남세균

3. 분류학

현재 알려져 있는 남세균은 약 150속 2,000종이다. 이들은 바닷물이나 민물뿐만 아니라 토양 속, 나무 줄기 위 등 다양한 환경에서 서식한다. 심지어 눈 속이나 80°C 이상의 온천에서도 발견될 정도로, 물이 있는 곳이라면 어디에서든 생존할 수 있다. 특이하게도 같은 종류가 한대, 온대, 열대 지역에 모두 분포하는 경우도 있다.^{[225][226][227]}

남세균은 다음과 같은 특징을 갖는다.

• 세포 내에 핵, 엽록체, 미토콘드리아, 액포 등의 세포 기관이 분화되어 있지 않다.
• 엽록소 a와 남조소, 홍조소 등의 피코빌린 색소를 가지고 광합성을 한다.
• 광합성 결과 생성되는 저장 물질은 남조 녹말이다.
• 세포벽은 뮤코펩티드라는 화합물로 구성되며, 세포벽 바깥쪽에는 점액질 층이 있다.
• 무성 생식만 하며, 유성 생식은 알려져 있지 않다.
• 생식 세포에 편모가 없어 헤엄칠 수 없다.

• 남구슬말목(Chroococcales) 또는 소구체목
• Chroococcidiopsidales
• Gloeobacterales
• 구슬말목(Nostocales) 또는 줄구슬말목, 염주말목
• 흔들말목(Oscillatoriales)
• 플레우로캅사목(Pleurocapsales) 또는 굳은막말목
• Spirulinales
• Synechococcales

3. 1. 하위 분류

남세균의 분류에서는 속(Genus) 수준에서도 비단일계통성이 나타나는 경우가 많다.^[525] 이러한 속의 일부는 게놈 수준의 분자 계통 분석에 기반하여 다수의 속으로 세분화하는 것이 제안되고 있다.^[521][526]

4. 형태

남세균은 단세포, 사상, 군체 등 다양한 형태를 띤다. 사상 형태는 이형세포 (질소 고정), 아키네트 (휴면 단계 세포), 호르모고니아 (생식성, 운동성 사상)와 같은 기능적 세포 분화를 나타낸다.^{[43][44][45][125]}

• 영양 세포 - 유리한 성장 조건에서 형성되는 정상적인 광합성 세포
• 아키네트 - 환경 조건이 악화될 때 형성될 수 있는 기후 저항성 포자
• 두꺼운 벽 이형세포 - 질소 고정에 필수적인 효소 질소 고정 효소를 포함한다.^[49][50][51] 산소에 대한 민감성으로 인해 혐기성 환경에서^[51] 생성된다.

• '''단세포성''' (unicellular): 몸이 1개의 세포로 이루어져 있다. 세포의 형태는 구형이나 간상인 것이 많고(예: Synechococcus, Synechocystis), 또한 이극성(heteropolarity; 기단과 선단에서 형태가 다름)을 나타내는 종도 있다(예: Chamaesiphon).^[289]

• '''군체성''' (colonial): 몸이 복수의 세포로 이루어져 있지만, 세포가 밀접하지 않고, 세포 분화가 보이지 않는 등 다세포라고 부르기 어려운 것이다(명확한 정의는 어렵다). 군체 전체의 형태는 다양하다(부정형, 구형, 다면체, 시트 모양, 끈 모양 등).^[289] 또한 군체 양식으로는, 다수의 세포가 공통의 점액질에 싸인 팔메라상 군체(palmelloid colony; 예: Aphanocapsa)가 많지만, 그 외에도 세포가 밀착하여 덩어리 모양이 되는 사르시나상 군체(sarcinoid colony; 예: Cyanosarcina)나, 분기하는 점액질 자루의 선단에 각각 세포가 위치하는 수지상 군체(dendroid colony) 등이 있다.^[289]

• '''사상성''' (filamentous):

5. 질소 고정

일부 남세균은 이형세포라는 특수한 세포를 통해 혐기성 조건에서 대기 중의 질소를 고정할 수 있다.^[50][51] 이형세포는 고정 질소가 부족할 때 적절한 환경 조건(무산소)에서 형성될 수도 있다. 이형세포를 형성하는 종은 질소 고정에 특화되어 있으며, 질소 가스를 암모니아(NH₃), 아질산염(NO₂^-) 또는 질산염(NO₃^-)으로 고정할 수 있다. 이는 식물이 흡수하여 단백질과 핵산으로 전환될 수 있다(대기 중의 질소는 내생 공생 질소 고정 세균을 가진 식물, 특히 콩과를 포함한 일부 식물을 제외하고는 식물에게 생물학적 이용 가능하지 않다).

