Potential for green microalgae to produce hydrogen, pharmaceuticals and other high value products in a combined process

Figures & data

Figure 1.  Production of valuable metabolites from algae in commercial use today (A), compared to the proposed processes where stress factors are applied to induce both hydrogen production and production of valuable metabolites simultaneously (B), or in sequence (C).

Figure 2.  Overview of the combined process for production of hydrogen and bioactive metabolites. Green microalgae can be cultured under optimal growth conditions, followed by exposure to stress conditions (high light intensity, nutrient deprivation). The algal biomass can be harvested and used for different purposes, for example direct use as food supplement, aquaculture and animal fodders. Several valuable components can be extracted for the purpose of pharmaceutical industry, cosmetics or other types of industrial purposes.

Table 1.  Algal species which have been studied and shown ability to produce hydrogen and/or metabolites with pharmaceutical/ industrial interest and algae which are being used today for commercial purposes.

Genus	Species^†	Hydrogen production^‡	Hydrogen –S^‡	Health food	Aquaculture/ animal feed	Valuable metabolites^§,#
Ankistrodesmus	spp.				FS (Turker et al., 2003)	α-linolenic acid (Ben-Amotz et al., 1985)
						Hexadecatetraenoic acid (Ben-Amotz et al., 1985)
	angustus	D (Brand et al., 1989)
	braunii	D (Kessler, 1973)				Astaxanthin (Borowitzka, 1988)
	falcatus	D (Bishop et al., 1977)
	spiralis					Mycosporine-like amino acids (Xiong et al., 1999)
Botryococcus	sppbraunii					α-linolenic acid (Chiang et al., 2004)
						Linoleic acid (Chiang et al., 2004)
						Butylated hydroxytoluene (Babu and Wu, 2008)
						Lutein (Rao et al., 2006)
						β-carotene (Borowitzka, 1988)
						Extracellular alkadienes (wax for cosmetics) (Mendes et al., 2003)
Brachiomonas	submarina					Glycerol (Ahmadand Hellebust, 1986)
Carteria	spp				BP (de Pauw and Persoone, 1988, Lavens and Sorgeloos, 1996)
	crusifera	D (Brand et al., 1989)
	eugametos	D (Brand et al., 1989)
Chlamydobotrys	stellata	D (Boichenko et al., 1992)
Chlamydocapsa	spp					Lutein (Leya et al., 2009)
						Cantaxanthin (Leya et al., 2009)
						Astaxanthin (Leya et al., 2009)
Chlamydomonas	spp.				BL, BP, FZ, MR, BS, BP,	Glycerol (Miyasaka et al., 1998)
					FS (Lavens and Sorgeloos, 1996, Opuszynski, 1981)	Vitamin B (Vilchez et al., 1997)
						α-linolenic acid (Poerschmann et al., 2004)
						Hexadecatetraenoic acid (Poerschmann et al., 2004)
	acidophila					Lutein (Garbayo et al., 2008)
						β-carotene (Garbayo et al., 2008)
						Violaxanthin (Garbayo et al., 2008)
	applanata	D (Brand et al., 1989)				Haemagglutinin (Chu et al., 2004)
	asymmetrica					Haemagglutinin (Chu et al., 2004)
	chlamydogama	D (Brand et al., 1989)
	debaryana	D (Healey, 1970)
	dorsoventralis	D (Brand et al., 1989)				Haemagglutinin (Chu et al., 2004)
	dysosmos	D (Healey, 1970)
	elliptica	D (Brand et al., 1989)
	eugametos	D (Kessler, 1974)				Vitamin B (Uhlik and Gowans, 1974)
	euryale	D (Skjånes et al., 2008)	D (Skjånes et al., 2008)
	hindakii	D (Brand et al., 1989) (noctigama)
	hydra	D (Brand et al., 1989)
	intermedia	D (Kessler, 1974)
