฉ๊ฒฉ๋‹จ๋ฐฑ์งˆ์ด๋‹ค์ด๋“ค์€ ๋ณดํ˜ธ์ž ์—ญํ• ์„ ํ•˜๋ฉด์„œ ๋‹ค๋ฅธ ๋‹จ๋ฐฑ์งˆ์ด๊ธฐ๋Šฅ์  ๋ชจ์–‘์œผ๋กœ ์ ‘ํžˆ๊ฒŒ ๋„์™€์ฃผ๊ฑฐ๋‚˜ํšจ์†Œ์™€ ๋‹จ๋ฐฑ์งˆ์ด ๋ณ€์„ฑ๋˜์ง€ ์•Š๋„๋ก ๋ณดํ˜ธํ•ด์ค€๋‹ค์‹๋ฌผ์€ ์„ธํฌ๋ง‰์˜ ์ง€์งˆ ๊ตฌ์„ฑ์„ ์กฐ์ ˆํ•˜์—ฌ์˜จ์ „ํ•จ์„ ์œ ์ง€ํ•˜๊ณ  ์ตœ์  ๋ง‰ ์œ ๋™์„ฑ์„ ์œ ์ง€ํ•˜๋Š”๋ฐ์ด๋Š” ์—ด๊ณผ ์ถ”์œ„ ์ŠคํŠธ๋ ˆ์Šค์— ๋Œ€ํ•œ ๋ฐ˜์‘์ด๋‹ค๋ง‰ ์œ ๋™์„ฑ์€ ๋ง‰์˜ ํˆฌ๊ณผ์„ฑ์— ์˜ํ–ฅ์„ ๋ฏธ์ณ์„œ๋ง‰์„ ํ†ต๊ณผํ•˜๋Š” ๋ถ„์ž์˜ ์›€์ง์ž„์„ ๊ทœ์ œํ•˜๊ณ ์„ธํฌ ์•ˆํŒŽ์œผ๋กœ ์ƒˆ๋Š” ๊ฒƒ์„ ๋ฐฉ์ง€ํ•œ๋‹ค์ธ์ง€์งˆ์€ ์ด์ค‘์ธต์œผ๋กœ ๋ฐฐ์—ด๋˜์–ด์›ํ˜•์งˆ ๋ง‰์˜ ๊ธฐ๋ณธ ๊ตฌ์กฐ๋ฅผ ํ˜•์„ฑํ•œ๋‹ค์ด ์ด์ค‘์ธต์˜ ์ง€๋ฐฉ์„ฑ๋ถ„์€ํฌํ™” ํ˜น์€ ๋ถˆํฌํ™” ์ง€๋ฐฉ์‚ฐ์œผ๋กœ ๊ตฌ์„ฑ๋˜์–ด ์žˆ๋‹ค์—ด ์ŠคํŠธ๋ ˆ์Šค ๋™์•ˆ์— ๋†’์€ ์˜จ๋„๋Š”์ง€๋ฐฉ ์ด์ค‘์ธต์ด ๋” ์œ ๋™์ ์ด๊ณ , ๋” ํˆฌ๊ณผํ•˜๊ฒŒ ํ˜น์€๋” ์ƒˆ๊ฒŒ ๋งŒ๋“ ๋‹ค์‹๋ฌผ์˜ ๋ฐ˜์‘์€ ๋ง‰์˜ ํฌํ™” ์ง€๋ฐฉ์‚ฐ ๋น„์œจ์„ ์ฆ๊ฐ€์‹œ์ผœ์„œ์—ด ์ €ํ•ญ์„ ํ–ฅ์ƒ์‹œํ‚ค๊ณ ๋ง‰์˜ ์œ ๋™ํ™”๋ฅผ ๋ง‰์•„์ค€๋‹ค์ถ”์œ„ ์ŠคํŠธ๋ ˆ์Šค ๋™์•ˆ์—๋Š” ๋‚ฎ์€ ์˜จ๋„๊ฐ€์ง€๋ฐฉ ์ด์ค‘์ธต์„ ๋” ๊ฒฌ๊ณ ํ•˜๊ฒŒ ๋งŒ๋“ค์–ด์„œํˆฌ๊ณผ์„ฑ์„ ๊ฐ์†Œ์‹œํ‚จ๋‹ค๋ฐ˜์‘์œผ๋กœ, ๋ง‰์— ์žˆ๋Š” ๋ถˆํฌํ™” ์ง€๋ฐฉ์‚ฐ ๋น„์œจ์ด๋Š˜์–ด๋‚˜์„œ ๋ง‰์˜ ๊ฒฝ์ง์„ฑ์ด ์ค„์–ด๋“ค๊ณ ์ตœ์ ์˜ ์œ ๋™์„ฑ์„ ์œ ์ง€ํ•˜๊ฒŒ ๋œ๋‹ค๋น™์ ํ•˜์˜ ์˜จ๋„์—์„œ ๋Œ€๋ถ€๋ถ„ ์‹๋ฌผ์˜ ์„ธํฌ๋ฒฝ๊ณผ์„ธํฌ๊ฐ„ ๊ณต๊ฐ„์— ์ƒ๊ธฐ๋Š” ์–ผ์Œ์€๋ฌผ์ด ์„ธํฌ์งˆ์„ ๋– ๋‚˜๊ฒŒ ๋งŒ๋“ค์–ด์„œ์„ธํฌ์งˆ ํƒˆ์ˆ˜๋ฅผ ์ผ์œผํ‚จ๋‹ค์ด๋ฅผ ๋ง‰๊ธฐ ์œ„ํ•˜์—ฌ ๋งŽ์€ ์„œ๋ฆฌ ๋‚ด์„ฑ ์‹๋ฌผ๋“ค์€๋‹น๊ณผ ๊ฐ™์€ ์šฉ์งˆ์„์„ธํฌ์งˆ์— ์ถ•์ ํ•˜์—ฌ ์‚ผํˆฌ ์ž ์žฌ๋ ฅ์„ ๊ทœ์ œํ•œ๋‹ค์—ด๊ณผ ์ถ”์œ„ ๋ฐ˜์‘์˜ ์ ์‘ ๊ธฐ์ œ๋Š”ํ•ญ์ƒ์„ฑ ์œ ์ง€๋ฅผ ๋„์™€์ฃผ๊ณ  ์‹๋ฌผ์˜ ์ƒ์กด์„ ํ™•์‹คํ•˜๊ฒŒ ํ•ด์ค€๋‹ค

