积累类胡萝卜素的绿藻类囊体膜蛋白组学研究
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摘要
绿藻在进化地位上介于高等植物和低等光合生物之间，在陆地和海洋环境中都有较广分布。其广泛的分布也使绿藻具有很强的对各种不同环境的适应能力。并且，绿藻生长所经历的多种生境也赋予了其抵抗逆境的生物学特性。尽管不同绿藻适应逆境的机制各不相同，但是很多绿藻在逆境下会合成并且积累类胡萝卜素。而且绿藻中含有的类胡萝卜素种类繁多，并且不同物种在逆境下以合成某一种或某几种类胡萝卜素为主。例如，单细胞淡水绿藻雨生红球藻（Haematococcuspluvialis）在多种逆境下会合成大量酮式类胡萝卜素——虾青素；盐、海水生的单细胞绿藻——杜氏盐藻（Dunaliella salina）在逆境下则合成β-胡萝卜素；而一些大型潮间带海藻如浒苔（Ulva prolifera），虽然也能耐受强光照、干旱等逆境，却并未出现其他两种绿藻具有的大量合成和积累类胡萝卜素的现象。因此，不同种类的类胡萝卜素之于不同的绿藻，在逆境响应中，可能存在不同的协同机制。
光合生物中，类胡萝卜素多与类囊体膜蛋白结合；绿藻作为光合生物，执行光合作用的类囊体膜蛋白极为重要，而且很容易受到逆境损伤。本文从蛋白组学的角度，利用不同的比较手段，研究了三种绿藻——雨生红球藻、盐藻和浒苔之间在蛋白组水平上的差异和变化。重点研究了雨生红球藻不同细胞阶段细胞总可溶蛋白组、类囊体膜蛋白组以及三种绿藻在强光照下类囊体膜蛋白组对强光照的响应。盐藻作为与雨生红球藻非常接近的单细胞绿藻，在逆境下积累虾青素合成的前体物质β-胡萝卜素，但是不会积累虾青素。因此，将这两种微藻放在一起比较，为研究虾青素在雨生红球藻光保护中的作用提供比较和参考的样本。另外，能耐受强光的潮间带的大型海藻浒苔，使用相同的强光照处理方式，研究其光合类囊体膜蛋白组的变化，寻找其与单细胞绿藻所不同的光响应形式。本研究论文关于三种绿藻细胞生理和蛋白组研究的主要内容和关键论点包括：
（1）研究了雨生红球藻不同阶段细胞的固碳能力和虾青素合成之间的关系。四种细胞中，虽然GV（绿色游动）细胞虾青素的成分最低；GV具有最高的CA酶（carbonic anhydrase,碳酸酐酶）活性，并且同位素示踪分析结果表明GV细胞有最高固定无机碳能力。虽然RC（红色不动孢子）含有最高含量的虾青素，但是其含有最低的CA酶活性。因此，虾青素快速合成的OR细胞具有较快地将无机碳转化为虾青素的能力。
（2）雨生红球藻不同细胞阶段的总可溶性蛋白差异荧光双向电泳比较不同生活史阶段细胞蛋白构成差异发现：GV细胞具有细胞快速分裂和生物量积累的蛋白基础；而处于逆境的细胞代谢缓慢。类囊体膜蛋白BN-PAGE和蔗糖密度梯度超速离心-DM增溶SDS聚丙烯酰胺电泳分析差异蛋白显示不同阶段的细胞类囊体膜蛋白保持了主要的光合蛋白。并且鉴定得到一些结合在类囊体膜上的调控蛋白。但是RC细胞可能存在捕光色素蛋白和反应中心蛋白形成的超级蛋白复合体。
（3）通过比较雨生红球藻和盐藻强光响应下类囊体膜蛋白组变化，揭示了雨生红球藻在同等光强条件下比盐藻更快地适应了强光。雨生红球藻可能在适应强光的过程中重新调整了碳流向，通过多种机制参与光保护和淀粉与虾青素的合成。在这种机制中，无糖磷酸途径可能参与了抗逆。通过比较研究浒苔强光响应下类囊体膜蛋白组变化，结果表明浒苔的两个捕光色素蛋白系统表现为不同步的变化；并且浒苔的PsbS蛋白在强光照下表现为上调。可能是通过PsbS来启动了NPQ来响应强光。
本研究通过对三种绿藻的比较分析，积累类胡萝卜素不同的绿藻对逆境响应的机制不同。先在叶绿体内合成β-胡萝卜素并且随后在细胞质内合成虾青素的雨生红球藻响应强光的机制相对特殊与高效。
Green algae evolved at the position between higher plants and lower plants. Andthey are widely distributed both on land and in the sea. The versatile and toughhabitats green algae live in enable them to survive and acclimate. Many green algaespecies synthesize and accumulate carotenoids in the process of stress response.Carotenoids include a wide array of secondary metabolites. For example, the freshwater unicellular microalgae Haematococcus pluvialis are characterized with theability of accumutaing astaxanthin under stress. And Dunaliella salina is able toaccumulate β-carotene intracellularly under stress. However, some macroalgae, Ulvaprolifera for example, show high stress tolerance ability without any carotenoidsaccumulated. Consequently, carotenoids accumulstion and stress acclimation areinterelated, with some synergy effects existed.
Thylakoid membrane is where the main process of photosynthesis occurs, makingit important for green algae. In this research, the three green algae, including H.pluvialis, D. salina and U. prolifera were comparatively studied from the thylakoidproteomic perspective. H. pluvialis, which firstly synthesizes β-carotene and convertsit to astaxanthin under stress, was extensively investigated in this research. However,D. salina synthesizes and accumulates β-carotene under stresses. Hence, D. salinacould be used as a suitable comparative object in H. pluvialis photoprotection research.Besides, the macroalga U. prolifera, which shows neither large amount of β-carotenenor astaxanthin accumulation under high light stress, could also cope with high light.The elucidation of high light responsive pattern in U. prolifera might provide someclue revealing the carotenoids role in high light response. Conseqently, the thylakoidmembranes of three green algae were comparatively studied under high light stress.The main research work and conclusions are as follows:
(1) The relation of carbon fixation and astaxanthin synthesis in H. pluvialisi wasinvestigated. Among the four types of cells, GV (green vegetative) cells showed the highest CA (carbonic anhydrase) activity and the lowest astaxanthin content level.Radioactive labeling and tracing results also confirmed GV showed highest carbonfixation capability. Comparatively, RC (red cyst) cells were highest in astaxanthincontent while lowest in CA activity. Notably, OR (orange resting) cells showedhighest capability of converting fixed carbon into astaxanthin. Those findings were inaccordance with the fact that OR cells were in the process of rapid astaxanthinsynthesis in H. pluvialis.
(2) Comparative analysis using2D DIGE identified some differential expressedproteins among the four types of H. pluvialis cells. The rapid proliferation andbiomass accumulation characteristics GV cells were supported by some up-regulatedproteins. And the other three types of cells, which were under various stress, showedhigher level of proteins involving in low cellular metabolism. Thylakoid membraneproteomics research using BN-PAGE and sucrose gradient ultracentrifugationidentified some regulatory proteins and results also showed that H. pluvialis indifferent stages conserved the photosynthetic proteins. However, RC cells showed thepossibility of LHC-PS super-complex forming as revealed by continuous sucrosegradient ultracentrifugation.
(3) By quantitatively monitoring the abundance variations of thylakoid membraneproteins, H. pluvialis showed great high light acclimation capability. Based on thefindings in this study and inspired by results from others, a hyperthetical carbon flowreorientation strategy was proposed, by which the H. pluvialis employed during highlight acclimation. It was theoretically possible that phosphate pentose pathway wasinvolved in stress induced metabolisms including starch and astaxanthin synthesis.Comparatively, U. prolifera showed inbalanced energy distribution pattern betweenthe two photosystems as indicated by the abundance alteration of LHCI and LHCII.Moreover, PsbS protein was monitored to be significantly up-regulated under highlight. Thus it is possible that PsbS activated and enhanced NPQ to funnel excessenergy for photoprotection.
光合生物中，类胡萝卜素多与类囊体膜蛋白结合；绿藻作为光合生物，执行光合作用的类囊体膜蛋白极为重要，而且很容易受到逆境损伤。本文从蛋白组学的角度，利用不同的比较手段，研究了三种绿藻——雨生红球藻、盐藻和浒苔之间在蛋白组水平上的差异和变化。重点研究了雨生红球藻不同细胞阶段细胞总可溶蛋白组、类囊体膜蛋白组以及三种绿藻在强光照下类囊体膜蛋白组对强光照的响应。盐藻作为与雨生红球藻非常接近的单细胞绿藻，在逆境下积累虾青素合成的前体物质β-胡萝卜素，但是不会积累虾青素。因此，将这两种微藻放在一起比较，为研究虾青素在雨生红球藻光保护中的作用提供比较和参考的样本。另外，能耐受强光的潮间带的大型海藻浒苔，使用相同的强光照处理方式，研究其光合类囊体膜蛋白组的变化，寻找其与单细胞绿藻所不同的光响应形式。本研究论文关于三种绿藻细胞生理和蛋白组研究的主要内容和关键论点包括：
（1）研究了雨生红球藻不同阶段细胞的固碳能力和虾青素合成之间的关系。四种细胞中，虽然GV（绿色游动）细胞虾青素的成分最低；GV具有最高的CA酶（carbonic anhydrase,碳酸酐酶）活性，并且同位素示踪分析结果表明GV细胞有最高固定无机碳能力。虽然RC（红色不动孢子）含有最高含量的虾青素，但是其含有最低的CA酶活性。因此，虾青素快速合成的OR细胞具有较快地将无机碳转化为虾青素的能力。
（2）雨生红球藻不同细胞阶段的总可溶性蛋白差异荧光双向电泳比较不同生活史阶段细胞蛋白构成差异发现：GV细胞具有细胞快速分裂和生物量积累的蛋白基础；而处于逆境的细胞代谢缓慢。类囊体膜蛋白BN-PAGE和蔗糖密度梯度超速离心-DM增溶SDS聚丙烯酰胺电泳分析差异蛋白显示不同阶段的细胞类囊体膜蛋白保持了主要的光合蛋白。并且鉴定得到一些结合在类囊体膜上的调控蛋白。但是RC细胞可能存在捕光色素蛋白和反应中心蛋白形成的超级蛋白复合体。
（3）通过比较雨生红球藻和盐藻强光响应下类囊体膜蛋白组变化，揭示了雨生红球藻在同等光强条件下比盐藻更快地适应了强光。雨生红球藻可能在适应强光的过程中重新调整了碳流向，通过多种机制参与光保护和淀粉与虾青素的合成。在这种机制中，无糖磷酸途径可能参与了抗逆。通过比较研究浒苔强光响应下类囊体膜蛋白组变化，结果表明浒苔的两个捕光色素蛋白系统表现为不同步的变化；并且浒苔的PsbS蛋白在强光照下表现为上调。可能是通过PsbS来启动了NPQ来响应强光。
本研究通过对三种绿藻的比较分析，积累类胡萝卜素不同的绿藻对逆境响应的机制不同。先在叶绿体内合成β-胡萝卜素并且随后在细胞质内合成虾青素的雨生红球藻响应强光的机制相对特殊与高效。
Green algae evolved at the position between higher plants and lower plants. Andthey are widely distributed both on land and in the sea. The versatile and toughhabitats green algae live in enable them to survive and acclimate. Many green algaespecies synthesize and accumulate carotenoids in the process of stress response.Carotenoids include a wide array of secondary metabolites. For example, the freshwater unicellular microalgae Haematococcus pluvialis are characterized with theability of accumutaing astaxanthin under stress. And Dunaliella salina is able toaccumulate β-carotene intracellularly under stress. However, some macroalgae, Ulvaprolifera for example, show high stress tolerance ability without any carotenoidsaccumulated. Consequently, carotenoids accumulstion and stress acclimation areinterelated, with some synergy effects existed.