	moewusii	D (Healey, 1970)	D (Meuser et al., 2009)			α-linolenic acid (Arisz et al., 2000)
						Linoleic acid (Arisz et al., 2000)
						Hexadecatetraenoic acid (Arisz et al., 2000)
	nivalis					Astaxanthin (Remias et al., 2005)
						Phenolic antioxidants (Duval et al., 2000)
						Mycosporine-like amino acids (Duval et al., 2000)
	noctigama	D (Winkler et al., 2002)	D (Skjånes et al., 2008)			Haemagglutinin (Chu et al., 2004)
	pulsatilla					Glycerol (Ahmadand Hellebust, 1986)
	pulvinata					Haemagglutinin (Chu et al., 2004)
	reinhardtii	D (Kessler, 1974)	D (Melis et al., 2000)			Transgenic proteins (Griesbeck et al., 2005)
						Toxic fatty acids (McCracken et al., 1980, Spruell, 1984)
						Glycerol (León and Galván, 1999)
						Vitamin C (Borowitzka, 1988)
						Vitamin E (Borowitzka, 1988)
	simplex					Haemagglutinin (Chu et al., 2004)
	texensis	D (Brand et al., 1989)
	ulvaensis					Haemagglutinin (Chu et al., 2004)
	vectensis	D (Skjånes et al., 2008)	D (Skjånes et al., 2008)
Chlorella	spp.		D (Chader et al., 2009, Timmins et al., 2009a)	NP (Pulz and Gross, 2004, Yamaguchi, 1997)	MR, BS, FS, BL, ML, FZ (Apt and Behrens, 1999, de Pauw and Persoone, 1988, Duerr et al., 1998, Pulz and Gross, 2004, Turker et al., 2003)	Vitamin C (Running et al., 2002)
						Vitamin B (Borowitzka, 1988)
						Extract w antimicrobial and anticancer activity (Ördög et al., 2004)
	ellipsiodea					Violaxanthin (Cha et al., 2008)
						Antheraxanthin (Cha et al., 2008)
						Zeaxanthin (Cha et al., 2008)
						Extract w antimicrobial activity (Chu et al., 2004)
						Extract w apoptose inducing effect (Cha et al., 2008)
						Haemagglutinin (Chu et al., 2004)
	emersonii					Canthaxanthin (Bhosale and Bernstein, 2005)
	fusca	D (Kessler, 1974)				Lutein (Del Campo et al., 2000)
						Astaxanthin (Borowitzka, 1988)
						Canthaxanthin (Borowitzka, 1988)
						Extract w antimicrobial activity (Chu et al., 2004)
						Haemagglutinin (Chu et al., 2004)
	homosphaera	D (Kessler, 1974)				α-linolenic acid (Ahlgren et al., 1992)
	kessleri	D (Kessler, 1974) (Parachlorella)				Extract w antimicrobial activity (Chu et al., 2004)
						Haemagglutinin (Chu et al., 2004)
	luteovirtidis					Haemagglutinin (Chu et al., 2004)
						Mycosporine-like amino acids (Karsten et al., 2007)
	minutissima				BL, PL (Duerr et al., 1998)	Eicosapentaenoic acid (Seto et al., 1984)
						Extract w antimicrobial and anticancer activity (Ördög et al., 2004)
						Mycosporine-like amino acids (Xiong et al., 1999)
	protothecoides	D (Bishop et al., 1977)				Lutein (Chen, 1998, Shi et al., 2006)
						Extract w antimicrobial activity (Chu et al., 2004)
						Haemagglutinin (Chu et al., 2004)
						Linoleic acid (Day et al., 2009)
						Vitamin C (Running et al., 2002)
	pyrenoidosa	D (Brand et al., 1989)	D (Skjånes et al., 2008)			Extract w antimicrobial/ antifungal activity (Abedin and Taha, 2008, Chu et al., 2004)
						Haemagglutinin (Chu et al., 2004)
						Polysaccharide w immunostim effect (Pugh et al., 2001, Yang et al., 2006)
						Polysaccharide w antitumor activity (Sheng et al., 2007)
						Lutein (Wu et al,. 2007)
						Inhibition of dioxin maternal transfer (Nakano et al., 2005)
						Vitamin B (Becker, 2004b)
						Vitamin E (Becker, 2004b)
