Photosynth Res

DOI 10.1007/s11120-014-9985-8
REVIEW
The plae o! al"ae #n a"$#%lt%$e& pol##es !o$ al"al '#o(ass
p$o)%t#on
E(#ly *. T$entaoste

+l#e *. *a$t#ne,

T#( -en.
Ree#/e)& 22 0an%a$y 2014 / +epte)& 12 1e'$%a$y 2014
The +%tho$2s3 2014. Th#s a$t#le #s p%'l#she) 4#th open aess at 5p$#n"e$l#n..o(
+'st$at +l"ae ha/e 'een %se) !o$ !oo) an)
n%t$ae%-t#als !o$ tho%san)s o! yea$s6 an) the
la$"e-sale %lt#/at#on o! al"ae6 o$ al"a%lt%$e6
has e7#ste) !o$ o/e$ hal! a ent%$y. *o$e $eently
al"ae ha/e 'een #)ent#!#e) an) )e/elope) as
$ene4a'le !%el so%$es6 an) the %lt#/at#on o!
al"al '#o(ass !o$ /a$#o%s p$o)%ts #s
t$ans#t#on#n" to o((e$#al-sale syste(s. It #s
$%#al )%$#n" th#s pe$#o) that #nst#t%t#onal
!$a(e4o$.s 2#.e.6 pol##es3 s%ppo$t an) p$o(ote
)e/elop-(ent an) o((e$#al#,at#on an)
ant##pate an) st#(%late the e/ol%t#on o! the al"al
'#o(ass #n)%st$y as a so%$e o! $ene4a'le !%els6
h#"h /al%e p$ote#n an) a$'ohy)$ates an) lo4-
ost )$%"s. 8a$"e-sale %lt#/at#on o! al"ae
(e$"es the !%n)a(ental aspets o! t$a)#t#onal
a"$#%lt%$al !a$(#n" an) a9%a%lt%$e. Desp#te
th#s o/e$lap6 al"a%lt%$e has not yet 'een
a!!o$)e) a pos#t#on 4#th#n a"$#%lt%$e o$ the
'ene!#ts asso#ate) 4#th #t. Va$#o%s !e)e$al an)
state a"$#%lt%$al s%ppo$t an) ass#stane
p$o"$a(s a$e %$$ently app$op$#ate) !o$ $ops6
'%t the#$ e7tens#on to al"al '#o(ass #s %ne$ta#n.
These p$o"$a(s a$e essent#al !o$ nasent
#n)%st$#es to eno%$a"e #n/est(ent6 '%#l)
#n!$ast$%t%$e6 )#sse(#nate tehn#al e7pe$#ene
an) #n!o$(at#on6 an) $eate (a$.ets. Th#s $e/#e4
)es$#'es the potent#al a"$#%lt%$al pol##es an)
:%est E)#to$& 5te/e *ay!#el).
E. *. T$entaoste 2&3
5$#pps Inst#t%t#on o! Oeano"$aphy6 ;n#/e$s#ty
o! <al#!o$n#a-5an D#e"o6 9500 :#l(an D$.6 8a
0olla6 <+ 9209=6 ;5+ e-(a#l&
et$enta>%s).e)%
+. *. *a$t#ne, T. -en.
5apph#$e Ene$"y6 In.6 =115 *e$$y!#el)
Ro46 5an D#e"o6
<+ 921216 ;5+
T. -en.
+l"ae ?#o(ass O$"an#,at#on6
125 5t. Pa%l 5t$eet6 P$eston6
*@ 559A56 ;5+
p$o"$a(s that o%l)
s%ppo$t al"al '#o(ass
%lt#/at#on6 an) the
'a$$#e$s to the e7pans#on
o! these p$o"$a(s to
al"ae.
Bey4o$)s +l"ae
'#o(ass
+"$#%lt%$e Pol#y
Rene4a'le ene$"y
+l"ae %lt#/at#on
Int$o)%t#on
+l"ae a$e s#(ple6
photosynthet#6 "ene$ally
a9%at# o$"an#s(s that6
l#.e plants6 %se ene$"y
!$o( s%nl#"ht to
se9%este$ a$'on )#o7#)e
2<O
2
3 !$o( the
at(osphe$e #nto '#o(ass
th$o%"h photosynthes#s.
Plants e/ol/e) !$o(
an#ent al"ae anesto$s6
an) the photosynthet#
(ah#ne$y #n 'oth plants
an) al"ae o$#"#nally a(e
!$o( the sa(e so%$e&
yano'ate$#a 21aloCn et
al. 2010 D 1ehl#n" et al.
20073. +ltho%"h al"ae
an) plants )#!!e$ #n (any
4ays6 the !%n)a(ental
p$oesses6 s%h as
photosynthes#s6 that (a.e
the( so )#st#n"%#she)
a(on" Ea$thEs o$"an#s(s
an) /al%-a'le as $ops6 a$e the sa(e.
<e$ta#n st$a#ns o! al"ae ha/e 'een %se) !o$
anth$opo-"en# p%$poses !o$ tho%san)s o!
yea$s6 #nl%)#n" as s%p-ple(ents an)