6. 광합성

남세균은 지구 역사상 최초로 광합성을 시작한 생물이다. 이들은 이산화 탄소, 메테인과 같은 온실 기체를 이용하여 산소를 만들어 냈다. 남세균의 광합성은 지금도 바다에 산소를 공급하고 있으며, 대기 산소 농도를 급증시켜 오존층을 형성하고 지표에 도달하는 자외선을 차단하여 해상 생물이 육상 생물로 진화하는 결과로 이어졌다.^[76]

6. 1. 탄소 고정

6. 2. 전자 전달

6. 2. 1. 전자 전달 사슬

6. 2. 2. 대사

7. 생태

남세균은 해양, 담수, 습한 토양, 사막, 극지방 등 거의 모든 환경에서 서식한다.^[85] 플랑크톤 세포로 존재하거나 광합성 바이오필름을 형성하며, 일부는 다른 생물과 공생 관계를 유지한다.^[85] 특히, 수생 남세균은 조류 번식(algal bloom)을 일으켜 수질 오염 문제를 야기하기도 한다.^[87] 남세균의 성장은 잔잔하고 따뜻한 물에서 활발하며,^[88] 높은 온도와 부영양화는 유해 남조류 대발생(녹조 현상)의 주요 원인으로 작용한다.^[88]

남세균은 매우 크고 다양한 문에 속하는 광합성 원핵생물이다.^[19] 이들은 독특한 생물학적 색소 조합과 산소 발생 광합성을 수행하는 능력을 가지고 있다.^[19] 종종 다양한 형태를 취할 수 있는 군체 집합체에 살기도 한다.^[159] 특히 사상 남세균은 온천, 고염수, 사막 및 극지방과 같은 극한 환경뿐만 아니라 일반적인 환경에서도 널리 분포한다.^[20][21] 일부 종은 질소 고정을 하며 다양한 습한 토양과 물에서 자유롭게 또는 식물이나 지의류를 형성하는 균류와 공생 관계를 유지한다.^[23]

남세균은 전 세계적으로 널리 퍼져 있으며, 전 세계적인 생지화학적 순환에 주요 기여를 한다.^[125] 이들은 유일한 산소 발생 광합성 원핵생물이며, 다양하고 극한 환경에서 번성한다.^[24] 지구에서 가장 오래된 생물 중 하나이며, 화석 기록은 최소 21억 년 전으로 거슬러 올라간다.^[171] 플랑크톤성 남세균은 해양 먹이 그물의 기본적인 구성 요소이며, 전 세계적인 탄소 순환과 질소 순환 흐름에 주요 기여를 한다.^[25][26] 일부 남세균은 유해 조류 번식을 형성하여 수생 생태계 서비스의 교란을 일으키고, 마이크로시스틴, 삭시톡신, 실린드로스퍼몹신과 같은 강력한 독소(시아노톡신)를 생성하여 야생 동물과 인간을 중독시킨다.^[27][28] 오늘날 남세균 번식은 수생 환경과 공중 보건에 심각한 위협이 되고 있으며, 전 세계적으로 빈도와 규모가 증가하고 있다.^[120][125]

해양 환경에서 남세균은 1차 생산자로서 중요한 역할을 하며, 해양 식물 플랑크톤의 일부로서 현재 지구 전체 1차 생산량의 거의 절반을 기여한다.^[29] 전 세계 해양 1차 생산량의 약 25%는 남세균에 의해 기여된다.^[30]

남세균 내에서, ''Crocosphaera''와 친척, 남세균 UCYN-A, ''Trichodesmium'', ''Prochlorococcus'', ''Synechococcus''와 같은 몇몇 계통만이 넓은 바다에 서식한다.^{[31][32][33][34]} 질소 고정 남세균은 질소 가스를 암모늄으로 변환하여 해양 1차 생산성과 심해로의 유기 탄소 수출을 통제하기 때문에 특히 중요하다. 암모늄은 나중에 아미노산과 단백질을 만드는 데 사용된다. 해양 피코남세균(''Prochlorococcus''와 ''Synechococcus'')은 현대 해양의 대부분의 식물 플랑크톤 집합체에서 수적으로 우위를 차지하며, 1차 생산성에 중요하게 기여한다.^[33][34][35] 일부 플랑크톤성 남세균은 단세포이며 자유 생활 세포(예: ''Crocosphaera'', ''Prochlorococcus'', ''Synechococcus'')이지만, 다른 것들은 햅토조류, 예를 들어 코콜리소포어와 공생 관계를 맺고 있다.^[32]