	saccharophila					Haemagglutinin (Chu et al., 2004)
	salina	D (Chader et al., 2009)	D (Chader et al., 2009)
	sorokiniana	D (Bishop et al., 1977)	D (Chader et al., 2009)			Fatty acid extract w/medical activity (Chou et al., 2008)
						Sulfoxidation biocatalyst activity (Daligault et al., 2006)
						Lutein (Matsukawa et al., 2000)
						α /β-carotene (Matsukawa et al., 2000)
						Vitamin E (Matsukawa et al., 2000)
						Haemagglutinin (Chu et al., 2004)
						Mycosporine-like amino acids (Xiong et al., 1999)
	sphaerica					Mycosporine-like amino acids (Karsten et al., 2007)
	stigmatophora					Polysaccharide w anti-innflammatory and immunosuppressive activity (Guzman et al., 2003)
						Extract w effect on the central nervous system (Laguna et al., 1993)
	vacuolata	D (Bishop et al., 1977)				Extract w antimicrobial activity (Chu et al., 2004)
	vulgaris	D (Healey, 1970)	D (Timmins et al., 2009a)	NP (Pulz and Gross, 2004, Raja et al., 2008, Rodriguez-Garcia and Guil-Guerrero, 2008)	FS, HF (Gouveia et al., 2002, Janczyk et al., 2009, Raja et al., 2008)	Lutein (Cha et al., 2008)
						Canthaxanthin (Gouveia et al., 1996, Mendes et al., 2003)
						Astaxanthin (Gouveia et al., 1996, Mendes et al., 2003)
						Vitamin C (Borowitzka, 1988)
						Vitamin E (Borowitzka, 1988)
						Glycoprotein w anti-cancer activity (Hasegawa et al., 2002, Tanaka et al., 1998)
						Sulfonated polysaccharide, antioxidant (Mohamed, 2008)
						Haemagglutinin (Chu et al., 2004)
						Eicosatrienoic acid (Becker, 2004b)
						α-linolenic acid (Piorreck et al., 1984)
						Linoleic acid (Piorreck et al., 1984)
						Extract w toxin-protective effect (Kim et al., 2009a)
	xanthella					Haemagglutinin (Chu et al., 2004)
	zofingiensis					Lutein (Bhosale and Bernstein, 2005, Del Campo et al., 2004)
						Astaxanthin (Bhosale and Bernstein, 2005, Del Campo et al., 2004)
						Canthaxanthin (Pelah et al., 2004)
						Haemagglutinin (Chu et al., 2004)
Chlorococcum	sp	D (Boichenko and Hoffmann, 1994)			BL, BP, FZ, MR, BS BP (Lavens and Sorgeloos, 1996)	Astaxanthine (Leya et al., 2009, Liu and Lee, 2000, Yuan et al., 2002)
						Adonixanthin (Yuan et al., 2002)
						Canthaxanthin (Yuan et al., 2002)
						Lutein (Leya et al., 2009)
						y-linolenic acid (Ohta et al., 1995)
	citriforme					Lutein (Del Campo et al., 2000)
	humicolum	D (Boichenko and Hoffmann, 1994)
	littorale	D (Ueno et al., 1999)				Ethanol (Ueno et al., 1998)
	submarinum	D (Kamp et al., 2008)	D (Kamp et al., 2008)			Glycerol (Blackwell and Gilmour, 1991)
	vacuolatum	D (Kessler, 1974)
Chlorogonium	elongatum	D (Kreuzberg, 1984)
Chloromonas	nivalis					Lutein (Leya et al., 2009)
						Zeaxanthin (Leya et al., 2009)
Chlorosarcinopsis	ereme	D (Boichenko and Hoffmann, 1994)
Chodatella	balatonica	D (Boichenko and Hoffmann, 1994)
Coelastrella	striolata					Canthaxanthin (Abe et al., 2007)
						Astaxanthin (Abe et al,. 2007)
						β-carotene (Abe et al., 2007)
Coelastrum	microporum				FS (Opuszynski, 1981)
	proboscideum	D (Bishop et al., 1977)				Lutein (Del Campo et al., 2000)
Coccomyxa	lacustris	D (Boichenko and Hoffmann, 1994)
Cosmarium	botrytis	D (Brand et al., 1989)
	turpinii	D (Brand et al., 1989)
Desmococcus	olivaceus					Extract w antimicrobial and anticancer activity (Ördög et al., 2004)