36.6:

์—ด๊ณผ ๋ƒ‰๊ธฐ ์ŠคํŠธ๋ ˆ์Šค์— ๋Œ€ํ•œ ๋ฐ˜์‘

๋ชจ๋“  ์œ ๊ธฐ์ฒด๋Š” ๊ฑด๊ฐ•ํ•œ ์„ฑ์žฅ๊ณผ ์ƒ๋ฆฌ์  ๊ธฐ๋Šฅ์„ ํ•  ์ˆ˜ ์žˆ๋Š” ์ตœ์ ์˜ ์˜จ๋„ ๋ฒ”์œ„๋ฅผ ๊ฐ€์ง€๊ณ  ์žˆ์Šต๋‹ˆ๋‹ค. ์ด ๋ฒ”์œ„์˜ ๋์—๋Š” ์ƒ๋ฌผํ•™์  ๊ณผ์ •์„ ๋ฐฉํ•ดํ•˜๋Š” ์ตœ์ € ์˜จ๋„์™€ ์ตœ๊ณ  ์˜จ๋„๊ฐ€ ์กด์žฌํ•ฉ๋‹ˆ๋‹ค.

ํ™˜๊ฒฝ ์—ญํ•™์ด ์ฃผ์–ด์ง„ ์ข…์ด ๊ฐ€์ง„ ์ตœ์ ์˜ ๋ฒ”์œ„๋ฅผ ๋ฒ—์–ด๋‚  ๋•Œ, ๋ฌผ์งˆ๋Œ€์‚ฌ์™€ ๊ธฐ๋Šฅ์— ๋ณ€ํ™”๊ฐ€ ์ด๋Š”๋ฐ, ์ด๋ฅผ ์ŠคํŠธ๋ ˆ์Šค๋ผ ์ •์˜ํ•ฉ๋‹ˆ๋‹ค. ์‹๋ฌผ์€ ์œ ์ „์ž ๋ฐœํ˜„์— ๋ณ€ํ™”๋ฅผ ์ผ์œผ์ผœ ์ŠคํŠธ๋ ˆ์Šค์— ๋ฐ˜์‘ํ•ฉ๋‹ˆ๋‹ค. ์‹๋ฌผ์˜ ์ŠคํŠธ๋ ˆํŠธ ๋ฐ˜์‘์€ ์‹๋ฌผ ๋ฌผ์งˆ๋Œ€์‚ฌ์™€ ํ•ญ์ƒ์„ฑ(homeostasis)์„ ์–ป๊ธฐ ์œ„ํ•œ ๋ฐœ์œก ์กฐ์ •์œผ๋กœ ์ด์–ด์ง‘๋‹ˆ๋‹ค.