Thylakoid membrane is where the main process of photosynthesis occurs, makingit important for green algae. In this research, the three green algae, including H.pluvialis, D. salina and U. prolifera were comparatively studied from the thylakoidproteomic perspective. H. pluvialis, which firstly synthesizes β-carotene and convertsit to astaxanthin under stress, was extensively investigated in this research. However,D. salina synthesizes and accumulates β-carotene under stresses. Hence, D. salinacould be used as a suitable comparative object in H. pluvialis photoprotection research.Besides, the macroalga U. prolifera, which shows neither large amount of β-carotenenor astaxanthin accumulation under high light stress, could also cope with high light.The elucidation of high light responsive pattern in U. prolifera might provide someclue revealing the carotenoids role in high light response. Conseqently, the thylakoidmembranes of three green algae were comparatively studied under high light stress.The main research work and conclusions are as follows:
(1) The relation of carbon fixation and astaxanthin synthesis in H. pluvialisi wasinvestigated. Among the four types of cells, GV (green vegetative) cells showed the highest CA (carbonic anhydrase) activity and the lowest astaxanthin content level.Radioactive labeling and tracing results also confirmed GV showed highest carbonfixation capability. Comparatively, RC (red cyst) cells were highest in astaxanthincontent while lowest in CA activity. Notably, OR (orange resting) cells showedhighest capability of converting fixed carbon into astaxanthin. Those findings were inaccordance with the fact that OR cells were in the process of rapid astaxanthinsynthesis in H. pluvialis.
(2) Comparative analysis using2D DIGE identified some differential expressedproteins among the four types of H. pluvialis cells. The rapid proliferation andbiomass accumulation characteristics GV cells were supported by some up-regulatedproteins. And the other three types of cells, which were under various stress, showedhigher level of proteins involving in low cellular metabolism. Thylakoid membraneproteomics research using BN-PAGE and sucrose gradient ultracentrifugationidentified some regulatory proteins and results also showed that H. pluvialis indifferent stages conserved the photosynthetic proteins. However, RC cells showed thepossibility of LHC-PS super-complex forming as revealed by continuous sucrosegradient ultracentrifugation.
(3) By quantitatively monitoring the abundance variations of thylakoid membraneproteins, H. pluvialis showed great high light acclimation capability. Based on thefindings in this study and inspired by results from others, a hyperthetical carbon flowreorientation strategy was proposed, by which the H. pluvialis employed during highlight acclimation. It was theoretically possible that phosphate pentose pathway wasinvolved in stress induced metabolisms including starch and astaxanthin synthesis.Comparatively, U. prolifera showed inbalanced energy distribution pattern betweenthe two photosystems as indicated by the abundance alteration of LHCI and LHCII.Moreover, PsbS protein was monitored to be significantly up-regulated under highlight. Thus it is possible that PsbS activated and enhanced NPQ to funnel excessenergy for photoprotection.
引文
丁兰平,栾日孝(2009)浒苔（Enteromorpha prolifera）的分类鉴定、生境习性及分布.海洋与湖沼40:68-71.
韦韬,顾文辉,李健,张波,潘光华,朱大玲,王广策(2012)不同碳氮浓度对雨生红球藻生长及虾青素累积的影响.海洋科学36:55-61.
韦韬,顾文辉,李健,张波,潘光华,朱大玲,王广策(2013)雨生红球藻的光周期效应.植物学报48:168-173.
Aflalo C, Meshulam Y, Zarka A, Boussiba S (2007) On the relative efficiency of two-vs. one-stage production of astaxanthin by the green alga Haematococcuspluvialis. Biotechnology and bioengineering98:300-305.
Alberte RS, Friedman AL, Gustafson DL, Rudnick MS, Lyman H (1981)Light-harvesting systems of brown algae and diatoms. Isolation andcharacterization of chlorophyll ac and chlorophyll afucoxanthinpigment-protein complexes. Biochimica et Biophysica Acta(BBA)-Bioenergetics635:304-316.
Alric J (2010) Cyclic electron flow around photosystem I in unicellular green algae.Photosynthesis Research106:47-56.
Apel K, Hirt H (2004) Reactive oxygen species: Metabolism, Oxidative Stress, andSignal Transduction. Annual Review of Plant Biology55:373-399.
Asara JM, Christofk HR, Freimark LM, Cantley LC (2008) A label‐freequantification method by MS/MS TIC compared to SILAC and spectralcounting in a proteomics screen. Proteomics8:994-999.
Baas-Becking L (1931) Salt effects on swarmers of Dunaliella viridis Teod. TheJournal of general physiology14:765.
Bachi A, Bonaldi T (2008) Quantitative proteomics as a new piece of the systemsbiology puzzle. Journal of Proteomics71:357-367.
Banse K (2002) Steemann Nielsen and the zooplankton. Hydrobiologia480:15-28.
Bar E, Rise M, Vishkautsan M, Arad S (1995) Pigment and Structural Changes inChlorella zofingiensis upon Light and Nitrogen Stress. Journal of PlantPhysiology146:527-534.
Bassi R, Caffarri S (2000) Lhc proteins and the regulation of photosynthetic lightharvesting function by xanthophylls. Photosynthesis Research64:243-256.
Ben-Amotz A (1995) New mode of Dunaliella biotechnology: two-phase growth forβ-carotene production. Journal ofApplied Phycology7:65-68.
Bj rn L, Papageorgiou G, Blankenship R, Govindjee (2009) A viewpoint: Whychlorophyll a? Photosynthesis Research99:85-98.
Blum H, Beier H, Gross HJ (1987) Improved silver staining of plant proteins, RNAand DNA in polyacrylamide gels. Electrophoresis8:93-99.
Borowitzka MA, Huisman JM, Osborn A (1991) Culture of the astaxanthin-producinggreen alga Haematococcus pluvialis1. Effects of nutrients on growth and celltype. Journal of Applied Phycology3:295-304.
Boussiba S (2000) Carotenogenesis in the green alga Haematococcus pluvialis:Cellular physiology and stress response. Physiologia Plantarum108:111-117.
Boussiba S, Bing W, Yuan J-P, Zarka A, Chen F (1999) Changes in pigments profile inthe green alga Haeamtococcus pluvialis exposed to environmental stresses.Biotechnology Letters21:601-604.
Boussiba S, Fan L, Vonshak A, Lester P (1992) Enhancement and determination ofastaxanthin accumulation in green alga Haematococcus pluvialis. San Diego,CA, ETATS-UNIS: Academic Press.
Bradford MM (1976) A rapid and sensitive method for the quantitation of microgramquantities of protein utilizing the principle of protein-dye binding. AnalyticalBiochemistry72:248-254.
Buchanan BB, Balmer Y (2005) Redox regulation: A broadening horizon. AnnualReview of Plant Biology56:187-220.
Buchanan BB, Gruissem W, Jones RL (2000) Biochemistry&molecular biology ofplants, Chapter12Photosynthesis: American Society of Plant PhysiologistsRockville.
Caffarri S, Kouril R, Kereiche S, Boekema EJ, Croce R (2009) Functionalarchitecture of higher plant photosystem II supercomplexes. EMBO J28:3052-3063.
Chang HI (2010) Protective effect of the natural pigment on naproxen induced gastriculceration in rat. Journal of Biotechnology150:68-68.
Chen Z, Wang G, Niu J (2012) Variation in Rubisco and other photosyntheticparameters in the life cycle of Haematococcus pluvialis. Chinese Journal ofOceanology and Limnology30:136-145.
Chua NH, Bennoun P (1975) Thylakoid membrane polypeptides of Chlamydomonasreinhardtii: wild-type and mutant strains deficient in photosystem II reactioncenter. Proceedings of the National Academy of Sciences72:2175-2179.
Collins AM, Jones HDT, Han D, Hu Q, Beechem TE, Timlin JA (2011) CarotenoidDistribution in Living Cells of Haematococcus pluvialis (Chlorophyceae).PLoS ONE6: e24302.
Cort A, Ozturk N, Akpinar D, Unal M, Yucel G, Ciftcioglu A, Yargicoglu P, Aslan M(2010) Suppressive effect of astaxanthin on retinal injury induced by elevatedintraocular pressure. Regulatory Toxicology and Pharmacology58:121-130.
Cunningham FX, Gantt E (1998) GENES AND ENZYMES OF CAROTENOIDBIOSYNTHESIS IN PLANTS. Annual Review of Plant Physiology and PlantMolecular Biology49:557-583.
Cunningham FX, Gantt E (2011) Elucidation of the Pathway to Astaxanthin in theFlowers of Adonis aestivalis. The Plant Cell Online23:3055-3069.
Czeczuga B (1974) Carotenoids in Euglena rubida Mainx. Comparative Biochemistryand Physiology Part B: Comparative Biochemistry48:349-354.
D'Amici GM, Huber CG, Zolla L (2009) Separation of thylakoid membrane proteinsby sucrose gradient ultracentrifugation or blue native-SDS-PAGEtwo-dimensional electrophoresis. Methods Mol Biol528:61-70.
Damiani MC, Popovich CA, Constenla D, Leonardi PI (2010) Lipid analysis inHaematococcus pluvialis to assess its potential use as a biodiesel feedstock.Bioresource Technology101:3801-3807.
Davison PA, Hunter CN, Horton P (2002) Overexpression of [beta]-carotenehydroxylase enhances stress tolerance in Arabidopsis. Nature418:203-206.
Del Campo JA, Rodríguez H, Moreno J, Vargas Má, Rivas J, Guerrero MG (2004)Accumulation of astaxanthin and lutein in Chlorella zofingiensis(Chlorophyta). Applied Microbiology and Biotechnology64:848-854.
Deo PM, Biswal B (2001) Response of senescing cotyledons of clusterbean to waterstress in moderate and low light: Possible photoprotective role of β‐carotene.Physiologia plantarum112:47-54.
Ding L, Fei X, Lu Q, Deng Y, Lian S (2009) The possibility analysis of habitats,origin and reappearance of bloom green alga (Enteromorpha prolifera) oninshore of western Yellow Sea. Chinese Journal of Oceanology and Limnology27:421-424.
Droop MR (1954) Conditions governing haematochrome formation and loss in thealga Haematococcus pluvialis flotow. Archiv für Mikrobiologie20:391-397.
Droop MR (1955) Carotenogenesis in Haematococcus pluvialis. Nature175:42-42.
Ellis RJ (1979) Most abundant protein in the world. Trends in Biochemical Sciences4:241-244.
Ellis RJ (1979) Msot abundant protein in the world. Trends in Biochemical Sciences4:241-244.
Ephritikhine G, Ferro M, Rolland N (2004) Plant membrane proteomics. PlantPhysiology and Biochemistry42:943-962.
Escoubas J-M, Lomas M, LaRoche J, Falkowski PG (1995) Light intensity regulationof cab gene transcription is signaled by the redox state of the plastoquinonepool. Proceedings of the National Academy of Sciences92:10237-10241.
Eshaghi S, Andersson B, Barber J (1999) Isolation of a highly active PSII-LHCIIsupercomplex from thylakoid membranes by a direct method. FEBS Letters446:23-26.
Fábregas J, Otero A, Maseda A, Domínguez A (2001) Two-stage cultures for theproduction of Astaxanthin from Haematococcus pluvialis. Journal ofBiotechnology89:65-71.
Fabregas J, Dominguez A, Regueiro M, Maseda A, Otero A (2000) Optimization ofculture medium for the continuous cultivation of the microalgaHaematococcus pluvialis. Applied Microbiology and Biotechnology53:530-535.
Fan L, Vonshak A, Boussiba S (1994) Effect of temperature and irradiance on growthof Haematococcus pluvialis (chlorophyceae)1. Journal of Phycology30:829-833.
Field CB, Behrenfeld MJ, Randerson JT, Falkowski P (1998) Primary Production ofthe Biosphere: Integrating Terrestrial and Oceanic Components. Science281:237-240.
Flotow J v (1844) Haematococcus pluvialis. Verh Kais Leopold-Carol Akad NaturfNova Acta Acad Caes Leopold Carolin Nat Curios:411-606.
Frey PA, Hegeman AD, Ruzicka FJ (2008) The Radical SAM Superfamily. CriticalReviews in Biochemistry and Molecular Biology43:63-88.
Fromm HJ, Hargrove MS (2012) Photosynthesis
Essentials of Biochemistry. Springer Berlin Heidelberg. pp.317-329.
Funk C, Schr der WP, Green BR, Renger G, Andersson B (1994) The intrinsic22kDaprotein is a chlorophyll-binding subunit of photosystem II. FEBS Letters342:261-266.