Desmodesmus	spp.	D (Timmins et al., 2009a)	D (Timmins et al., 2009a)
	subspicatus	D (Skjånes et al., 2008)	D (Skjånes et al., 2008)
	quadricauda	D (Healey, 1970) (Scenedesmus)
Dictyococcus	pseudovarians	D (Boichenko and Hoffmann, 1994)
Dunaliella	sp.			NP (Finney et al., 1984, Pulz and Gross, 2004, Yamaguchi, 1997)	BP, BS, MR, PL (Apt and Behrens, 1999, D'Souza and Loneragan, 1999, Lavens and Sorgeloos, 1996, Pulz and Gross, 2004)	Glutathione (Li et al., 2004)
						Glycerol (Kaçka and Dönmez, 2008)
	bardawil			NP (Mokady et al., 1989)	FS (Shpigel et al., 2006)	β-carotene (Rabbani et al., 1998)
						Extract w anti-tumor effect (Fujii et al., 1993)
						Hexadecatetraenoic acid (Fried et al., 1982)
						Linoleic acid (Fried et al., 1982)
						α-linolenic acid (Fried et al., 1982)
	primolecta					y-linolenic acid (Ohta et al., 1995)
						Pheophoride-like comp w anti-virus activity (Ohta et al., 1995)
	salina			NP (Pulz and Gross, 2004, Raja et al., 2008)	NS (Raja et al., 2008)	β-carotene (Coesel et al., 2008, Hejazi and Wijffels, 2003)
						Zeaxanthin (Bhosale and Bernstein, 2005)
						Extract w protective effect against fibrosarcoma (Raja et al., 2007)
						Extract w antimicrobial activity (Herrero et al., 2006)
						Glycerol (Hadi et al., 2008)
						α-linolenic acid (Ben-Amotz et al., 1985)
	tertiolecta				BL, FS (Carballo-Cárdenas et al., 2003)	Lutein (Barbosa et al., 2005)
						β-carotene (Barbosa et al., 2005)
						Vitamin C (Barbosa et al., 2005)
						Vitamin E (Carballo-Cárdenas et al., 2003)
						Extract w central nervous system effect (Villar et al., 1992)
						Hexadecatetraenoic acid (D'Souza and Loneragan, 1999, Gouveia and Oliveira, 2009)
						α-linolenic acid (D'Souza and Loneragan, 1999, Gouveia and Oliveira, 2009)
						Linoleic acid (Gouveia and Oliveira, 2009)
Eudorina	elegans	D (Brand et al., 1989)			FS (Opuszynski, 1981)
Golenkinia	sp.	D (Boichenko and Hoffmann, 1994)
Gonium	sociale	D (Brand et al., 1989)
Haematococcus	spp.	D (Boichenko and Hoffmann, 1994)		NP (Lorenz et al., 2000, Raja et al., 2008)	NS (Lorenz et al., 2000, Pulz and Gross, 2004)	Astaxanthin (Ceron et al., 2007, He et al., 2007, Jin et al., 2006, Kang et al., 2005, Lorenz et al., 2000)
	droebakensis	D (Brand et al., 1989)
	lacustris					Canthaxanthin (Chattopadhyay et al., 2008)
						Astaxanthin (Chattopadhyay et al., 2008)
	pluvalis	D (Boichenko and Hoffmann, 1994)		NP (Pulz and Gross, 2004, Raja et al., 2008)	FS (Choubert et al., 2006, Raja et al., 2008)	Astaxanthin (Domínguez-Bocanegra et al., 2004, Kang et al., 2005, Zhekisheva et al., 2005)
Halochlorococcum	sp.	D (Boichenko and Hoffmann, 1994)
	saccatum	D (Greenbaum et al., 1983)
Kirchneriella	lunaris	D (Bishop et al., 1977)
	obesa	D (Boichenko and Hoffmann, 1994)
Lobochlamys	culleus	D (Meuser et al., 2009)	D (Meuser et al., 2009)
	segnis	D (Winkler et al., 2002)
Micromonas	sp				BL, BP (Lavens and Sorgeloos, 1996, Raja et al., 2008)
	pusilla				BL (Martinez-Fernandez and Southgate, 2007, Martinez-Fernandez et al., 2006)	α-Linolenic acid (Martinez-Fernandez et al., 2006)
						Stearidonic acid (Martinez-Fernandez et al., 2006)
						Neoxanthin (Egeland et al., 1995)
						Prasonoxanthin (Egeland et al., 1995)
						Micromonal (pigment) (Egeland et al., 1995, Latasa et al., 2004)