์‹๋ฌผ์€ ์˜จ๋„ ๋ณ€๋™ ์‹œ ๋ง‰ ์œ ๋™์„ฑ์„ ์œ ์ง€ํ•ฉ๋‹ˆ๋‹ค

์‹๋ฌผ์˜ ์„ธํฌ๋ง‰์€ ๋ณดํ†ต ์ฃผ์œ„ ์˜จ๋„ ๋ณ€ํ™”์— ์˜ํ•ด ์˜ํ–ฅ์„ ๋ฐ›๋Š” ์ฒซ ๋ฒˆ์งธ ๊ตฌ์กฐ ์ค‘ ํ•˜๋‚˜์ž…๋‹ˆ๋‹ค. ์‹๋ฌผ์˜ ์„ธํฌ๋ง‰์€ ์ฃผ๋กœ ์ธ์‚ฐ, ์ฝœ๋ ˆ์Šคํ…Œ๋กค, ๊ทธ๋ฆฌ๊ณ  ๋‹จ๋ฐฑ์งˆ๋กœ ๊ตฌ์„ฑ๋˜๊ณ , ์ง€์งˆ ๋ถ€๋ถ„์€ ๋ถˆํฌํ™” ์ง€๋ฐฉ์‚ฐ ๋˜๋Š” ํฌํ™” ์ง€๋ฐฉ์‚ฐ์˜ ๊ธด ์‚ฌ์Šฌ๋กœ ๊ตฌ์„ฑ๋ฉ๋‹ˆ๋‹ค. ์˜จ๋„ ๋ณ€ํ™” ํ•˜์—์„œ ์‹๋ฌผ๋“ค์ด ์„ ํƒํ•  ์ˆ˜ ์žˆ๋Š” ์ฃผ์š” ์ „๋žต ์ค‘ ํ•˜๋‚˜๋Š” ์„ธํฌ๋ง‰์˜ ์ง€์งˆ ์„ฑ๋ถ„์„ ๋ฐ”๊พธ๋Š” ๊ฒƒ์ž…๋‹ˆ๋‹ค. ์‹๋ฌผ์€ ๊ณ ์˜จ์—์„œ ๋ง‰ ์ง€์งˆ์˜ ๋ถˆํฌํ™” ์ •๋„๋ฅผ ๊ฐ์†Œ์‹œํ‚ค๊ณ  ์ €์˜จ์—์„  ์ฆ๊ฐ€ ์‹œ์ผœ ๋ง‰์˜ ์œ ๋™์„ฑ์„ ์œ ์ง€ํ•ฉ๋‹ˆ๋‹ค.

์—ด์ถฉ๊ฒฉ๋‹จ๋ฐฑ์งˆ(heat shock protein)

์‹๋ฌผ ์กฐ์ง์ด๋‚˜ ์„ธํฌ๊ฐ€ ๊ฐ‘์ž‘์Šค๋Ÿฌ์šด ๊ณ ์˜จ ์ŠคํŠธ๋ ˆ์Šค์— ๋…ธ์ถœ๋˜๋ฉด ์—ด์ถฉ๊ฒฉ๋‹จ๋ฐฑ์งˆ(heat shock protein, ์ค„์—ฌ์„œ HSP)์ด ์ผ์‹œ์ ์œผ๋กœ ๋ฐœํ˜„๋ฉ๋‹ˆ๋‹ค. ์—ด์ถฉ๊ฒฉ๋‹จ๋ฐฑ์งˆ์€ ์ƒคํŽ˜๋ก (chaperone)์œผ๋กœ์„œ ํ•„์ˆ˜ ์ƒ๋ฆฌ ๊ธฐ๋Šฅ์„ ์ˆ˜ํ–‰ํ•˜๊ฑฐ๋‚˜, ๋ณ€์„ฑ๋œ ๋‹จ๋ฐฑ์งˆ์˜ ์‘์ง‘์„ ๋ฐฉ์ง€ํ•˜๊ฑฐ๋‚˜, ์‘์ง‘๋œ ๋‹จ๋ฐฑ์งˆ ๋ถ„์ž์˜ ์›ํ˜•ํšŒ๋ณต์„ ์ด‰์ง„ํ•ฉ๋‹ˆ๋‹ค.