Gao S, Chen X, Yi Q, Wang G, Pan G, Lin A, Peng G (2010) A Strategy for theProliferation of Ulva prolifera, Main Causative Species of Green Tides, withFormation of Sporangia by Fragmentation. PLoS ONE5: e8571.
Gao S, Shen S, Wang G, Niu J, Lin A, Pan G (2011) PSI-driven cyclic electron flowallows intertidal macro-algae Ulva sp.(Chlorophyta) to survive in desiccatedconditions. Plant Cell Physiol52:885-893.
Goodwin TW (1955) Carotenoids. Annual Review of Biochemistry24:497-522.
Green B, Durnford D (1996) The chlorophyll-carotenoid proteins of oxygenicphotosynthesis. Annual review of plant biology47:685-714.
Gu W, Xie X, Gao S, Zhou W, Pan G, Wang G (2013) Comparison of different cells ofHaematococcus pluvialis reveals an extensive acclimation mechanism duringits aging process: From a perspective of photosynthesis. PLoS One8.
Guerin M, Huntley ME, Olaizola M (2003) Haematococcus astaxanthin: applicationsfor human health and nutrition. Trends in Biotechnology21:210-216.
Hagen C, Grünewald K, Schmidt S, Müller J (2000) Accumulation of secondarycarotenoids in flagellates of Haematococcus pluvialis (Chlorophyta) isaccompanied by an increase in per unit chlorophyll productivity ofphotosynthesis. European Journal of Phycology35:75-82.
Hagen C, Grünewald K, Xyl nder M, Rothe E (2001) Effect of cultivation parameterson growth and pigment biosynthesis in flagellated cells of Haematococcuspluvialis. Journal of Applied Phycology13:79-87.
Haglund K, Bj rk M, Ramazanov Z, García-Reina G, Pedersén M (1992) Role ofcarbonic anhydrase in photosynthesis and inorganic-carbon assimilation in thered alga Gracilaria tenuistipitata. Planta187:275-281.
Han D, Li Y, Hu Q (2013) Astaxanthin in microalgae: pathways, functions andbiotechnological implications. Algae28:131-147.
Han DX, Wang JF, Sommerfeld M, Hu Q (2012) Susceptibility and protectivemechanisms of motile and non motile cells of Haematococcus Pluvialis(Chlorophyceae) to photooxidative stress. Journal of Phycology48:693-705.
Hankamer B, Morris E, Nield J, Gerle C, Barber J (2001) Three-dimensional structureof the photosystem II core dimer of higher plants determined by electronmicroscopy. Journal of structural biology135:262-269.
Harker M, Hirschberg J (1997) Biosynthesis of ketocarotenoids in transgeniccyanobacteria expressing the algal gene for [beta]-C-4-oxygenase, crtO. FEBSLetters404:129-134.
Harker M, Tsavalos AJ, Young AJ (1996) Factors responsible for astaxanthinformation in the Chlorophyte Haematococcus pluvialis. BioresourceTechnology55:207-214.
Hejazi M, Holwerda E, Wijffels R (2004) Milking microalga Dunaliella salina for β‐carotene production in two‐phase bioreactors. Biotechnology andbioengineering85:475-481.
Helbig AO, Heck AJR, Slijper M (2010) Exploring the membraneproteome--Challenges and analytical strategies. Journal of Proteomics73:868-878.
Higuera-Ciapara I, Félix-Valenzuela L, Goycoolea FM (2006) Astaxanthin: A Reviewof its Chemistry and Applications. Critical Reviews in Food Science andNutrition46:185-196.
Holmgren A, Johansson C, Berndt C, Lonn M, Hudemann C, Lillig C (2005) Thiolredox control via thioredoxin and glutaredoxin systems. Biochemical SocietyTransactions33:1375-1377.
Holtin K, Kuehnle M, Rehbein J, Schuler P, Nicholson G, Albert K (2009)Determination of astaxanthin and astaxanthin esters in the microalgaeHaematococcus pluvialis by LC-(APCI)MS and characterization ofpredominant carotenoid isomers by NMR spectroscopy. Analytical andBioanalytical Chemistry395:1613-1622.
Horton P, Ruban AV, Walters RG (1996) Regulation of light harvesting in green plants.Annual Review of Plant Physiology and Plant Molecular Biology47:655-684.
Ivanov AG, Rosso D, Savitch LV, Stachula P, Rosembert M, Oquist G, Hurry V, HünerNPA (2012) Implications of alternative electron sinks in increased resistanceof PSII and PSI photochemistry to high light stress in cold-acclimatedArabidopsis thaliana. Photosynthesis Research113:191-206.
Jahns P, Holzwarth AR (2012) The role of the xanthophyll cycle and of lutein inphotoprotection of photosystem II. Biochim Biophys Acta1817:182-193.
Jiang C-D, Gao H-Y, Zou Q, Jiang G-M, Li L-H (2006) Leaf orientation,photorespiration and xanthophyll cycle protect young soybean leaves againsthigh irradiance in field. Environmental and Experimental Botany55:87-96.
JIN E, LEE C (2006) Secondary Carotenoid Accumulation in Haematococcus(Chlorophyceae): Biosynthesis, Regulation, and Biotechnology. J MicrobiolBiotechnol16:821-831.
Johnson EA, Lewis MJ (1979) Astaxanthin formation by the yeast Phaffia rhodozyma.J Gen Microbiol115:173-183.
Kügler M, J nsch L, Kruft V, Schmitz UK, Braun H-P (1997) Analysis of thechloroplast protein complexes by blue-native polyacrylamide gelelectrophoresis (BN-PAGE). Photosynthesis Research53:35-44.
Kajiwara S, Kakizono T, Saito T, Kondo K, Ohtani T, Nishio N, Nagai S, Misawa N(1995) Isolation and functional identification of a novel cDNA for astaxanthinbiosynthesis from Haematococcus pluvialis, and astaxanthin synthesis inEscherichia coli. Plant Molecular Biology29:343-352.
Kamath BS, Srikanta BM, Dharmesh SM, Sarada R, Ravishankar GA (2008) Ulcerpreventive and antioxidative properties of astaxanthin from Haematococcuspluvialis. European Journal of Pharmacology590:387-395.
Kang C, Lee J, Park T, Sim S (2007) Complementary limiting factors of astaxanthinsynthesis during photoautotrophic induction of Haematococcus pluvialis: C/Nratio and light intensity. Applied Microbiology and Biotechnology74:987-994.
Kargul J, Barber J (2008) Photosynthetic acclimation: Structural reorganisation oflight harvesting antenna–role of redox-dependent phosphorylation of majorand minor chlorophyll a/b binding proteins. FEBS Journal275:1056-1068.
Kasahara M, Kagawa T, Oikawa K, Suetsugu N, Miyao M, Wada M (2002)Chloroplast avoidance movement reduces photodamage in plants. Nature420:829-832.
Katz A, Waridel P, Shevchenko A, Pick U (2007) Salt-induced changes in the plasmamembrane proteome of the halotolerant alga Dunaliella salina as revealed byblue native gel electrophoresis and nano-LC-MS/MS analysis. Molecular&Cellular Proteomics6:1459-1472.
Kim S-J, Chang HI (2010) Astaxanthin represses adipogenesis in3T3-L1cells.Journal of Biotechnology150:308-308.
Kim S, Sandusky P, Bowlby NR, Aebersold R, Green BR, Vlahaskis S, Yocum CF,Pichersky E (1992) Characterization of a spinach psbS cDNA encoding the22kDa protein of photosystem II. FEBS Letters314:67-71.
Kindlund P (2011) Astaxanthin. Nutrafoods10:27-31.
Kobayashi M (2003) Astaxanthin biosynthesis enhanced by reactive oxygen species inthe green alga Haematococcus pluvialis. Biotechnology and BioprocessEngineering8:322-330.
Kobayashi M, Kakizono T, Nagai S (1993) Enhanced carotenoid biosynthesis byoxidative stress in acetate-induced cyst cells of a green unicellular alga,Haematococcus pluvialis. Applied And Environmental Microbiology59:867.
Kobayashi M, Kakizono T, Yamaguchi K, Nishio N, Nagai S (1992) Growth andastaxanthin formation of Haematococcus pluvialis in heterotrophic andmixotrophic conditions. Journal of Fermentation and Bioengineering74:17-20.
Kobayashi M, Katsuragi T, Tani Y (2001) Enlarged and astaxanthin-accumulating cystcells of the green alga Haematococcus pluvialis. Journal of Bioscience andBioengineering92:565-568.
Kobayashi M, Kurimura Y, Kakizono T, Nishio N, Tsuji Y (1997) Morphologicalchanges in the life cycle of the green alga Haematococcus pluvialis. Journal ofFermentation and Bioengineering84:94-97.
Kota U, Goshe MB (2011) Advances in qualitative and quantitative plant membraneproteomics. Phytochemistry72:1040-1060.
Kristoffersen AS, Svensen, Ssebiyonga N, Erga SR, Stamnes JJ, Frette (2012)Chlorophyll a and NADPH Fluorescence Lifetimes in the MicroalgaeHaematococcus pluvialis (Chlorophyceae) under Normal andAstaxanthin-Accumulating Conditions. Appl Spectrosc66:1216-1225.
Lamers PP, van de Laak CCW, Kaasenbrood PS, Lorier J, Janssen M, De Vos RCH,Bino RJ, Wijffels RH (2010) Carotenoid and fatty acid metabolism inlight-stressed Dunaliella salina. Biotechnology and Bioengineering106:638-648.
Layer G, Heinz DW, Jahn D, Schubert W-D (2004) Structure and function of radicalSAM enzymes. Current Opinion in Chemical Biology8:468-476.
Leister D (2003) Chloroplast research in the genomic age. Trends in Genetics19:47-56.
Lello Z, Anna MT, Christian GH (2004) Thylakoid Membrane Proteome: Separationby HPLC and Identification by Accurate Molecular Mass Determinations.Current Proteomics1:247-260.
Lemoine Y, Schoefs B (2010) Secondary ketocarotenoid astaxanthin biosynthesis inalgae: a multifunctional response to stress. Photosynthesis Research106:155-177.
Li J, Zhu D, Niu J, Shen S, Wang G (2011) An economic assessment of astaxanthinproduction by large scale cultivation of Haematococcus pluvialis.Biotechnology Advances29:568-574.
Li X-P, Bj rkman O, Shih C, Grossman AR, Rosenquist M, Jansson S, Niyogi KK(2000) A pigment-binding protein essential for regulation of photosyntheticlight harvesting. Nature403:391-395.
Li X-P, Gilmore AM, Caffarri S, Bassi R, Golan T, Kramer D, Niyogi KK (2004)Regulation of Photosynthetic Light Harvesting Involves Intrathylakoid LumenpH Sensing by the PsbS Protein. Journal of Biological Chemistry279:22866-22874.
Li X-P, Müller-Moulé P, Gilmore AM, Niyogi KK (2002) PsbS-dependentenhancement of feedback de-excitation protects photosystem II fromphotoinhibition. Proceedings of the National Academy of Sciences99:15222-15227.
Li Y, Sommerfeld M, Chen F, Hu Q (2008) Consumption of oxygen by astaxanthinbiosynthesis: a protective mechanism against oxidative stress inHaematococcus pluvialis (Chlorophyceae). J Plant Physiol165:1783-1797.
Li Y, Sommerfeld M, Chen F, Hu Q (2010) Effect of photon flux densities onregulation of carotenogenesis and cell viability of Haematococcus pluvialis(Chlorophyceae). Journal of Applied Phycology22:253-263.
Li Z, Ahn TK, Avenson TJ, Ballottari M, Cruz JA, Kramer DM, Bassi R, Fleming GR,Keasling JD, Niyogi KK (2009) Lutein Accumulation in the Absence ofZeaxanthin Restores Nonphotochemical Quenching in the Arabidopsisthaliana npq1Mutant. The Plant Cell Online21:1798-1812.
Li Z, Wakao S, Fischer BB, Niyogi KK (2009) Sensing and responding to excess light.Annu Rev Plant Biol60:239-260.