Monoraphidium						Astaxanthin (Fujii et al., 2008)
Muriella	aurantiaca					Lutein (Del Campo et al., 2000)
Muriellopsis	sp.					Lutein (Blanco et al., 2007, Del Campo et al., 2000)
						α-Linolenic acid (Blanco et al., 2007)
Nannochloris	sp.				BP, MR, SC (de Pauw and Persoone, 1988)	Eicosapentaenoic acid (Ben-Amotz et al., 1985)
	atomus					Eicosapentaenoic acid (Reitan et al., 1994)
						α-Linolenic acid (Reitan et al., 1994)
						Arachidonic acid (Reitan et al., 1994)
						Linoleic acid (Reitan et al., 1994)
	oculata				MR, BS (Duerr et al., 1998)
Neochloris	minuta	D (Boichenko and Hoffmann, 1994)
	oleaobundans				BP (Barnhart, 2006, Gatenby et al., 2003)	Linoleic acid (Gatenby et al., 2003)
						Linolenic acid (Gatenby et al., 2003)
Neospongiococcum	spp.					Astaxanthin (Borowitzka, 1988)
						Canthaxanthin (Borowitzka, 1988)
	gelatinosum					Lutein (Del Campo et al., 2000)
Oocystis	sp.	D (Skjånes et al., 2008)			BL, BS (Myrand and de la Noüe, 1983)	α-Linolenic acid (Patil et al., 2007)
						Linoleic acid (Patil et al., 2007)
	elliptica				FS (Opuszynski, 1981)
	parva	D (Boichenko and Hoffmann, 1994)
Pandorina	morum	D (Brand et al., 1989)			FS (Elliott et al., 2008, Opuszynski, 1981)
Parachlorella	kessleri	D (Kessler, 1974)
		(Chlorella)
Parietochloris	incisa					Arachidonic acid (Zhang et al., 2002a)
Pediastrum	boryanum	D (Boichenko and Hoffmann, 1994)			FS (Opuszynski, 1981)
	duplex				FS (Opuszynski, 1981)
Platymonas	striata					Lutein (Egeland et al., 1997)
	subcordiformis	D (Guan et al., 2004)	D (Guan et al., 2004)		PL, BL, BP, AL, BS, MR (Lavens and Sorgeloos, 1996, Tang et al., 2006)	Lutein (Egeland et al., 1997)
	(Tetraselmis)					Violaxanthin (Egeland et al., 1997)
	tetrahele					Lutein (Egeland et al., 1997)
						Neoxanthin (Egeland et al., 1997)
Prototheca	moriformis					Vitamin C (Running et al., 2002)
Pseudokirchneriella	subcapitata	D (Skjånes et al., 2008)			BS, MR (Belgis and Guido, 2003)	α-Linolenic acid (Patil et al., 2007)
						Linoleic acid (Patil et al., 2007)
Pseudospongiococcum	protococcoides	D (Boichenko and Hoffmann, 1994)
Pyramimonas	sp				BL, BP (Lavens and Sorgeloos, 1996, Raja et al., 2008)	Docosahexaenoic acid (Tzovenis et al., 2009)
						Lutein (Egeland et al., 1997)
	amylifera					β-carotene (Egeland et al., 1997)
						Siphonaxanthin (Egeland et al., 1997)
	cf. cordata					Sterols (Ponomarenko et al., 2004)
	obovata					β-carotene (Egeland et al., 1997)
	urceolata					Lutein (Egeland et al., 1997)
Scenedesmus	sp				FS, FZ, MR, BR (de Pauw and Persoone, 1988, Pulz and Gross, 2004, Turker et al., 2003)	Extract w antimicrobial and anticancer activity (Shon et al., 2004, Ördög et al., 2004)
						Mycosporine-like amino acids (Xiong et al., 1999)
	acutiformis	D (Soeder and Hegewald, 1988)
	acutus			NP (Kim et al., 2007)	FS (Kim et al., 2007)	α-linolenic acid (Ahlgren et al., 1992)
						Vitamin B (Borowitzka, 1988)
						Vitamin C (Borowitzka, 1988)
						Vitamin E (Borowitzka, 1988)
	almeriensis					Lutein (Ceron et al., 2008, Sánchez et al., 2008)
	armatus	D (Soeder and Hegewald, 1988)				Lutein (Tukaj et al., 2003)
	bicellularis	D (Soeder and Hegewald, 1988)
	communis	D (Soeder and Hegewald, 1988)
	komarekii	D (Soeder and Hegewald, 1988)