๊ธฐ๊ณต ์ „๋„๋„

์ผ์˜จ๋„๊ฐ€ ์ผ๋ฐ˜์ ์ธ ํ‰๊ท  ๋ฒ”์œ„ ์ด์ƒ์œผ๋กœ ์ฆ๊ฐ€ํ•˜๋ฉด ์‹๋ฌผ์˜ ๊ด‘ํ•ฉ์„ฑ ํ™œ๋™๊ณผ ๊ธฐ๊ณต ์ƒ๋ฆฌ์— ์˜ํ–ฅ์„ ๋ฏธ์นฉ๋‹ˆ๋‹ค. ์˜จ๋„๊ฐ€ ์ƒ์Šนํ•จ์— ๋”ฐ๋ผ, ์‹๋ฌผ์€ ๊ธฐ๊ณต์„ ๋‹ซ์•„ ๊ธฐ๊ณต ์ „๋„ ๋ฐ ์ฆ์‚ฐ์œผ๋กœ ์ธํ•œ ์ˆ˜๋ถ„ ์†์‹ค์„ ์ค„์ผ ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค.

์‹๋ฌผ ์„ธํฌ ๋‚ด ์šฉ์งˆ ์ถ•์ 

๊ทน๋„๋กœ ๋‚ฎ์€ ์˜จ๋„๋Š” ๋‚ฎ์€ ์ˆ˜๋ถ„ํผํ…์…œ(water potential; ์ˆ˜๋ถ„ํฌํ…์…œ)๋กœ ์ธํ•ด ์ˆ˜๋ถ„ ํก์ˆ˜๋ฅผ ๊ฐ์†Œ ์‹œ์ผœ ํƒˆ์ˆ˜๋ฅผ ์œ ๋ฐœํ•  ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค. ์—ฌ๋Ÿฌ ์‹๋ฌผ์€ ๊ทธ๋“ค์˜ ์‚ผํˆฌํผํ…์…œ(osmotic potential)์„ ์กฐ์ ˆํ•˜๊ณ  ๋‹น๊ณผ ๊ฐ™์€ ์šฉ์งˆ(์ˆ˜ํฌ๋กœ์Šค, ํฌ๋„๋‹น, ๊ณผ๋‹น)์˜ ์„ธํฌ ๋‚ด ์ถ•์ ์„ ํ†ตํ•ด ์ˆ˜๋ถ„ ํ•จ๋Ÿ‰์„ ์œ ์ง€ํ•ฉ๋‹ˆ๋‹ค. ์ด๋Ÿฌํ•œ ์šฉ์งˆ์˜ ์ถ•์  ๋˜ํ•œ ๋™๊ฒฐ์ ์„ ๋‚ฎ์ถฐ ์กฐ์ง์˜ ๋ฌผ ๋™๊ฒฐ์„ ์ง€์—ฐ์‹œํ‚ฌ ์ˆ˜ ์žˆ์Šต๋‹ˆ๋‹ค.

Suggested Reading

Nievola, Catarina C, Camila P Carvalho, Victória Carvalho, and Edson Rodrigues. "Rapid Responses of Plants to Temperature Changes." Temperature. 4 (4)2017: 371–405. [Source]

Zheng, Guowei, Bo Tian, Fujuan Zhang, Faqing Tao, and Weiqi Li. "Plant Adaptation to Frequent Alterations between High and Low Temperatures: Remodeling of Membrane Lipids and Maintenance of Unsaturation Levels." Plant, Cell & Environment. 34 (9)2011: 431–1442. [Source]

Tarkowski, ลukasz P., and Wim Van den Ende. "Cold tolerance triggered by soluble sugars: a multifaceted countermeasure." Frontiers in plant science 6 (2015): 203. [Source]