Lichtenthaler HK, Babani F (2004) Light Adaptation and Senescence of thePhotosynthetic Apparatus. Changes in Pigment Composition, ChlorophyllFluorescence Parameters and Photosynthetic Activity. In: Papageorgiou GC,Govindjee, editors. Chlorophyll a Fluorescence: Springer Netherlands. pp.713-736.
Lin A, Shen S, Wang J, Yan B (2008) Reproduction diversity of Enteromorphaprolifera. Journal of integrative plant biology50:622-629.
Lin R (2002) Xanthophyll Cycle and Its Molecular Mechanism in Photoprotection.Acta Botanica Sinica44.
Liska AJ, Shevchenko A, Pick U, Katz A (2004) Enhanced photosynthesis and redoxenergy production contribute to salinity tolerance in Dunaliella as revealed byhomology-based proteomics. Plant physiology136:2806-2817.
Long SP (1994) Photoinhibition of photosynthesis in nature. Annu Rev Plant PhysiolPlant Mol Biol45:633-662.
Mackinney G (1952) Carotenoids. Annual Review of Biochemistry21:473-492.
Majeran W, Zybailov B, Ytterberg AJ, Dunsmore J, Sun Q, van Wijk KJ (2008)Consequences of C4Differentiation for Chloroplast Membrane Proteomes inMaize Mesophyll and Bundle Sheath Cells. Molecular&Cellular Proteomics7:1609-1638.
Marouga R, David S, Hawkins E (2005) The development of the DIGE system:2Dfluorescence difference gel analysis technology. Analytical and BioanalyticalChemistry382:669-678.
Maxwell DP, Laudenbach DE, Huner N (1995) Redox Regulation of Light-HarvestingComplex II and cab mRNA Abundance in Dunaliella salina. Plant Physiology109:787-795.
McDonald AE, Ivanov AG, Bode R, Maxwell DP, Rodermel SR, Hüner NPA (2011)Flexibility in photosynthetic electron transport: The physiological role ofplastoquinol terminal oxidase (PTOX). Biochimica et Biophysica Acta (BBA)-Bioenergetics1807:954-967.
McLean K, Sabri M, Marshall K, Lawson R, Lewis D, Clift D, Balding P, Dunford A,Warman A, McVey J (2005) Biodiversity of cytochrome P450redox systems.Biochemical Society Transactions33:796-801.
Melkozernov AN, Barber J, Blankenship RE (2005) Light Harvesting in PhotosystemI Supercomplexes,. Biochemistry45:331-345.
Miflin BJ, Lea PJ (1976) The pathway of nitrogen assimilation in plants.Phytochemistry15:873-885.
Miki W (1991) Biological function and activity of animal carotenoids. Pure ApplChem:141-146.
Milledge J (2010) Commercial application of microalgae other than as biofuels: abrief review. Reviews in Environmental Science and Biotechnology:1-11.
Monteoliva L, Albar JP (2004) Differential proteomics: an overview of gel andnon-gel based approaches. Briefings in functional genomics&proteomics3:220-239.
Moore GF, Brudvig GW (2011) Energy Conversion in Photosynthesis: A Paradigm forSolar Fuel Production. Annual Review of Condensed Matter Physics2:303-327.
Mou S, Zhang X, Dong M, Fan X, Xu J, Cao S, Xu D, Wang W, Ye N (2013)Photoprotection in the green tidal alga Ulva prolifera: role of LHCSR andPsbS proteins in response to high light stress. Plant Biol (Stuttg).
Murakami A (1997) Quantitative analysis of77K fluorescence emission spectra inSynechocystis sp. PCC6714and Chlamydomonas reinhardtii with variable PSI/PS II stoichiometries. Photosynthesis Research53:141-148.
Murata N, Nishimura M, Takamiya A (1966) Fluorescence of chlorophyll inphotosynthetic systems III. Emission and action spectra offluorescence—Three emission bands of chlorophyll a and the energy transferbetween two pigment systems. Biochimica et Biophysica Acta (BBA)-Biophysics including Photosynthesis126:234-243.
Naumann B, Hippler M (2007) Insights into chloroplast proteomics: from basicprinciples to new horizonsCell and Molecular Biology of Plastids. In: Bock R, editor. Cell and MolecularBiology of Plastids: Springer Berlin/Heidelberg. pp.371-407.
Neidhardt J, Benemann J, Zhang L, Melis A (1998) Photosystem-II repair andchloroplast recovery from irradiance stress: relationship between chronicphotoinhibition, light-harvesting chlorophyll antenna size and photosyntheticproductivity in Dunaliella salina (green algae). Photosynthesis Research56:175-184.
Neilson KA, Ali NA, Muralidharan S, Mirzaei M, Mariani M, Assadourian G, Lee A,van Sluyter SC, Haynes PA (2011) Less label, more free: Approaches inlabel-free quantitative mass spectrometry. PROTEOMICS11:535-553.
Nielsen ES (1952) The Use of Radio-active Carbon (C14) for Measuring OrganicProduction in the Sea. Journal du Conseil18:117-140.
Niu J, Hu H, Hu S, Wang G, Peng G, Sun S (2010) Analysis of expressed sequencetags from the Ulva prolifera (Chlorophyta). Chinese Journal of Oceanologyand Limnology28:26-36.
Niyogi KK (1999) Photoprotection revisited: genetic and molecular approaches.Annual Review of Plant Physiology and Plant Molecular Biology50:333-359.
Oeljeklaus S, Meyer HE, Warscheid B (2009) Advancements in plant proteomicsusing quantitative mass spectrometry. Journal of Proteomics72:545-554.
Old WM, Meyer-Arendt K, Aveline-Wolf L, Pierce KG, Mendoza A, Sevinsky JR,Resing KA, Ahn NG (2005) Comparison of Label-free Methods forQuantifying Human Proteins by Shotgun Proteomics. Molecular&CellularProteomics4:1487-1502.
Oren A (2005) A hundred years of Dunaliella research:1905-2005. Saline Systems1:2.
Oren A (2005) A hundred years of Dunaliella research:1905–2005. Saline systems1:1-14.
Orosa M, Franqueira D, Cid A, Abalde J (2001) Carotenoid accumulation inHaematococcus pluvialis in mixotrophic growth. Biotechnology Letters23:373-378.
Park J, Chyun J, Kim Y, Line L, Chew B (2010) Astaxanthin decreased oxidativestress and inflammation and enhanced immune response in humans. Nutrition&Metabolism7:18.
Pascal AA, Liu Z, Broess K, van Oort B, van Amerongen H, Wang C, Horton P,Robert B, Chang W, Ruban A (2005) Molecular basis of photoprotection andcontrol of photosynthetic light-harvesting. Nature436:134-137.
Pastenes C, Pimentel P, Lillo J (2005) Leaf movements and photoinhibition in relationto water stress in field-grown beans. Journal of Experimental Botany56:425-433.
Patel VJ, Thalassinos K, Slade SE, Connolly JB, Crombie A, Murrell JC, Scrivens JH(2009) A Comparison of Labeling and Label-Free Mass Spectrometry-BasedProteomics Approaches. Journal of Proteome Research8:3752-3759.
PEARSON CK, WILSON SB, SCHAFFER R, ROSS AW (1993) NAD turnover andutilisation of metabolites for RNA synthesis in a reaction sensing the redoxstate of the cytochrome b6f complex in isolated chloroplasts. European journalof biochemistry218:397-404.
Peled E, Pick U, Zarka A, Shimoni E, Leu S, Boussiba S (2012) Light-induced oilglobule migration in Haematococcus pluvialis (Chlorophyceae). Journal ofPhycology48:1209-1219.
Perkins PD, Miller CA, Kuhlmann FE (2005) Comparison of Different Approachesfor the Label-Free Relative Quantitation of proteins.53rd ASMS Conference，2005.
Pink M, Verma N, Rettenmeier AW, Schmitz-Spanke S (2010) CBB staining protocolwith higher sensitivity and mass spectrometric compatibility. Electrophoresis31:593-598.
Rabbani S, Beyer P, Lintig Jv, Hugueney P, Kleinig H (1998) Induced β-CaroteneSynthesis Driven by Triacylglycerol Deposition in the Unicellular AlgaDunaliella bardawil. Plant Physiology116:1239-1248.
Raja R, Hemaiswarya S, Rengasamy R (2007) Exploitation of Dunaliella forβ-carotene production.Applied microbiology and biotechnology74:517-523.
Raven JA (2013) Rubisco: still the most abundant protein of Earth? New Phytologist198:1-3.
Rismani-Yazdi H, Haznedaroglu BZ, Bibby K, Peccia J (2011) Transcriptomesequencing and annotation of the microalgae Dunaliella tertiolecta: pathwaydescription and gene discovery for production of next-generation biofuels.BMC genomics12:148.
Rumeau D, Peltier G, Cournac L (2007) Chlororespiration and cyclic electron flowaround PSI during photosynthesis and plant stress response. Plant, Cell&Environment30:1041-1051.
Saito K (2004) Sulfur assimilatory metabolism. The long and smelling road. PlantPhysiology136:2443-2450.
Sandmann G (1995) Carotenoid biosynthesis in microorganisms and plants. EJBReviews1994: Springer Berlin Heidelberg. pp.129-146.
Sarada R, Tripathi U, Ravishankar GA (2002) Influence of stress on astaxanthinproduction in Haematococcus pluvialis grown under different cultureconditions. Process Biochemistry37:623-627.
Sch gger H (2006) Tricine-SDS-PAGE. Nat Protocols1:16-22.
Schluesener D, Fischer F, Kruip J, R gner M, Poetsch A (2005) Mapping themembrane proteome of Corynebacterium glutamicum. PROTEOMICS5:1317-1330.
Schoefs B, Rmiki N-E, Rachadi J, Lemoine Y (2001) Astaxanthin accumulation inHaematococcus requires a cytochrome P450hydroxylase and an activesynthesis of fatty acids. FEBS Letters500:125-128.
Schwender J, Goffman F, Ohlrogge JB, Shachar-Hill Y (2004) Rubisco without theCalvin cycle improves the carbon efficiency of developing green seeds. Nature432:779-782.
Shaish A, Avron M, Pick U, Ben-Amotz A (1993) Are active oxygen species involvedin induction of β-carotene in Dunaliella bardawil? Planta190:363-368.
Shen Y, Zhao R, Berger SJ, Anderson GA, Rodriguez N, Smith RD (2002)High-efficiency nanoscale liquid chromatography coupled on-line with massspectrometry using nanoelectrospray ionization for proteomics. Analyticalchemistry74:4235-4249.
Shikanai T (2007) Cyclic Electron Transport Around Photosystem I: GeneticApproaches. Annual Review of Plant Biology58:199-217.
Smith BM, Morrissey PJ, Guenther JE, Nemson JA, Harrison MA, Allen JF, Melis A(1990) Response of the Photosynthetic Apparatus in Dunaliella salina (GreenAlgae) to Irradiance Stress. Plant physiology93:1433-1440.
Smith RD (2002) Trends in mass spectrometry instrumentation for proteomics. Trendsin Biotechnology20: s3-s7.
Spolaore P, Joannis-Cassan C, Duran E, Isambert A (2006) Commercial applicationsof microalgae. Journal of Bioscience and Bioengineering101:87-96.
Staehelin LA (2005) Chloroplast structure: from chlorophyll granules tosupra-molecular architecture of thylakoid membranes
Discoveries in Photosynthesis. In: Govindjee G, Beatty JT, Gest H, Allen JF, editors.Discoveries in photosynthesis: Springer Netherlands. pp.717-728.
Stoll DR, Wang X, Carr PW (2008) Comparison of the practical resolving power ofone-and two-dimensional high-performance liquid chromatography analysis ofmetabolomic samples. Analytical chemistry80:268-278.
Strain HH (1944) Chloroplast Pigments. Annual Review of Biochemistry13:591-610.
Sun S, Wang F, Li C, Qin S, Zhou M, Ding L, Pang S, Duan D, Wang G, Yin B (2008)Emerging challenges: Massive green algae blooms in the Yellow Sea. NaturePrecedings.
Suzuki K, Masaki H, Takei M (1996) External preparation for skin. Japanese Patent#08073312[in Japanese].
Tan S, Cunningham FX, Youmans M, Grabowski B, Sun Z, Gantt E (1995)Cytochrome f loss in astaxanthin-accumulating red cells of Haematococcuspluvialis (Chlorophyceae): Comparasion of photosynthetic activity,photosynthetic enzymes, and thylakoid membrane polyppeptides in red andgreen cells. Journal of Phycology31:897-905.