해양 남세균은 가장 작은 광합성 유기체를 포함한다. 가장 작은 것 중 하나인 ''Prochlorococcus''는 폭이 0.5µm~0.8µm에 불과하다.^[39] 개체 수 측면에서, ''Prochlorococcus''는 지구에서 가장 풍부한 속일 가능성이 있다. 표면 해수 1밀리리터는 이 속의 세포 100,000개 이상을 포함할 수 있으며, 전 세계적으로 여러 옥틸리언(10²⁷, 100경) 개체가 있는 것으로 추정된다.^[40] ''Prochlorococcus''는 위도 40°N과 40°S 사이에서 어디에나 존재하며, 해양의 빈영양(영양분이 부족한) 지역에서 우세하다.^[41] 이 박테리아는 지구 대기 중 산소의 약 20%를 차지한다.^[42]

남세균은 육상 서식지와 유기체 군집에서 중요한 역할을 하는 것으로 밝혀졌다. 남세균 토양 피막은 침식을 방지하고 물을 유지하기 위해 토양을 안정화하는 데 도움이 된다고 보고되었다.^[91]

7. 1. 시아노비온트 (Cyanobionts)

7. 2. 집단 행동

7. 3. 세포 사멸

7. 4. 남세균 파지 (Cyanophages)

8. 운동

오랫동안 사상 남세균은 표면 운동을 수행하며 이러한 움직임이 IV형 선모에서 비롯된다는 것이 알려져 왔다.^{[147][129][148]} 또한 해양 남세균인 ''Synechococcus''는 세균 편모와는 다른 메커니즘으로 25μm/s의 속도로 헤엄치는 것으로 알려져 있다.^[149] 남세균 표면에서 파도가 형성되어 주변 물을 뒤로 밀어내는 것으로 생각된다.^[150][151] 세포는 활공 방식으로 운동성을 갖는 것으로 알려져 있으며,^[152] 편모 운동과 관련이 없는 새롭고 특징이 없는 비광주성 수영 방식도 있다.^[153]

많은 남세균 종이 활공할 수 있다. 활공은 기어다니거나 헤엄치는 것과는 다른 형태의 세포 운동으로, 뚜렷한 외부 기관이나 세포 형태의 변화에 의존하지 않으며, 기질이 있는 경우에만 발생한다.^[154][155] 사상 남세균의 활공은 세포가 수화되면서 빠르게 팽창하는 젤을 배출하여 추진력을 제공하는 "점액 제트" 메커니즘에 의해 구동되는 것으로 보이지만,^[156][157] 일부 단세포 남세균은 활공을 위해 IV형 선모를 사용한다.^[158][21]

남세균은 엄격한 광 요구 사항을 가지고 있다. 빛이 너무 적으면 에너지 생성이 불충분해질 수 있으며, 일부 종에서는 세포가 종속 영양 호흡에 의존하게 될 수 있다.^[20] 빛이 너무 많으면 세포가 억제되고 광합성 효율이 감소하며 표백에 의한 손상을 일으킬 수 있다. UV 방사선은 남세균에 특히 치명적이며, 일반적인 태양 수준은 경우에 따라 이러한 미생물에 심각한 해를 끼칠 수 있다.^{[159][160][21]}

미생물 매트에서 사는 사상 남세균은 광합성을 위한 빛 요구 사항과 광손상에 대한 민감성을 균형 있게 유지하기 위해 매트 내에서 수직 및 수평으로 이동하는 경우가 많다. 예를 들어, 멕시코 게레로 네그로의 과염수 벤토스 매트에서 발견되는 사상 남세균인 ''Oscillatoria'' sp.와 ''Spirulina subsalsa''는 강렬한 햇빛을 피하기 위해 낮에는 하층으로 이동한 다음 황혼에 표면으로 올라온다.^[161] 반대로, 프랑스 카마르그의 과염수 매트에서 발견되는 ''Microcoleus chthonoplastes'' 개체군은 낮 동안 매트의 상층으로 이동하고 밤에는 매트를 통해 균일하게 퍼진다.^[162] ''Phormidium uncinatum''을 사용한 시험관 내 실험에서도 이 종이 손상 방사선을 피하기 위해 이동하는 경향이 있음을 보여주었다.^[159][160] 이러한 이동은 일반적으로 일종의 광운동의 결과이지만 다른 형태의 택시스도 역할을 할 수 있다.^[163][21]