	obliquus	D (Gaffron and Rubin, 1942)				Astaxanthin (Qin et al., 2008)
						Vitamin B (Borowitzka, 1988)
						Vitamin C (Borowitzka, 1988)
						Vitamin E (Borowitzka, 1988)
						Hexadecatetraenoic acid (Becker, 2004b)
						α-Linolenic acid (Becker, 2004b)
						Linoleic acid (Gouveia and Oliveira, 2009)
	quadricauda	D (Healey, 1970)		NP (Kim et al., 2007)	FS (Kim et al., 2007, Opuszynski, 1981)	Haemagglutinin (Chu et al., 2004)
		(Desmodesmus)				α-linolenic acid (Ahlgren et al., 1992)
						Sulfonated polysaccharide (Mohamed, 2008)
						Vitamin B (Becker, 2004b)
						Vitamin C (Becker, 2004b)
						Vitamin E (Becker, 2004b)
						Extract w antimicrobial/ antifungal activity (Abedin and Taha , 2008)
	spinosus			NP (Kim et al., 2007)	FS (Kim et al., 2007)
Scotiella	chlorelloidea					Mycosporine-like amino acids (Xiong et al., 1999)
	nivalis	D (Boichenko and Hoffmann, 1994)
Selenastrum	sp.	D (Boichenko and Hoffmann, 1994)
	capricornutum					Haemagglutinin (Chu et al., 2004)
	gracile	D (Kessler, 1974)
Stichococcus	sp.					Mycosporine-like amino acids (Karsten et al., 2007)
Tetraedron	sp.				FS (Turker et al., 2003)
	bitridens	D (Brand et al., 1989)
	caudatum	D (Boichenko and Hoffmann, 1994)
Tetracystis	intermedium					Lutein (Del Campo et al., 2000)
	tetrasporum					Lutein (Del Campo et al., 2000)
Tetraselmis	sp				PL, BL, BP, AL, BS, MR, SC (D'Souza and Loneragan, 1999, Lavens and Sorgeloos, 1996, Puello-Cruz et al., 2009, Raja et al., 2008, Zmora et al., 2005)	Eicosapentaenoic acid (Patil et al., 2007, Reitan et al., 1994, Tzovenis et al., 2009)
						α-Linolenic acid (Patil et al., 2007, Reitan et al., 1994)
						β-carotene (Egeland et al., 1995)
						Lutein (Egeland et al., 1995)
						Violoaxanthin (Egeland et al., 1995)
						Neoxanthin (Egeland et al., 1995)
	chui				FS, BS, SC, MR, PL (Cunha et al., 2008, Duerr et al., 1998)
	kochinensis	D (Bhosale et al., 2009)
	subcordiformis	D (Guan et al., 2004)	D (Guan et al., 2004)		BL, BP (Duerr et al., 1998)
	suecica				BL, FS, PL, SC, MR, BS (Carballo-Cárdenas et al., 2003, D'Souza and Kelly,. 2000, Dahl et al., 2009, Puello-Cruz et al., 2009, Seixas et al., 2008, Souto et al., 2008)	Vitamin E (Carballo-Cárdenas et al., 2003)
						Sterols (Cardozo et al., 2007)
						Linoleic acid (D'Souza and Loneragan, 1999)
						α-Linolenic acid (D'Souza and Loneragan, 1999)
						Stearidonic acid (D'Souza and Loneragan, 1999)
						Eicosapentaenoic acid (D'Souza and Loneragan, 1999)
						Hexadecatetraenoic acid (D'Souza and Loneragan, 1999)
	tetrahele				FS, BS, SC, MR (Duerr et al., 1998)
	wettsteinii					β -carotene (Egeland et al., 1995)
						Lutein (Egeland et al., 1995)
Tetraspora	sp.	D (Maneeruttanarungroj et al., 2010)	D (Maneeruttanarungroj et al., 2010)
	cylindrica					Hexadecaedienoic acid (Ghazala et al., 2004)
						Heptadecaedienoic acid (Ghazala et al., 2004)
						Trimethyl dodecatrienoic acid (Ghazala et al., 2004)
						Extracts w antifungal, antibacterial and phytotoxic activity (Ghazala et al., 2004)
	gelatinosa					Trimethyl dodecatrienoic acid (Ghazala et al., 2004)
						Heptadecatrienoic acid (Ghazala et al., 2004)
						Pentadecatetraenoate (Ghazala et al., 2004)
						Extracts w phytotoxic activity (Ghazala et al., 2004)