Tanaka A, Melis A (1997) Irradiance-Dependent Changes in the Size and Compositionof the Chlorophyll a-b Light-Harvesting Complex in the Green AlgaDunaliella salina. Plant and Cell Physiology38:17-24.
Thornber JP (1975) Chlorophyll-proteins: light-harvesting and reaction centercomponents of plants. Annual review of plant physiology26:127-158.
Tran N-P, Park J-K, Kim ZH, Lee C-G (2009) Influence of sodium orthovanadate onthe production of astaxanthin from green algae Haematococcus lacustris.Biotechnology and Bioprocess Engineering14:322-329.
Tran N-P, Park J-K, Lee C-G (2009) Proteomics analysis of proteins in green algaHaematococcus lacustris (Chlorophyceae) expressed under combined stress ofnitrogen starvation and high irradiance. Enzyme and Microbial Technology45:241-246.
Tripathi U, Sarada R, Rao SR, Ravishankar GA (1999) Production of astaxanthin inHaematococcus pluvialis cultured in various media. Bioresource Technology68:197-199.
Tripathi U, Sarada R, Ravishankar G (2002) Effect of culture conditions on growth ofgreen alga—Haematococcus pluvialis and astaxanthin production. ActaPhysiologiae Plantarum24:323-329.
Turujman SA, Wamer WG, Wei RR, Albert RH (1997) Rapid liquid chromatographicmethod to distinguish wild salmon from aquacultured salmon fed syntheticastaxanthin. Journal of AOAC International80:622-632.
Vasil'ev S, Bruce D (2004) Optimization and Evolution of Light Harvesting inPhotosynthesis: The Role of Antenna Chlorophyll Conserved betweenPhotosystem II and Photosystem I. The Plant Cell Online16:3059-3068.
Vasilikiotis C, Melis A (1995) The role of chloroplast-encoded protein biosynthesis onthe rate of D1protein degradation in Dunaliella salina. PhotosynthesisResearch45:147-155.
Wang B, Zarka A, Trebst A, Boussiba S (2003) Astaxanthin accumulation inHaematococcus pluvialis (Chlorophyceae) as an active photoprotectiveprocess under high irradiance. Journal of Phycology39:1116-1124.
Wang J, Sommerfeld M, Hu Q (2009) Occurrence and environmental stress responsesof two plastid terminal oxidases in Haematococcus pluvialis (Chlorophyceae).Planta230:191-203.
Wang S-B, Chen F, Sommerfeld M, Hu Q (2004) Proteomic analysis of molecularresponse to oxidative stress by the green alga Haematococcus pluvialis(Chlorophyceae). Planta220:17-29.
Wang S-B, Hu Q, Sommerfeld M, Chen F (2003) An optimized protocol for isolationof soluble proteins from microalgae for two-dimensional gel electrophoresisanalysis. Journal of Applied Phycology15:485-496.
Wang SB, Hu Q, Sommerfeld M, Chen F (2004) Cell wall proteomics of the greenalga Haematococcus pluvialis (Chlorophyceae). Proteomics4:692-708.
Wang X, Willén R, Wadstr m T (2000) Astaxanthin-Rich Algal Meal and Vitamin CInhibitHelicobacter pylori Infection in BALB/cA Mice. Antimicrobial Agentsand Chemotherapy44:2452-2457.
Wayama M, Ota S, Matsuura H, Nango N, Hirata A, Kawano S (2013)Three-dimensional ultrastructural study of oil and astaxanthin accumulationduring encystment in the green alga Haematococcus pluvialis. PLoS One8:e53618.
Whitelegge J (2003) Thylakoid membrane proteomics. Photosynthesis Research78:265-277.
Wittig I, Braun H-P, Schagger H (2006) Blue native PAGE. Nat Protocols1:418-428.
Wright PC, Noirel J, Ow SY, Fazeli A (2012) A review of current proteomicstechnologies with a survey on their widespread use in reproductive biologyinvestigations. Theriogenology77:738-765.e752.
Wright S, Jeffrey S, Mantoura R (2005) Phytoplankton pigments in oceanography:guidelines to modern methods: Unesco Pub.
Xiao F-G, Shen L, Ji H-F (2011) On photoprotective mechanisms of carotenoids inlight harvesting complex. Biochemical and Biophysical ResearchCommunications414:1-4.
Xie X, Gao S, Gu W, Pan G, Wang G (2013) Desiccation induces accumulations ofantheraxanthin and zeaxanthin in intertidal macro-alga Ulva pertusa(Chlorophyta). PLoS ONE8: e72929.
Xie X, Gu W, Gao S, Lu S, Li J, Pan G, Wang G, Shen S (2013) Alternative electrontransports participate in the maintenance of violaxanthin de-epoxidase activityof Ulva sp. under low irradiance. PLoS ONE8: e78211.
Yamazaki J-y, Kozu A, Fukunaga Y (2006) Characterization of Chlorophyll–proteinComplexes Isolated from Two Marine Green Algae, Bryopsis maxima andUlva pertusa, Growing in the Intertidal Zone. Photosynthesis Research89:19-25.
Yamazaki J-y, Suzuki T, Maruta E, Kamimura Y (2005) The stoichiometry andantenna size of the two photosystems in marine green algae, Bryopsis maximaand Ulva pertusa, in relation to the light environment of their natural habitat.Journal of Experimental Botany56:1517-1523.
Yates JR, Ruse CI, Nakorchevsky A (2009) Proteomics by Mass Spectrometry:Approaches, Advances, and Applications. Annual Review of BiomedicalEngineering11:49-79.
Ye V, Bhatia S (2012) Pathway engineering strategies for production of beneficialcarotenoids in microbial hosts. Biotechnology Letters:1-10.
Yoshimura S, Ranjbar R, Inoue R, Katsuda T, Katoh S (2006) Effective utilization oftransmitted light for astaxanthin production by Haematococcus pluvialis.Journal of Bioscience and Bioengineering102:97-101.
Z rb C, Schmitt S, Neeb A, Karl S, Linder M, Schubert S (2004) The biochemicalreaction of maize (Zea mays L.) to salt stress is characterized by a mitigationof symptoms and not by a specific adaptation. Plant Science167:91-100.
Zhang BY, Geng YH, Li ZK, Hu HJ, Li YG (2009) Production of astaxanthin fromHaematococcus in open pond by two-stage growth one-step process.Aquaculture295:275-281.
Zhang XW, Gong XD, Chen F (1999) Dynamics and stability analysis of the growthand astaxanthin production system of Haematococcus pluvialis. Journal ofIndustrial Microbiology and Biotechnology23:133-137.
Zhekisheva M, Boussiba S, Khozin-Goldberg I, Zarka A, Cohen Z (2002)Accumulation of oleic acid in Haematococcus pluvialis (Chlorophyceae)under nitrogen starvation or high light is correlated with that of astaxanthinesters. Journal of Phycology38:325-331.
Zhou D, Zhang L, Zhang S, Fu H, Chen J (2010) Hydrothermal liquefaction ofmacroalgae Enteromorpha prolifera to bio-oil. Energy&Fuels24:4054-4061.
Zhu H, Bilgin M, Snyder M (2003) Proteomics. Annual Review of Biochemistry72:783-812.
Zhu W, Smith JW, Huang CM (2010) Mass spectrometry-based label-free quantitativeproteomics. J Biomed Biotechnol2010:840518.
韦韬,顾文辉,李健,张波,潘光华,朱大玲,王广策(2012)不同碳氮浓度对雨生红球藻生长及虾青素累积的影响.海洋科学36:55-61.
韦韬,顾文辉,李健,张波,潘光华,朱大玲,王广策(2013)雨生红球藻的光周期效应.植物学报48:168-173.
Aflalo C, Meshulam Y, Zarka A, Boussiba S (2007) On the relative efficiency of two-vs. one-stage production of astaxanthin by the green alga Haematococcuspluvialis. Biotechnology and bioengineering98:300-305.
Alberte RS, Friedman AL, Gustafson DL, Rudnick MS, Lyman H (1981)Light-harvesting systems of brown algae and diatoms. Isolation andcharacterization of chlorophyll ac and chlorophyll afucoxanthinpigment-protein complexes. Biochimica et Biophysica Acta(BBA)-Bioenergetics635:304-316.
Alric J (2010) Cyclic electron flow around photosystem I in unicellular green algae.Photosynthesis Research106:47-56.
Apel K, Hirt H (2004) Reactive oxygen species: Metabolism, Oxidative Stress, andSignal Transduction. Annual Review of Plant Biology55:373-399.
Asara JM, Christofk HR, Freimark LM, Cantley LC (2008) A label‐freequantification method by MS/MS TIC compared to SILAC and spectralcounting in a proteomics screen. Proteomics8:994-999.
Baas-Becking L (1931) Salt effects on swarmers of Dunaliella viridis Teod. TheJournal of general physiology14:765.
Bachi A, Bonaldi T (2008) Quantitative proteomics as a new piece of the systemsbiology puzzle. Journal of Proteomics71:357-367.
Banse K (2002) Steemann Nielsen and the zooplankton. Hydrobiologia480:15-28.
Bar E, Rise M, Vishkautsan M, Arad S (1995) Pigment and Structural Changes inChlorella zofingiensis upon Light and Nitrogen Stress. Journal of PlantPhysiology146:527-534.
Bassi R, Caffarri S (2000) Lhc proteins and the regulation of photosynthetic lightharvesting function by xanthophylls. Photosynthesis Research64:243-256.
Ben-Amotz A (1995) New mode of Dunaliella biotechnology: two-phase growth forβ-carotene production. Journal ofApplied Phycology7:65-68.
Bj rn L, Papageorgiou G, Blankenship R, Govindjee (2009) A viewpoint: Whychlorophyll a? Photosynthesis Research99:85-98.
Blum H, Beier H, Gross HJ (1987) Improved silver staining of plant proteins, RNAand DNA in polyacrylamide gels. Electrophoresis8:93-99.
Borowitzka MA, Huisman JM, Osborn A (1991) Culture of the astaxanthin-producinggreen alga Haematococcus pluvialis1. Effects of nutrients on growth and celltype. Journal of Applied Phycology3:295-304.
Boussiba S (2000) Carotenogenesis in the green alga Haematococcus pluvialis:Cellular physiology and stress response. Physiologia Plantarum108:111-117.
Boussiba S, Bing W, Yuan J-P, Zarka A, Chen F (1999) Changes in pigments profile inthe green alga Haeamtococcus pluvialis exposed to environmental stresses.Biotechnology Letters21:601-604.
Boussiba S, Fan L, Vonshak A, Lester P (1992) Enhancement and determination ofastaxanthin accumulation in green alga Haematococcus pluvialis. San Diego,CA, ETATS-UNIS: Academic Press.
Bradford MM (1976) A rapid and sensitive method for the quantitation of microgramquantities of protein utilizing the principle of protein-dye binding. AnalyticalBiochemistry72:248-254.
Buchanan BB, Balmer Y (2005) Redox regulation: A broadening horizon. AnnualReview of Plant Biology56:187-220.
Buchanan BB, Gruissem W, Jones RL (2000) Biochemistry&molecular biology ofplants, Chapter12Photosynthesis: American Society of Plant PhysiologistsRockville.
Caffarri S, Kouril R, Kereiche S, Boekema EJ, Croce R (2009) Functionalarchitecture of higher plant photosystem II supercomplexes. EMBO J28:3052-3063.
Chang HI (2010) Protective effect of the natural pigment on naproxen induced gastriculceration in rat. Journal of Biotechnology150:68-68.
Chen Z, Wang G, Niu J (2012) Variation in Rubisco and other photosyntheticparameters in the life cycle of Haematococcus pluvialis. Chinese Journal ofOceanology and Limnology30:136-145.
Chua NH, Bennoun P (1975) Thylakoid membrane polypeptides of Chlamydomonasreinhardtii: wild-type and mutant strains deficient in photosystem II reactioncenter. Proceedings of the National Academy of Sciences72:2175-2179.
Collins AM, Jones HDT, Han D, Hu Q, Beechem TE, Timlin JA (2011) CarotenoidDistribution in Living Cells of Haematococcus pluvialis (Chlorophyceae).PLoS ONE6: e24302.