광운동(입사광의 함수로서의 세포 운동의 조절)은 남세균이 환경에서 최적의 광 조건을 찾기 위한 수단으로 사용한다. 광운동에는 광운동, 주광성 및 광 공포 반응의 세 가지 유형이 있다.^{[164][165][166][21]}

광운동성 미생물은 입사광 강도에 따라 활공 속도를 조절한다. 예를 들어, ''Phormidium autumnale''의 활공 속도는 입사광 강도에 따라 선형적으로 증가한다.^[167][21]

주광성 미생물은 환경 내 빛의 방향에 따라 움직이며, 양성 주광성 종은 빛과 평행하게 움직여 광원을 향하는 경향이 있다. ''Phormidium uncinatum''과 같은 종은 빛을 향해 직접 조종할 수 없지만, 올바른 방향으로 스스로를 정렬하기 위해 무작위 충돌에 의존한 후 빛을 향해 더 많이 이동하는 경향이 있다. ''Anabaena variabilis''와 같은 다른 종은 트리콤을 구부려 조종할 수 있다.^[168][21]

마지막으로, 광공포성 미생물은 공간적 및 시간적 빛 기울기에 반응한다. 단계적 광공포 반응은 유기체가 어두운 곳에서 더 밝은 영역으로 들어가 방향을 바꾸어 밝은 빛을 피할 때 발생한다. 반대 반응인 단계적 반응은 유기체가 밝은 곳에서 어두운 영역으로 들어가 방향을 바꾸어 빛 속에 머무를 때 발생한다.^[21]

9. 진화

계통 분석에 따르면, 남세균은 세균 중에서 클로로플렉수스 문, 데이노코쿠스-테르무스 문, 방선균 문, 피르미쿠테스 문 등과 가까운 관계가 있으며, 이들을 테라박테리아로 묶는 것이 제안되었다.^[484][485] 남세균은 최초로 산소 발생형 광합성 능력을 획득한 생물군으로 여겨진다. 메타게놈 연구를 통해 동물의 장, 토양, 오수 등 다양한 환경에서 남세균과 가까운 비광합성 세균들(메라이나박테리아 등)이 발견되었다.^{[486][487][488]} 이 세균들이 광합성 능력의 흔적이 없다는 점에서, 남세균의 조상은 혐기성 비광합성 생물이었으며, 유전자 수평 이동 등으로 산소 발생형 광합성 능력을 얻었다고 생각된다.^[487][489]

남세균 이외의 광합성 세균은 남세균이 가진 광화학계 I 또는 광화학계 II 중 하나만 가지고 있다.^[490] 이러한 시스템에서는 물의 광분해가 일어나지 않아 비산소 발생형 광합성이 수행된다. 두 광화학계가 연결된 산소 발생형 광합성을 하는 세균은 남세균뿐이다.

남세균의 진화는 화석, 바이오마커, 분자 시계 계산 등 다양한 방법으로 연구되고 있지만, 아직 널리 합의된 가설은 없다. 약 24억 년 전 지구 환경이 급격히 변화한 '''대산소화 사건'''은 광합성으로 생성된 산소가 원인으로 추정되며, 남세균은 적어도 이 시기까지는 탄생했을 것으로 보인다.^[494] 그러나 대산소화 사건 이전에도 산소가 존재했다는 증거가 있어, 남세균의 탄생 시기는 논쟁 중이다.^[495][496]

9. 1. 지구 역사

9. 2. 광합성의 기원

9. 3. 엽록체의 기원

9. 4. 해양 기원

10. 유전학

남세균은 자연적으로 유전자 형질전환이 가능하다.^{[220][221][222]} 자연적 유전자 형질전환은 외부 환경으로부터 외래 DNA를 직접 흡수하고 통합하여 세포의 유전적 변형을 일으키는 과정이다. 세균의 형질전환이 일어나려면 수용 세균이 역능 상태여야 하는데, 이는 기아, 높은 세포 밀도, DNA 손상 물질 노출과 같은 조건에 대한 반응으로 자연계에서 발생할 수 있다. 염색체 형질전환에서 상동 형질전환 DNA는 상동 재조합에 의해 수용체 게놈에 통합될 수 있으며, 이 과정은 DNA 손상 복구에 대한 적응으로 보인다.^[223]