The metabolites are produced under different culturing conditions, see text.

D, detected; NP, human consumption for nutritional purposes; HF, hen feed; PL, penaeid shrimp larvae feed; BL, bivalve mollusc larvae feed; ML, freshwater prawn larvae feed; BP, bivalve mollusc postlarvae feed; AL, abalone larvae feed; MR, marine rotifers feed; BS, brine shrimp feed; SC, saltwater copepods feed; FZ, freshwater zooplankton feed; FS, fish feed; NS, organisms not specified.

^†Some of the species might have changed names since the cited work was published. This has not been taken into account in this study, although in some cases synonyms are presented in parentheses below species names.

^‡Hydrogen production and hydrogen production during sulfur deprivation, refers to species where hydrogen production has been detected, or where expression of hydrogenase enzyme has been seen.

^§Polyunsaturated fatty acids (PUFA): Hexadecatetraenoic acid 16:4 (ω-3), α-linolenic acid (ALA) 18:3 (ω-3), Stearidonic acid (STD) 18:4 (ω-3), Eicosapentaenoic acid (EPA) 20:5 (ω-3), Eicosatrienoic acid (ETA) 20:3 (ω-3), Docosahexaenoic acid (DHA) 22:6 (ω-3), γ-linolenic acid (GLA) 18:3 (ω-6), Linoleic acid (LA) 18:2 (ω-6), Arachidonic acid (AA) 20:4(ω-6), Trimethyl-dodecatrienoate (15:3), Hexadecaedienoic acid (16:2), Heptadecaedienoic acid (17:2), Heptadecatrienoic acid (17:3), Pentadecatetraenoic acid (15:4).