Cort A, Ozturk N, Akpinar D, Unal M, Yucel G, Ciftcioglu A, Yargicoglu P, Aslan M(2010) Suppressive effect of astaxanthin on retinal injury induced by elevatedintraocular pressure. Regulatory Toxicology and Pharmacology58:121-130.
Cunningham FX, Gantt E (1998) GENES AND ENZYMES OF CAROTENOIDBIOSYNTHESIS IN PLANTS. Annual Review of Plant Physiology and PlantMolecular Biology49:557-583.
Cunningham FX, Gantt E (2011) Elucidation of the Pathway to Astaxanthin in theFlowers of Adonis aestivalis. The Plant Cell Online23:3055-3069.
Czeczuga B (1974) Carotenoids in Euglena rubida Mainx. Comparative Biochemistryand Physiology Part B: Comparative Biochemistry48:349-354.
D'Amici GM, Huber CG, Zolla L (2009) Separation of thylakoid membrane proteinsby sucrose gradient ultracentrifugation or blue native-SDS-PAGEtwo-dimensional electrophoresis. Methods Mol Biol528:61-70.
Damiani MC, Popovich CA, Constenla D, Leonardi PI (2010) Lipid analysis inHaematococcus pluvialis to assess its potential use as a biodiesel feedstock.Bioresource Technology101:3801-3807.
Davison PA, Hunter CN, Horton P (2002) Overexpression of [beta]-carotenehydroxylase enhances stress tolerance in Arabidopsis. Nature418:203-206.
Del Campo JA, Rodríguez H, Moreno J, Vargas Má, Rivas J, Guerrero MG (2004)Accumulation of astaxanthin and lutein in Chlorella zofingiensis(Chlorophyta). Applied Microbiology and Biotechnology64:848-854.
Deo PM, Biswal B (2001) Response of senescing cotyledons of clusterbean to waterstress in moderate and low light: Possible photoprotective role of β‐carotene.Physiologia plantarum112:47-54.
Ding L, Fei X, Lu Q, Deng Y, Lian S (2009) The possibility analysis of habitats,origin and reappearance of bloom green alga (Enteromorpha prolifera) oninshore of western Yellow Sea. Chinese Journal of Oceanology and Limnology27:421-424.
Droop MR (1954) Conditions governing haematochrome formation and loss in thealga Haematococcus pluvialis flotow. Archiv für Mikrobiologie20:391-397.
Droop MR (1955) Carotenogenesis in Haematococcus pluvialis. Nature175:42-42.
Ellis RJ (1979) Most abundant protein in the world. Trends in Biochemical Sciences4:241-244.
Ellis RJ (1979) Msot abundant protein in the world. Trends in Biochemical Sciences4:241-244.
Ephritikhine G, Ferro M, Rolland N (2004) Plant membrane proteomics. PlantPhysiology and Biochemistry42:943-962.
Escoubas J-M, Lomas M, LaRoche J, Falkowski PG (1995) Light intensity regulationof cab gene transcription is signaled by the redox state of the plastoquinonepool. Proceedings of the National Academy of Sciences92:10237-10241.
Eshaghi S, Andersson B, Barber J (1999) Isolation of a highly active PSII-LHCIIsupercomplex from thylakoid membranes by a direct method. FEBS Letters446:23-26.
Fábregas J, Otero A, Maseda A, Domínguez A (2001) Two-stage cultures for theproduction of Astaxanthin from Haematococcus pluvialis. Journal ofBiotechnology89:65-71.
Fabregas J, Dominguez A, Regueiro M, Maseda A, Otero A (2000) Optimization ofculture medium for the continuous cultivation of the microalgaHaematococcus pluvialis. Applied Microbiology and Biotechnology53:530-535.
Fan L, Vonshak A, Boussiba S (1994) Effect of temperature and irradiance on growthof Haematococcus pluvialis (chlorophyceae)1. Journal of Phycology30:829-833.
Field CB, Behrenfeld MJ, Randerson JT, Falkowski P (1998) Primary Production ofthe Biosphere: Integrating Terrestrial and Oceanic Components. Science281:237-240.
Flotow J v (1844) Haematococcus pluvialis. Verh Kais Leopold-Carol Akad NaturfNova Acta Acad Caes Leopold Carolin Nat Curios:411-606.
Frey PA, Hegeman AD, Ruzicka FJ (2008) The Radical SAM Superfamily. CriticalReviews in Biochemistry and Molecular Biology43:63-88.
Fromm HJ, Hargrove MS (2012) Photosynthesis
Essentials of Biochemistry. Springer Berlin Heidelberg. pp.317-329.
Funk C, Schr der WP, Green BR, Renger G, Andersson B (1994) The intrinsic22kDaprotein is a chlorophyll-binding subunit of photosystem II. FEBS Letters342:261-266.
Gao S, Chen X, Yi Q, Wang G, Pan G, Lin A, Peng G (2010) A Strategy for theProliferation of Ulva prolifera, Main Causative Species of Green Tides, withFormation of Sporangia by Fragmentation. PLoS ONE5: e8571.
Gao S, Shen S, Wang G, Niu J, Lin A, Pan G (2011) PSI-driven cyclic electron flowallows intertidal macro-algae Ulva sp.(Chlorophyta) to survive in desiccatedconditions. Plant Cell Physiol52:885-893.
Goodwin TW (1955) Carotenoids. Annual Review of Biochemistry24:497-522.
Green B, Durnford D (1996) The chlorophyll-carotenoid proteins of oxygenicphotosynthesis. Annual review of plant biology47:685-714.
Gu W, Xie X, Gao S, Zhou W, Pan G, Wang G (2013) Comparison of different cells ofHaematococcus pluvialis reveals an extensive acclimation mechanism duringits aging process: From a perspective of photosynthesis. PLoS One8.
Guerin M, Huntley ME, Olaizola M (2003) Haematococcus astaxanthin: applicationsfor human health and nutrition. Trends in Biotechnology21:210-216.
Hagen C, Grünewald K, Schmidt S, Müller J (2000) Accumulation of secondarycarotenoids in flagellates of Haematococcus pluvialis (Chlorophyta) isaccompanied by an increase in per unit chlorophyll productivity ofphotosynthesis. European Journal of Phycology35:75-82.
Hagen C, Grünewald K, Xyl nder M, Rothe E (2001) Effect of cultivation parameterson growth and pigment biosynthesis in flagellated cells of Haematococcuspluvialis. Journal of Applied Phycology13:79-87.
Haglund K, Bj rk M, Ramazanov Z, García-Reina G, Pedersén M (1992) Role ofcarbonic anhydrase in photosynthesis and inorganic-carbon assimilation in thered alga Gracilaria tenuistipitata. Planta187:275-281.
Han D, Li Y, Hu Q (2013) Astaxanthin in microalgae: pathways, functions andbiotechnological implications. Algae28:131-147.
Han DX, Wang JF, Sommerfeld M, Hu Q (2012) Susceptibility and protectivemechanisms of motile and non motile cells of Haematococcus Pluvialis(Chlorophyceae) to photooxidative stress. Journal of Phycology48:693-705.
Hankamer B, Morris E, Nield J, Gerle C, Barber J (2001) Three-dimensional structureof the photosystem II core dimer of higher plants determined by electronmicroscopy. Journal of structural biology135:262-269.
Harker M, Hirschberg J (1997) Biosynthesis of ketocarotenoids in transgeniccyanobacteria expressing the algal gene for [beta]-C-4-oxygenase, crtO. FEBSLetters404:129-134.
Harker M, Tsavalos AJ, Young AJ (1996) Factors responsible for astaxanthinformation in the Chlorophyte Haematococcus pluvialis. BioresourceTechnology55:207-214.
Hejazi M, Holwerda E, Wijffels R (2004) Milking microalga Dunaliella salina for β‐carotene production in two‐phase bioreactors. Biotechnology andbioengineering85:475-481.
Helbig AO, Heck AJR, Slijper M (2010) Exploring the membraneproteome--Challenges and analytical strategies. Journal of Proteomics73:868-878.
Higuera-Ciapara I, Félix-Valenzuela L, Goycoolea FM (2006) Astaxanthin: A Reviewof its Chemistry and Applications. Critical Reviews in Food Science andNutrition46:185-196.
Holmgren A, Johansson C, Berndt C, Lonn M, Hudemann C, Lillig C (2005) Thiolredox control via thioredoxin and glutaredoxin systems. Biochemical SocietyTransactions33:1375-1377.
Holtin K, Kuehnle M, Rehbein J, Schuler P, Nicholson G, Albert K (2009)Determination of astaxanthin and astaxanthin esters in the microalgaeHaematococcus pluvialis by LC-(APCI)MS and characterization ofpredominant carotenoid isomers by NMR spectroscopy. Analytical andBioanalytical Chemistry395:1613-1622.
Horton P, Ruban AV, Walters RG (1996) Regulation of light harvesting in green plants.Annual Review of Plant Physiology and Plant Molecular Biology47:655-684.
Ivanov AG, Rosso D, Savitch LV, Stachula P, Rosembert M, Oquist G, Hurry V, HünerNPA (2012) Implications of alternative electron sinks in increased resistanceof PSII and PSI photochemistry to high light stress in cold-acclimatedArabidopsis thaliana. Photosynthesis Research113:191-206.
Jahns P, Holzwarth AR (2012) The role of the xanthophyll cycle and of lutein inphotoprotection of photosystem II. Biochim Biophys Acta1817:182-193.
Jiang C-D, Gao H-Y, Zou Q, Jiang G-M, Li L-H (2006) Leaf orientation,photorespiration and xanthophyll cycle protect young soybean leaves againsthigh irradiance in field. Environmental and Experimental Botany55:87-96.
JIN E, LEE C (2006) Secondary Carotenoid Accumulation in Haematococcus(Chlorophyceae): Biosynthesis, Regulation, and Biotechnology. J MicrobiolBiotechnol16:821-831.
Johnson EA, Lewis MJ (1979) Astaxanthin formation by the yeast Phaffia rhodozyma.J Gen Microbiol115:173-183.
Kügler M, J nsch L, Kruft V, Schmitz UK, Braun H-P (1997) Analysis of thechloroplast protein complexes by blue-native polyacrylamide gelelectrophoresis (BN-PAGE). Photosynthesis Research53:35-44.
Kajiwara S, Kakizono T, Saito T, Kondo K, Ohtani T, Nishio N, Nagai S, Misawa N(1995) Isolation and functional identification of a novel cDNA for astaxanthinbiosynthesis from Haematococcus pluvialis, and astaxanthin synthesis inEscherichia coli. Plant Molecular Biology29:343-352.
Kamath BS, Srikanta BM, Dharmesh SM, Sarada R, Ravishankar GA (2008) Ulcerpreventive and antioxidative properties of astaxanthin from Haematococcuspluvialis. European Journal of Pharmacology590:387-395.
Kang C, Lee J, Park T, Sim S (2007) Complementary limiting factors of astaxanthinsynthesis during photoautotrophic induction of Haematococcus pluvialis: C/Nratio and light intensity. Applied Microbiology and Biotechnology74:987-994.
Kargul J, Barber J (2008) Photosynthetic acclimation: Structural reorganisation oflight harvesting antenna–role of redox-dependent phosphorylation of majorand minor chlorophyll a/b binding proteins. FEBS Journal275:1056-1068.
Kasahara M, Kagawa T, Oikawa K, Suetsugu N, Miyao M, Wada M (2002)Chloroplast avoidance movement reduces photodamage in plants. Nature420:829-832.
Katz A, Waridel P, Shevchenko A, Pick U (2007) Salt-induced changes in the plasmamembrane proteome of the halotolerant alga Dunaliella salina as revealed byblue native gel electrophoresis and nano-LC-MS/MS analysis. Molecular&Cellular Proteomics6:1459-1472.
Kim S-J, Chang HI (2010) Astaxanthin represses adipogenesis in3T3-L1cells.Journal of Biotechnology150:308-308.
Kim S, Sandusky P, Bowlby NR, Aebersold R, Green BR, Vlahaskis S, Yocum CF,Pichersky E (1992) Characterization of a spinach psbS cDNA encoding the22kDa protein of photosystem II. FEBS Letters314:67-71.