10. 1. DNA 복구

11. 인간과의 관계

남세균은 질소 고정 능력을 가지고 있어 유기 비료로 사용되기도 한다. 예를 들어, 개구리밥류는 잎 내부에 남세균을 공생시켜 질소를 고정하며, 이는 논의 녹비로 이용된다.^[473]

남세균이 생성하는 다양한 생리 활성 물질과 세포 외피의 보수성, 자외선 방어 능력은 여러 분야에서 활용 가치가 높아 연구가 진행되고 있다.^{[474][475][476][304][477]}

또한 남세균을 이용한 재생 에너지 연구도 활발하다. 유전자 변형을 통해 광합성으로 에탄올을 생산하거나,^[480] 질소 고정 과정에서 발생하는 수소를 에너지원으로 활용하는 연구가 진행 중이다.^[478] 남세균의 광합성을 직접 전기로 변환하는 연구도 이루어지고 있다.^[481][482]

미국 NASA는 화성 테라포밍에 남세균을 이용하는 방안을 언급하기도 했다.^[483]

11. 1. 생명공학

11. 2. 건강 위험

11. 3. 화학적 제어

11. 4. 기후 변화

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_[226] 웹사이트 LTP_all tree in newick format https://imedea.uib-c[...] 2021-02-23
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_[244] 논문 Genome sequence of the cyanobacterium Prochlorococcus marinus SS120, a nearly minimal oxyphototrophic genome 2003-08
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_[249] 웹사이트 Joule wins key patent for GMO cyanobacteria that create fuels from sunlight, CO2 and water : Biofuels Digest https://www.biofuels[...] 2022-08-10
_[250] 논문 Ethanol synthesis by genetic engineering in cyanobacteria 1999-02
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_[252] 논문 Engineered cyanobacteria with additional overexpression of selected Calvin-Benson-Bassham enzymes show further increased ethanol production 2021-06
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_[260] 논문 Edible blue-green algae reduce the production of pro-inflammatory cytokines by inhibiting NF-κB pathway in macrophages and splenocytes 2013-04
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_[266] 웹사이트 Professor Linda Lawton https://rgu-reposito[...] 2021-06-25
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_[293] 간행물 Determining cell shape: adaptive regulation of cyanobacterial cellular differentiation and morphology
_[294] 간행물 Cyanobacterial cell walls: News from an unusual prokaryotic envelope
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_[296] 간행물 Septal junctions in filamentous heterocyst-forming Cyanobacteria
_[297] 간행물 Role of two cell wall amidases in septal junction and nanopore formation in the multicellular cyanobacterium ''Anabaena'' sp. PCC 7120
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_[301] 간행물 A giant cell surface protein in ''Synechococcus'' WH8102 inhibits feeding by a dinoflagellate predator
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_[304] 간행물 Exocellular polysaccharides from cyanobacteria and their possible applications
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_[362] 논문 Nitrogen fixation and photosynthetic oxygen evolution in cyanobacteria
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_[397] 서적 アオコ : その出現と毒素 東京大学出版会
_[398] 논문 Dynamics of cyanophage-like particles and algicidal bacteria causing ''Microcystis aeruginosa'' mortality
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_[408] 논문 Isolation of salt-induced periplasmic proteins from ''Synechocystis'' sp. strain PCC 6803
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_[411] 서적 The Prokaryotes. A Handbook on the Biology of Bacteria, vol 1, 3rd ed. Symbiotic Associations, Biotechnology, Applied Microbiology Springer
_[412] 논문 Marine cyanobacterial symbioses
_[413] 서적 Algae and Symbioses: Plants, Animals, Fungi, Viruses, Interactions Explored Biopress Limited
_[414] 서적 Marine Protists https://link.springe[...] Springer Japan
_[415] 서적 Cyanobacteria in Symbiosis Kluwer Academic Publishers, Dordrecht
_[416] 논문 ''Geosiphon pyriforme'', a fungus forming endocytobiosis withNostoc (Cyanobacteria), is an ancestral member of the glomales: evidence by SSU rRNA analysis