^#Valuable metabolites listed in the table include components that are either produced in high amounts under stress conditions, or components that are produced under environmental conditions optimal for growth. In the case of carotenoids and fatty acids, the listed variants of these are in some cases produced in high amounts by the specific algae compared to the production by other algae species, in other cases represent dominant variants within the strain. See cited papers for details in each case.

Figure 3.  Schematic overview of suggested mechanisms for hydrogen production during sulfur deprivation in light, as it has been described for Chlamydomonas reinhardtii. Deprivation from sulfur leads to a degradation of PSII components, which partly inhibits the oxidation of water, and less oxygen is thereby produced in the photosystem. The low level of oxygen that is still produced in PSII is continuously consumed by the respiration, and the culture becomes anaerobic. Sulfur deprivation also leads to degradation of the enzymes in the Calvin cycle, causing this CO₂ fixation pathway and energy sink to come to a halt. When the Calvin cycle is no longer available for reducing CO₂, the whole system of PSII and PSI is reduced, creating a potentially dangerous situation for the algae. To remove the reductive pressure, the algae dispose of the electrons by transferring electrons from ferredoxin to hydrogenase. This enzyme then uses the reductive energy to form hydrogen which can easily be released from the cell. Depending on culturing conditions and other factors, a certain amount of electrons released in the form of hydrogen may originate from degradation of starch. This reducing power enters the electron transport chain from the PQ pool.

Figure 4.  Graphic summary of the most common stress reactions having influence on the synthesis of some important valuable metabolites in green algae.

Table 2.  Companies producing green microalgae for health food/ aquaculture/ animal feed/ pharmaceutical and industrial purposes.

Algae	Common products	Company	Country
Dunaliella	Biomass for health food, cosmetics or aquaculture feed, produced in the form of powder, paste, capsules or tablets. Extracts of β-carotene as color and antioxidant in food and cosmetics,produced in the form of powder or oil extract.	Aqua Carotene Ltd	Australia
		Betatene Ltd.	Australia
		Cognis Nutrition and Health	Australia
		Western Biotechnology Ltd.	Australia
		Inner Mongolia Biological Eng.	P.R.China
		Tianjin Lantai Biotechnology	P.R.China
		Cyanotech Corp.	Hawaii, USA
		ABL Biotechnologies Ltd.	India
		Parry Nutraceuticals	India
		Proalgen Biotech	India
		Nature Beta Technologies Ltd.	Israel
		Seambiotic	Israel
		Nikken Sohonsha Corporation	Japan
		Dutch State Mines	The Netherlands
		Easy Algae	Spain
Haematococcus	Biomass for health food, cosmetics or aquaculture feed, produced in the form of powder, paste, capsules or tablets. Extracts of astaxanthin for color and antioxidants in food and cosmetics,produced in the form of powder or oil extract.	WefirstBiotechnologyCo.,Ltd	P.R. China
		Blue BioTech Int.	Germany
		Cyanotech Corp.	Hawaii, USA
		Mera Pharmaceuticals	Hawaii, USA
		Parry Nutraceuticals	India
		AlgaTechnologies	Israel
		Dutch State Mines	The Netherlands
		Bioreal Inc.	Sweden, USA, Japan
Chlorella	Biomass for health food, cosmetics or animal feed, produced in the form of powder, paste, capsules or tablets.	Ocean Nutrition	Canada
		Bioprodukte Prof. Steinberg Produktions- undVertriebsGmbH& Co KG	Germany
		Blue BioTech Int.	Germany
		Nikken Sohonsha Corporation	Japan
		Sun Chlorella Corp.	Japan
		YaeyamaShokusanCO.Ltd.	Japan
		Taiwan Chlorella Manufacturing and Co.	Taiwan
Tetraselmis	Biomass for aquaculture feed, produced in the form of powder, paste or tablets.	Astaxa GmbH	Germany
		Seambiotic	Israel
		EasyAlgae	Spain
		Reed Mariculture	USA
Nannochloris	Biomass for aquaculture feed.	Seambiotic	Israel
Chlorococcum	Biomass for aquaculture feed.	Seambiotic	Israel

See text for details regarding use of algae products.

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