Kindlund P (2011) Astaxanthin. Nutrafoods10:27-31.
Kobayashi M (2003) Astaxanthin biosynthesis enhanced by reactive oxygen species inthe green alga Haematococcus pluvialis. Biotechnology and BioprocessEngineering8:322-330.
Kobayashi M, Kakizono T, Nagai S (1993) Enhanced carotenoid biosynthesis byoxidative stress in acetate-induced cyst cells of a green unicellular alga,Haematococcus pluvialis. Applied And Environmental Microbiology59:867.
Kobayashi M, Kakizono T, Yamaguchi K, Nishio N, Nagai S (1992) Growth andastaxanthin formation of Haematococcus pluvialis in heterotrophic andmixotrophic conditions. Journal of Fermentation and Bioengineering74:17-20.
Kobayashi M, Katsuragi T, Tani Y (2001) Enlarged and astaxanthin-accumulating cystcells of the green alga Haematococcus pluvialis. Journal of Bioscience andBioengineering92:565-568.
Kobayashi M, Kurimura Y, Kakizono T, Nishio N, Tsuji Y (1997) Morphologicalchanges in the life cycle of the green alga Haematococcus pluvialis. Journal ofFermentation and Bioengineering84:94-97.
Kota U, Goshe MB (2011) Advances in qualitative and quantitative plant membraneproteomics. Phytochemistry72:1040-1060.
Kristoffersen AS, Svensen, Ssebiyonga N, Erga SR, Stamnes JJ, Frette (2012)Chlorophyll a and NADPH Fluorescence Lifetimes in the MicroalgaeHaematococcus pluvialis (Chlorophyceae) under Normal andAstaxanthin-Accumulating Conditions. Appl Spectrosc66:1216-1225.
Lamers PP, van de Laak CCW, Kaasenbrood PS, Lorier J, Janssen M, De Vos RCH,Bino RJ, Wijffels RH (2010) Carotenoid and fatty acid metabolism inlight-stressed Dunaliella salina. Biotechnology and Bioengineering106:638-648.
Layer G, Heinz DW, Jahn D, Schubert W-D (2004) Structure and function of radicalSAM enzymes. Current Opinion in Chemical Biology8:468-476.
Leister D (2003) Chloroplast research in the genomic age. Trends in Genetics19:47-56.
Lello Z, Anna MT, Christian GH (2004) Thylakoid Membrane Proteome: Separationby HPLC and Identification by Accurate Molecular Mass Determinations.Current Proteomics1:247-260.
Lemoine Y, Schoefs B (2010) Secondary ketocarotenoid astaxanthin biosynthesis inalgae: a multifunctional response to stress. Photosynthesis Research106:155-177.
Li J, Zhu D, Niu J, Shen S, Wang G (2011) An economic assessment of astaxanthinproduction by large scale cultivation of Haematococcus pluvialis.Biotechnology Advances29:568-574.
Li X-P, Bj rkman O, Shih C, Grossman AR, Rosenquist M, Jansson S, Niyogi KK(2000) A pigment-binding protein essential for regulation of photosyntheticlight harvesting. Nature403:391-395.
Li X-P, Gilmore AM, Caffarri S, Bassi R, Golan T, Kramer D, Niyogi KK (2004)Regulation of Photosynthetic Light Harvesting Involves Intrathylakoid LumenpH Sensing by the PsbS Protein. Journal of Biological Chemistry279:22866-22874.
Li X-P, Müller-Moulé P, Gilmore AM, Niyogi KK (2002) PsbS-dependentenhancement of feedback de-excitation protects photosystem II fromphotoinhibition. Proceedings of the National Academy of Sciences99:15222-15227.
Li Y, Sommerfeld M, Chen F, Hu Q (2008) Consumption of oxygen by astaxanthinbiosynthesis: a protective mechanism against oxidative stress inHaematococcus pluvialis (Chlorophyceae). J Plant Physiol165:1783-1797.
Li Y, Sommerfeld M, Chen F, Hu Q (2010) Effect of photon flux densities onregulation of carotenogenesis and cell viability of Haematococcus pluvialis(Chlorophyceae). Journal of Applied Phycology22:253-263.
Li Z, Ahn TK, Avenson TJ, Ballottari M, Cruz JA, Kramer DM, Bassi R, Fleming GR,Keasling JD, Niyogi KK (2009) Lutein Accumulation in the Absence ofZeaxanthin Restores Nonphotochemical Quenching in the Arabidopsisthaliana npq1Mutant. The Plant Cell Online21:1798-1812.
Li Z, Wakao S, Fischer BB, Niyogi KK (2009) Sensing and responding to excess light.Annu Rev Plant Biol60:239-260.
Lichtenthaler HK, Babani F (2004) Light Adaptation and Senescence of thePhotosynthetic Apparatus. Changes in Pigment Composition, ChlorophyllFluorescence Parameters and Photosynthetic Activity. In: Papageorgiou GC,Govindjee, editors. Chlorophyll a Fluorescence: Springer Netherlands. pp.713-736.
Lin A, Shen S, Wang J, Yan B (2008) Reproduction diversity of Enteromorphaprolifera. Journal of integrative plant biology50:622-629.
Lin R (2002) Xanthophyll Cycle and Its Molecular Mechanism in Photoprotection.Acta Botanica Sinica44.
Liska AJ, Shevchenko A, Pick U, Katz A (2004) Enhanced photosynthesis and redoxenergy production contribute to salinity tolerance in Dunaliella as revealed byhomology-based proteomics. Plant physiology136:2806-2817.
Long SP (1994) Photoinhibition of photosynthesis in nature. Annu Rev Plant PhysiolPlant Mol Biol45:633-662.
Mackinney G (1952) Carotenoids. Annual Review of Biochemistry21:473-492.
Majeran W, Zybailov B, Ytterberg AJ, Dunsmore J, Sun Q, van Wijk KJ (2008)Consequences of C4Differentiation for Chloroplast Membrane Proteomes inMaize Mesophyll and Bundle Sheath Cells. Molecular&Cellular Proteomics7:1609-1638.
Marouga R, David S, Hawkins E (2005) The development of the DIGE system:2Dfluorescence difference gel analysis technology. Analytical and BioanalyticalChemistry382:669-678.
Maxwell DP, Laudenbach DE, Huner N (1995) Redox Regulation of Light-HarvestingComplex II and cab mRNA Abundance in Dunaliella salina. Plant Physiology109:787-795.
McDonald AE, Ivanov AG, Bode R, Maxwell DP, Rodermel SR, Hüner NPA (2011)Flexibility in photosynthetic electron transport: The physiological role ofplastoquinol terminal oxidase (PTOX). Biochimica et Biophysica Acta (BBA)-Bioenergetics1807:954-967.
McLean K, Sabri M, Marshall K, Lawson R, Lewis D, Clift D, Balding P, Dunford A,Warman A, McVey J (2005) Biodiversity of cytochrome P450redox systems.Biochemical Society Transactions33:796-801.
Melkozernov AN, Barber J, Blankenship RE (2005) Light Harvesting in PhotosystemI Supercomplexes,. Biochemistry45:331-345.
Miflin BJ, Lea PJ (1976) The pathway of nitrogen assimilation in plants.Phytochemistry15:873-885.
Miki W (1991) Biological function and activity of animal carotenoids. Pure ApplChem:141-146.
Milledge J (2010) Commercial application of microalgae other than as biofuels: abrief review. Reviews in Environmental Science and Biotechnology:1-11.
Monteoliva L, Albar JP (2004) Differential proteomics: an overview of gel andnon-gel based approaches. Briefings in functional genomics&proteomics3:220-239.
Moore GF, Brudvig GW (2011) Energy Conversion in Photosynthesis: A Paradigm forSolar Fuel Production. Annual Review of Condensed Matter Physics2:303-327.
Mou S, Zhang X, Dong M, Fan X, Xu J, Cao S, Xu D, Wang W, Ye N (2013)Photoprotection in the green tidal alga Ulva prolifera: role of LHCSR andPsbS proteins in response to high light stress. Plant Biol (Stuttg).
Murakami A (1997) Quantitative analysis of77K fluorescence emission spectra inSynechocystis sp. PCC6714and Chlamydomonas reinhardtii with variable PSI/PS II stoichiometries. Photosynthesis Research53:141-148.
Murata N, Nishimura M, Takamiya A (1966) Fluorescence of chlorophyll inphotosynthetic systems III. Emission and action spectra offluorescence—Three emission bands of chlorophyll a and the energy transferbetween two pigment systems. Biochimica et Biophysica Acta (BBA)-Biophysics including Photosynthesis126:234-243.
Naumann B, Hippler M (2007) Insights into chloroplast proteomics: from basicprinciples to new horizonsCell and Molecular Biology of Plastids. In: Bock R, editor. Cell and MolecularBiology of Plastids: Springer Berlin/Heidelberg. pp.371-407.
Neidhardt J, Benemann J, Zhang L, Melis A (1998) Photosystem-II repair andchloroplast recovery from irradiance stress: relationship between chronicphotoinhibition, light-harvesting chlorophyll antenna size and photosyntheticproductivity in Dunaliella salina (green algae). Photosynthesis Research56:175-184.
Neilson KA, Ali NA, Muralidharan S, Mirzaei M, Mariani M, Assadourian G, Lee A,van Sluyter SC, Haynes PA (2011) Less label, more free: Approaches inlabel-free quantitative mass spectrometry. PROTEOMICS11:535-553.
Nielsen ES (1952) The Use of Radio-active Carbon (C14) for Measuring OrganicProduction in the Sea. Journal du Conseil18:117-140.
Niu J, Hu H, Hu S, Wang G, Peng G, Sun S (2010) Analysis of expressed sequencetags from the Ulva prolifera (Chlorophyta). Chinese Journal of Oceanologyand Limnology28:26-36.
Niyogi KK (1999) Photoprotection revisited: genetic and molecular approaches.Annual Review of Plant Physiology and Plant Molecular Biology50:333-359.
Oeljeklaus S, Meyer HE, Warscheid B (2009) Advancements in plant proteomicsusing quantitative mass spectrometry. Journal of Proteomics72:545-554.
Old WM, Meyer-Arendt K, Aveline-Wolf L, Pierce KG, Mendoza A, Sevinsky JR,Resing KA, Ahn NG (2005) Comparison of Label-free Methods forQuantifying Human Proteins by Shotgun Proteomics. Molecular&CellularProteomics4:1487-1502.
Oren A (2005) A hundred years of Dunaliella research:1905-2005. Saline Systems1:2.
Oren A (2005) A hundred years of Dunaliella research:1905–2005. Saline systems1:1-14.
Orosa M, Franqueira D, Cid A, Abalde J (2001) Carotenoid accumulation inHaematococcus pluvialis in mixotrophic growth. Biotechnology Letters23:373-378.
Park J, Chyun J, Kim Y, Line L, Chew B (2010) Astaxanthin decreased oxidativestress and inflammation and enhanced immune response in humans. Nutrition&Metabolism7:18.
Pascal AA, Liu Z, Broess K, van Oort B, van Amerongen H, Wang C, Horton P,Robert B, Chang W, Ruban A (2005) Molecular basis of photoprotection andcontrol of photosynthetic light-harvesting. Nature436:134-137.
Pastenes C, Pimentel P, Lillo J (2005) Leaf movements and photoinhibition in relationto water stress in field-grown beans. Journal of Experimental Botany56:425-433.
Patel VJ, Thalassinos K, Slade SE, Connolly JB, Crombie A, Murrell JC, Scrivens JH(2009) A Comparison of Labeling and Label-Free Mass Spectrometry-BasedProteomics Approaches. Journal of Proteome Research8:3752-3759.
PEARSON CK, WILSON SB, SCHAFFER R, ROSS AW (1993) NAD turnover andutilisation of metabolites for RNA synthesis in a reaction sensing the redoxstate of the cytochrome b6f complex in isolated chloroplasts. European journalof biochemistry218:397-404.
Peled E, Pick U, Zarka A, Shimoni E, Leu S, Boussiba S (2012) Light-induced oilglobule migration in Haematococcus pluvialis (Chlorophyceae). Journal ofPhycology48:1209-1219.