_[417] 논문 Studies on initiation and development of the partner association in ''Geosiphon pyriforme'' (Kütz.) v. Wettstein, a unique endocytobiotic system of a fungus (Glomales) and the cyanobacterium ''Nostoc punctiforme'' (Kütz.) Hariot
_[418] 서적 In Vitro Culture of Mycorrhizas. Soil Biology, Volume 4, Part V null
_[419] 논문 The ecology and phylogeny of cyanobacterial symbionts in sponges
_[420] 논문 Episymbiotic microbes as food and defence for marine isopods: unique symbioses in a hostile environment
_[421] 논문 Host specificity and phylogeography of the prochlorophyte ''Prochloron'' sp., an obligate symbiont in didemnid ascidians
_[422] 논문 Unicellular cyanobionts in open ocean dinoflagellates, radiolarians, and tintinnids: ultrastructural characterization and immuno-localization of phycoerythrin and nitrogenase
_[423] 논문 Reverse transcription PCR amplification of cyanobacterial symbiont 16S rRNA sequences from single non-photosynthetic eukaryotic marine planktonic host cells
_[424] 논문 Algal symbiosis in larger foraminifera
_[425] 논문 Cyanobacterial endosymbionts in the benthic dinoflagellate Sinophysis canaliculata (Dinophysiales, Dinophyceae)
_[426] 논문 Environmentally-related seasonal variation in symbiotic associations of heterotrophic dinoflagellates with cyanobacteria in the western Bay of Bengal http://drs.nio.org/d[...]
_[427] 논문 The puzzle of plastid evolution
_[428] 논문 もう一つの一次共生？－''Paulinella chromatophora'' とそのシアネレ― https://doi.org/10.1[...] 日本原生生物学会
_[429] 논문 Cyanobacterium-plant symbioses
_[430] 논문 Cyanobacteria-bryophyte symbioses
_[431] 논문 ''Azolla'' and other plant-cyanobacteria symbioses - aspects of form and function
_[432] 논문 Diversity and host specificity of genus ''Azolla'' cyanobionts
_[433] 서적 Cyanobacteria in Symbiosis Kluwer Academic Publishers, Dordrecht
_[434] 서적 Cyanobacteria in Symbiosis Kluwer Academic Publishers
_[435] 논문 The ''Gunnera''-''Nostoc'' symbiosis https://www.jstor.or[...]
_[436] 논문 Biomagnification of cyanobacterial neurotoxins and neurodegenerative disease among the Chamorro people of Guam
_[437] 논문 Intracellular cyanobiont ''Richelia intracellularis'': ultrastructure and immuno-localisation of phycoerythrin, nitrogenase, Rubisco and glutamine synthetase
_[438] 논문 Characterization of diatom-cyanobacteria symbioses on the basis of ''nifH'', ''hetR'' and 16S rRNA sequences
_[439] 논문 Nitrogen fixation and transfer in open ocean diatom-cyanobacterial symbioses
_[440] 논문 Distribution and activity of diazotrophs in the Eastern Equatorial Atlantic
_[441] 논문 Summer surface waters in the Gulf of California: prime habitat for biological nitrogen fixation
_[442] 논문 Discovery of an endosymbiotic nitrogen-fixing cyanobacterium UCYN-A in ''Braarudosphaera bigelowii'' (Prymnesiophyceae)
_[443] 논문 Genetic diversity of the unicellular nitrogen‐fixing cyanobacteria UCYN‐A and its prymnesiophyte host
_[444] 논문 The cyanobacterial endosymbiont of the unicellular algae Rhopalodia gibba shows reductive genome evolution
_[445] 논문 Discovery of symbiotic nitrogen-fixing cyanobacteria in corals
_[446] 논문 Nitrogen fixation by symbiotic cyanobacteria provides a source of nitrogen for the scleractinian coral ''Montastraea cavernosa''
_[447] 논문 Symbiosis between diatoms and cyanobacterial colonies
_[448] 논문 Epiphytic cyanobacteria maintain shifts to macroalgal dominance on coral reefs following ENSO disturbance
_[449] 논문 Molecular detection of epiphytic ''Acaryochloris'' spp. on marine macroalgae
_[450] 논문 Epiphytic cyanobacteria on Chara vulgaris are the main contributors to N2 fixation in rice fields
_[451] 논문 Transfer of fixed-N from N2-fixing cyanobacteria associated with the moss ''Sphagnum riparium'' results in enhanced growth of the moss
_[452] 논문 Cyanobacteria in Symbiosis Kluwer Academic Publishers, Dordrecht
_[453] 논문 The distribution of algae epiphytic on pneumatophores of the mangrove, ''Avicennia marina'', at different salinities in the Kosi System
_[454] 논문 Nitrogen fixation by epiphytic and epibenthic diazotrophs associated with seagrass meadows along the Tanzanian coast, Western Indian Ocean
_[455] 논문 Characterization and com- parison of prokaryotic epiphytes associated with three East African seagrasses
_[456] 논문 Associative cyanobacteria isolated from the roots of epiphytic orchids
_[457] 논문 Cyanobacterial and eukaryotic algal odour compounds: signals or by-products? A review of their biological activity
_[458] 서적 藻類ハンドブック エヌ・ティー・エス
_[459] 서적 藻類ハンドブック エヌ・ティー・エス
_[460] 논문 Cyanobacterial toxins: risk management for health protection https://doi.org/10.1[...]
_[461] 논문 Toxin production of cyanobacteria is increased by exposure to zooplankton
_[462] 논문 Ecotoxicological effects of selected cyanobacterial secondary metabolites a short review https://doi.org/10.1[...]
_[463] 논문 2-II-17 機能性食品スピルリナ錠剤に含まれるビタミンB_<12>同族体の単離と同定 https://doi.org/10.2[...] 日本ビタミン学会
_[464] 웹사이트 食品安全関係情報詳細 資料管理ID:syu04830460475 https://www.fsc.go.j[...]
_[465] 웹사이트 【初心者向き】水槽のコケ対策と種類をプロがアドバイス！ https://tropica.jp/2[...] 株式会社東京アクアガーデン 2021-09-23
_[466] 웹사이트 【コケ種類別】熱帯魚飼育におすすめのコケ取り名人 https://www.gex-fp.c[...] GEX 2021-09-23
_[467] 서적 藻類ハンドブック エヌ・ティー・エス
_[468] 서적 Ecology of Cyanobacteria II Springer Netherlands
_[469] 웹사이트 フィコシアニン（天然系青色素　リナブルー®） https://www.dic-glob[...] DIC 2022-11-05
_[470] 서적 藻類ハンドブック エヌ・ティー・エス
_[471] 서적 藻類ハンドブック エヌ・ティー・エス
_[472] 서적 藻類ハンドブック エヌ・ティー・エス
_[473] 논문 アカウキクサ-ラン藻の共生による生物的窒素固定とその利用 https://doi.org/10.2[...] 日本土壌肥料學會
_[474] 논문 Structure and biosynthesis of borophycin, a new boeseken complex of boric acid from a marine strain of the blue-green alga ''Nostoc linckia''
_[475] 논문 Blue-green algae as an immuno-enhancer and biomodulator https://www.ganzheit[...]
_[476] 논문 Honaucins A−C, potent inhibitors of inflammation and bacterial quorum sensing: synthetic derivatives and structure-activity relationships
_[477] 서적 Ecology of Cyanobacteria II Springer Netherlands
_[478] 논문 多彩な戦略で挑むシアノバクテリア由来の燃料生産:持続可能な第三世代バイオ燃料生産の最前線 https://doi.org/10.1[...] 日本農芸化学会
_[479] 논문 シアノバクテリアの光合成能力を利用したバイオ燃料生産 https://dl.ndl.go.jp[...] 日本生物工学会
_[480] 논문 Algenol hits 9K gallons/acre mark for algae-to-ethanol process] http://www.biofuelsd[...]
_[481] 논문 Light-dependent electrogenic activity of cyanobacteria
_[482] 논문 Renewable energy from Cyanobacteria: energy production optimization by metabolic pathway engineering
_[483] 논문 Sustainable life support on Mars–the potential roles of cyanobacteria
_[484] 논문 A major clade of prokaryotes with ancient adaptations to life on land
_[485] 논문 Insights into the phylogeny and coding potential of microbial dark matter
_[486] 논문 An expanded genomic representation of the phylum Cyanobacteria
_[487] 논문 On the origins of oxygenic photosynthesis and aerobic respiration in Cyanobacteria
_[488] 논문 Hydrogen-based metabolism as an ancestral trait in lineages sibling to the Cyanobacteria
_[489] 논문 Crown group Oxyphotobacteria postdate the rise of oxygen
_[490] 간행물 The Phototrophic Way of Life https://doi.org/10.1[...] Springer 2013
_[491] 논문 Evolution of the 3-hydroxypropionate bicycle and recent transfer of anoxygenic photosynthesis into the Chloroflexi https://www.pnas.org[...] 2017-10-03
_[492] 논문 Phototrophy and carbon fixation in Chlorobi postdate the rise of oxygen https://www.biorxiv.[...] 2021-01-24
_[493] 논문 Evolutionary Implications of Anoxygenic Phototrophy in the Bacterial Phylum Candidatus Eremiobacterota (WPS-2) https://www.frontier[...] 2019