Perkins PD, Miller CA, Kuhlmann FE (2005) Comparison of Different Approachesfor the Label-Free Relative Quantitation of proteins.53rd ASMS Conference，2005.
Pink M, Verma N, Rettenmeier AW, Schmitz-Spanke S (2010) CBB staining protocolwith higher sensitivity and mass spectrometric compatibility. Electrophoresis31:593-598.
Rabbani S, Beyer P, Lintig Jv, Hugueney P, Kleinig H (1998) Induced β-CaroteneSynthesis Driven by Triacylglycerol Deposition in the Unicellular AlgaDunaliella bardawil. Plant Physiology116:1239-1248.
Raja R, Hemaiswarya S, Rengasamy R (2007) Exploitation of Dunaliella forβ-carotene production.Applied microbiology and biotechnology74:517-523.
Raven JA (2013) Rubisco: still the most abundant protein of Earth? New Phytologist198:1-3.
Rismani-Yazdi H, Haznedaroglu BZ, Bibby K, Peccia J (2011) Transcriptomesequencing and annotation of the microalgae Dunaliella tertiolecta: pathwaydescription and gene discovery for production of next-generation biofuels.BMC genomics12:148.
Rumeau D, Peltier G, Cournac L (2007) Chlororespiration and cyclic electron flowaround PSI during photosynthesis and plant stress response. Plant, Cell&Environment30:1041-1051.
Saito K (2004) Sulfur assimilatory metabolism. The long and smelling road. PlantPhysiology136:2443-2450.
Sandmann G (1995) Carotenoid biosynthesis in microorganisms and plants. EJBReviews1994: Springer Berlin Heidelberg. pp.129-146.
Sarada R, Tripathi U, Ravishankar GA (2002) Influence of stress on astaxanthinproduction in Haematococcus pluvialis grown under different cultureconditions. Process Biochemistry37:623-627.
Sch gger H (2006) Tricine-SDS-PAGE. Nat Protocols1:16-22.
Schluesener D, Fischer F, Kruip J, R gner M, Poetsch A (2005) Mapping themembrane proteome of Corynebacterium glutamicum. PROTEOMICS5:1317-1330.
Schoefs B, Rmiki N-E, Rachadi J, Lemoine Y (2001) Astaxanthin accumulation inHaematococcus requires a cytochrome P450hydroxylase and an activesynthesis of fatty acids. FEBS Letters500:125-128.
Schwender J, Goffman F, Ohlrogge JB, Shachar-Hill Y (2004) Rubisco without theCalvin cycle improves the carbon efficiency of developing green seeds. Nature432:779-782.
Shaish A, Avron M, Pick U, Ben-Amotz A (1993) Are active oxygen species involvedin induction of β-carotene in Dunaliella bardawil? Planta190:363-368.
Shen Y, Zhao R, Berger SJ, Anderson GA, Rodriguez N, Smith RD (2002)High-efficiency nanoscale liquid chromatography coupled on-line with massspectrometry using nanoelectrospray ionization for proteomics. Analyticalchemistry74:4235-4249.
Shikanai T (2007) Cyclic Electron Transport Around Photosystem I: GeneticApproaches. Annual Review of Plant Biology58:199-217.
Smith BM, Morrissey PJ, Guenther JE, Nemson JA, Harrison MA, Allen JF, Melis A(1990) Response of the Photosynthetic Apparatus in Dunaliella salina (GreenAlgae) to Irradiance Stress. Plant physiology93:1433-1440.
Smith RD (2002) Trends in mass spectrometry instrumentation for proteomics. Trendsin Biotechnology20: s3-s7.
Spolaore P, Joannis-Cassan C, Duran E, Isambert A (2006) Commercial applicationsof microalgae. Journal of Bioscience and Bioengineering101:87-96.
Staehelin LA (2005) Chloroplast structure: from chlorophyll granules tosupra-molecular architecture of thylakoid membranes
Discoveries in Photosynthesis. In: Govindjee G, Beatty JT, Gest H, Allen JF, editors.Discoveries in photosynthesis: Springer Netherlands. pp.717-728.
Stoll DR, Wang X, Carr PW (2008) Comparison of the practical resolving power ofone-and two-dimensional high-performance liquid chromatography analysis ofmetabolomic samples. Analytical chemistry80:268-278.
Strain HH (1944) Chloroplast Pigments. Annual Review of Biochemistry13:591-610.
Sun S, Wang F, Li C, Qin S, Zhou M, Ding L, Pang S, Duan D, Wang G, Yin B (2008)Emerging challenges: Massive green algae blooms in the Yellow Sea. NaturePrecedings.
Suzuki K, Masaki H, Takei M (1996) External preparation for skin. Japanese Patent#08073312[in Japanese].
Tan S, Cunningham FX, Youmans M, Grabowski B, Sun Z, Gantt E (1995)Cytochrome f loss in astaxanthin-accumulating red cells of Haematococcuspluvialis (Chlorophyceae): Comparasion of photosynthetic activity,photosynthetic enzymes, and thylakoid membrane polyppeptides in red andgreen cells. Journal of Phycology31:897-905.
Tanaka A, Melis A (1997) Irradiance-Dependent Changes in the Size and Compositionof the Chlorophyll a-b Light-Harvesting Complex in the Green AlgaDunaliella salina. Plant and Cell Physiology38:17-24.
Thornber JP (1975) Chlorophyll-proteins: light-harvesting and reaction centercomponents of plants. Annual review of plant physiology26:127-158.
Tran N-P, Park J-K, Kim ZH, Lee C-G (2009) Influence of sodium orthovanadate onthe production of astaxanthin from green algae Haematococcus lacustris.Biotechnology and Bioprocess Engineering14:322-329.
Tran N-P, Park J-K, Lee C-G (2009) Proteomics analysis of proteins in green algaHaematococcus lacustris (Chlorophyceae) expressed under combined stress ofnitrogen starvation and high irradiance. Enzyme and Microbial Technology45:241-246.
Tripathi U, Sarada R, Rao SR, Ravishankar GA (1999) Production of astaxanthin inHaematococcus pluvialis cultured in various media. Bioresource Technology68:197-199.
Tripathi U, Sarada R, Ravishankar G (2002) Effect of culture conditions on growth ofgreen alga—Haematococcus pluvialis and astaxanthin production. ActaPhysiologiae Plantarum24:323-329.
Turujman SA, Wamer WG, Wei RR, Albert RH (1997) Rapid liquid chromatographicmethod to distinguish wild salmon from aquacultured salmon fed syntheticastaxanthin. Journal of AOAC International80:622-632.
Vasil'ev S, Bruce D (2004) Optimization and Evolution of Light Harvesting inPhotosynthesis: The Role of Antenna Chlorophyll Conserved betweenPhotosystem II and Photosystem I. The Plant Cell Online16:3059-3068.
Vasilikiotis C, Melis A (1995) The role of chloroplast-encoded protein biosynthesis onthe rate of D1protein degradation in Dunaliella salina. PhotosynthesisResearch45:147-155.
Wang B, Zarka A, Trebst A, Boussiba S (2003) Astaxanthin accumulation inHaematococcus pluvialis (Chlorophyceae) as an active photoprotectiveprocess under high irradiance. Journal of Phycology39:1116-1124.
Wang J, Sommerfeld M, Hu Q (2009) Occurrence and environmental stress responsesof two plastid terminal oxidases in Haematococcus pluvialis (Chlorophyceae).Planta230:191-203.
Wang S-B, Chen F, Sommerfeld M, Hu Q (2004) Proteomic analysis of molecularresponse to oxidative stress by the green alga Haematococcus pluvialis(Chlorophyceae). Planta220:17-29.
Wang S-B, Hu Q, Sommerfeld M, Chen F (2003) An optimized protocol for isolationof soluble proteins from microalgae for two-dimensional gel electrophoresisanalysis. Journal of Applied Phycology15:485-496.
Wang SB, Hu Q, Sommerfeld M, Chen F (2004) Cell wall proteomics of the greenalga Haematococcus pluvialis (Chlorophyceae). Proteomics4:692-708.
Wang X, Willén R, Wadstr m T (2000) Astaxanthin-Rich Algal Meal and Vitamin CInhibitHelicobacter pylori Infection in BALB/cA Mice. Antimicrobial Agentsand Chemotherapy44:2452-2457.
Wayama M, Ota S, Matsuura H, Nango N, Hirata A, Kawano S (2013)Three-dimensional ultrastructural study of oil and astaxanthin accumulationduring encystment in the green alga Haematococcus pluvialis. PLoS One8:e53618.
Whitelegge J (2003) Thylakoid membrane proteomics. Photosynthesis Research78:265-277.
Wittig I, Braun H-P, Schagger H (2006) Blue native PAGE. Nat Protocols1:418-428.
Wright PC, Noirel J, Ow SY, Fazeli A (2012) A review of current proteomicstechnologies with a survey on their widespread use in reproductive biologyinvestigations. Theriogenology77:738-765.e752.
Wright S, Jeffrey S, Mantoura R (2005) Phytoplankton pigments in oceanography:guidelines to modern methods: Unesco Pub.
Xiao F-G, Shen L, Ji H-F (2011) On photoprotective mechanisms of carotenoids inlight harvesting complex. Biochemical and Biophysical ResearchCommunications414:1-4.
Xie X, Gao S, Gu W, Pan G, Wang G (2013) Desiccation induces accumulations ofantheraxanthin and zeaxanthin in intertidal macro-alga Ulva pertusa(Chlorophyta). PLoS ONE8: e72929.
Xie X, Gu W, Gao S, Lu S, Li J, Pan G, Wang G, Shen S (2013) Alternative electrontransports participate in the maintenance of violaxanthin de-epoxidase activityof Ulva sp. under low irradiance. PLoS ONE8: e78211.
Yamazaki J-y, Kozu A, Fukunaga Y (2006) Characterization of Chlorophyll–proteinComplexes Isolated from Two Marine Green Algae, Bryopsis maxima andUlva pertusa, Growing in the Intertidal Zone. Photosynthesis Research89:19-25.
Yamazaki J-y, Suzuki T, Maruta E, Kamimura Y (2005) The stoichiometry andantenna size of the two photosystems in marine green algae, Bryopsis maximaand Ulva pertusa, in relation to the light environment of their natural habitat.Journal of Experimental Botany56:1517-1523.
Yates JR, Ruse CI, Nakorchevsky A (2009) Proteomics by Mass Spectrometry:Approaches, Advances, and Applications. Annual Review of BiomedicalEngineering11:49-79.
Ye V, Bhatia S (2012) Pathway engineering strategies for production of beneficialcarotenoids in microbial hosts. Biotechnology Letters:1-10.
Yoshimura S, Ranjbar R, Inoue R, Katsuda T, Katoh S (2006) Effective utilization oftransmitted light for astaxanthin production by Haematococcus pluvialis.Journal of Bioscience and Bioengineering102:97-101.
Z rb C, Schmitt S, Neeb A, Karl S, Linder M, Schubert S (2004) The biochemicalreaction of maize (Zea mays L.) to salt stress is characterized by a mitigationof symptoms and not by a specific adaptation. Plant Science167:91-100.
Zhang BY, Geng YH, Li ZK, Hu HJ, Li YG (2009) Production of astaxanthin fromHaematococcus in open pond by two-stage growth one-step process.Aquaculture295:275-281.
Zhang XW, Gong XD, Chen F (1999) Dynamics and stability analysis of the growthand astaxanthin production system of Haematococcus pluvialis. Journal ofIndustrial Microbiology and Biotechnology23:133-137.
Zhekisheva M, Boussiba S, Khozin-Goldberg I, Zarka A, Cohen Z (2002)Accumulation of oleic acid in Haematococcus pluvialis (Chlorophyceae)under nitrogen starvation or high light is correlated with that of astaxanthinesters. Journal of Phycology38:325-331.
Zhou D, Zhang L, Zhang S, Fu H, Chen J (2010) Hydrothermal liquefaction ofmacroalgae Enteromorpha prolifera to bio-oil. Energy&Fuels24:4054-4061.
Zhu H, Bilgin M, Snyder M (2003) Proteomics. Annual Review of Biochemistry72:783-812.
Zhu W, Smith JW, Huang CM (2010) Mass spectrometry-based label-free quantitativeproteomics. J Biomed Biotechnol2010:840518.