_[494] 논문 The oxygenation of the atmosphere and oceans
_[495] 논문 A Whiff of Oxygen Before the Great Oxidation Event? https://www.science.[...] 2007-09-28
_[496] 논문 Reconciling evidence of oxidative weathering and atmospheric anoxia on Archean Earth https://www.science.[...]
_[497] 논문 Reappraisal of hydrocarbon biomarkers in Archean rocks https://www.pnas.org[...] 2015-05-12
_[498] 논문 Identification of a methylase required for 2-methylhopanoid production and implications for the interpretation of sedimentary hopanes https://www.pnas.org[...] 2010-05-11
_[499] 논문 ストロマトライト研究の歴史と今後の展望 https://doi.org/10.5[...] 東京地学協会
_[500] 논문 Microbially influenced formation of 2,724-million-year-old stromatolites
_[501] 논문 Fossil evidence of Archaean life
_[502] 논문 The Meaning of Stromatolites https://www.annualre[...] 2013-05-30
_[503] 논문 The evolutionary diversification of cyanobacteria: molecular-phylogenetic and paleontological perspectives
_[504] 논문 Multiple sulfur isotopes and the evolution of the atmosphere
_[505] 논문 A molecular timeline for the origin of photosynthetic eukaryotes
_[506] 논문 Plastid evolution
_[507] 논문 An early-branching freshwater cyanobacterium at the origin of plastids
_[508] 논문 Systematische Übersicht über die mit Flagellaten in Zusammenhang stehenden Algenreihen und Versuch einer Einreihung dieser Algenstämme in die Stämme des Pflanzenreiches
_[509] 서적 The Biology of the Algae. 2nd Edition
_[510] 웹사이트 藻類 https://photosyn.jp/[...] 日本光合成学会 null
_[511] 서적 キャンベル生物学 原書11版 丸善出版
_[512] 논문 The relationship between bacteria and Myxophyceae
_[513] 논문 What's in a name? The case of cyanobacteria
_[514] 논문 A proposal for further integration of the cyanobacteria under the Bacteriological Code
_[515] 논문 Prochlorophyta as a proposed new division of algae
_[516] 논문 Modern approach to the classification system of cyanophytes. 1. Introduction
_[517] 논문 The origin of multicellularity in cyanobacteria
_[518] 논문 System of cyanoprokaryotes (cyanobacteria) - state in 2004
_[519] 논문 Cyanobacteria and the Great Oxidation Event: evidence from genes and fossils
_[520] 논문 Improving the coverage of the cyanobacterial phylum using diversity-driven genome sequencing
_[521] 논문 Ecogenomics and taxonomy of Cyanobacteria phylum
_[522] 논문 A comprehensive study of cyanobacterial morphological and ecological evolutionary dynamics through deep geologic time
_[523] 논문 Taxonomic classification of cyanoprokaryotes (cyanobacterial genera) 2014, using a polyphasic approach
_[524] 간행물 AlgaeBase 2019
_[525] 논문 Several problems of the polyphasic approach in the modern cyanobacterial system
_[526] 논문 Comparative genomics of ''Synechococcus'' and proposal of the new genus ''Parasynechococcus''
_[527] 서적 日本淡水藻図鑑 内田老鶴圃
_[528] 서적 바이오디버シティ・シリーズ (3) 藻類の多様性と系統 裳華房
_[529] 서적 바이오디버シティ・シリーズ (3) 藻類の多様性と系統 裳華房
_[530] 논문 Cyanobacterial systematics and nomenclature as featured in the international bulletin of bacteriological nomenclature and taxonomy/international journal of systematic bacteriology/international journal of systematic and evolutionary microbiology
_[531] 웹사이트 CyanoDB 2.0 - On-line database of cyanobacterial genera http://www.cyanodb.c[...] Univ. of South Bohemia & Inst. of Botany AS CR 2019
_[532] 서적 Syllabus of Plant Families. A. Engler's Syllabus der Pflanzenfamilien Part 1/1 Borntraeger
_[533] 웹인용 Cyanophyceae http://www.accesssci[...] Access Science 2011-04-21
_[534] 간행물 A proposal for further integration of the cyanobacteria under the Bacteriological Code
_[535] 뉴스 100년 남극 기후 변화의 지표, 남세균 https://news.naver.c[...] 동아사이언스 2018-04-30
_[536] 뉴스 전기 생산하는 양송이버섯…버섯서 남세균 생존한계 극복 https://news.naver.c[...] 연합뉴스 2018-11-09