n%t$ae%t#als 2B#ple an) O$nelas 2000 3 an)
#n the !e$t#l#,at#on o! $#e pa))#es 2T%n" an)
5hen 1985 3. +s ea$ly as the 1940s6 othe$
st$a#ns 4e$e #)ent#!#e) as poss#'le !%el so%$es
2?o$o4#t,.a 201=a 3 'ea%se o! the#$ a'#l#ty to
p$o)%e !%el o$ !%el p$e%$so$ (ole%les.
8a$"e-sale p$o)%t#on
an) %lt#/at#on syste(s6
#nl%)#n" pho-
to'#o$eato$s an) o%t)oo$
open pon)s6 4e$e
)e/elope) #n the ea$ly
1950s #n the ;.5.6
:e$(any6 0apan6 an) the
@ethe$lan)s 2?o$o4#t,.a
201='D Ta(#ya 1957 3.
?y the onset o! the ;.5.
Depa$t(ent o! Ene$"yEs
2DOE3 a9%at# spe#es
p$o"$a( 2+5P3 #n the
;.5. #n 19806 /a$#o%s
spe#es o! (#$oal"ae an)
yano'ate$#a 4e$e 'e#n"
p$o)%e) an) !a$(e) on
o((e$#al sales
a$o%n) the 4o$l)6 an)
ha)
123
Photosynth
Res
Ta'le 1 <o((e$#al p$o)%ts
P$o)%t ;se
E7a(ple
so%$e Re!e$ene
!$o( al"ae
'-<a$otene 5%pple(ent D%nal#ella 8a(e$s et al. 2 2008 3
+sta7anth#n 5%pple(ent
Fae(atoo%
s 8o$en, an) <yse4s.# 2 2000 3
Whole-ell
n%t$ae%t#als 5%pple(ent 5p#$%l#na Bhan et al. 2 2005 3
<hlo$ella :oG$s et al. 2 2010 3
+9%a%lt%$e !ee) +n#(al !ee) Tet$asel(#s :la)%e an) *a7ey 2 1994 3
Isoh$ys#s :la)%e an) *a7ey 2 1994 3
Poly%nsat%$ate) !atty 5%pple(ent
<$yptheo)#n#%
( 0#an" et al. 2 1999 3
a#)s 2P;1+s3 5h#,ohyt$#%( 5polao$e et al. 2 200A 3
Phyoe$yth$#n ?#otehnolo"y Re) al"ae P%l, an) :$oss 2 2004 3
1%el (ole%les Ene$"y ?ot$yoo%s
+sho..%(a$ an) Ren"asa(y 2
20123
5ene)es(%s *an)al an) *all#. 2 2009 3
@eohlo$#s :o%/e#a et al. 2 2009 3
+nt#ane$ )$%"s
Pha$(a%et#a
ls 5y(ploa <oates et al. 2 201= 3
'een !o$ o/e$ 20 yea$s6 (ostly !o$ the health !oo)
an) n%t$#t#onal s%pple(ent #n)%st$#es 2?o$o4#t,.a
201='3.
*#$oal"ae ha/e e/ol/e) to 'e p$at#ally
%'#9%#to%s th$o%"ho%t the "lo'e6 an) the#$ /a$#e)
)#st$#'%t#ons an) e/ol%t#ona$y h#sto$#es 21ehl#n" et
al. 2007 3 a$e $e!lete) #n e7t$e(ely )#/e$se
(eta'ol# apa'#l#t#es 'et4een spe#es 2+n)e$sen
201=3. These )#/e$se (eta'ol#s(s p$o)%e a
(y$#a) o! o(po%n)s 4#th anth$opo"en#
$ele/ane #nl%)#n" n%t$ae%t#als6 s%h as the
a$oteno#)s p$o)%e) 'y D%nal#ella an)
Fae(atoo%s 2?o$o4#t,.a 201=a 6 ' 36 the
poly%nsat%$ate) !atty a#)s 2P;1+s3 p$o)%e) 'y
/a$-#o%s spe#es 2Ratle)"e 2004 36 an) the h#"h-
/al%e p$ote#ns an) a$'ohy)$ates a/a#la'le #n
4hole-ell s%pple(ents o! 5p#$%l#na an) <hlo$ella
2:oG$s et al. 2010 D Bhan et al. 2005 3. 5o(e
(#$oal"ae p$o)%e o(po%n)s o! '#oteh-
nolo"#al #nte$est #nl%)#n" !l%o$esent
o(po%n)s6 s%h as phyoe$yth$#n6 an) (any
p$o)%e #sop$eno#) (ole%les that an 'e %se) #n
!oo) an) o/e$-the-o%nte$ p$o)%ts 2+n)e$sen
201=3.
*#$oal"ae ha/e also 'een #)ent#!#e) as
att$at#/e so%$es o! '#o!%el 'ea%se )#!!e$ent
spe#es an p$o)%e a /a$#ety o! !%el p$o)%ts.
Va$#o%s (#$oal"al spe#es ha/e the a'#l#ty to
p$o)%e la$"e 9%ant#t#es o! l#p#) 4h#le se9%este$#n"
<O
2
6 pa$t#%la$ly ne%t$al l#p#)s #n the !o$( o!
t$#ayl"lye$ol 2T+:36 4h#h an 'e on/e$te) to
!atty a#) (ethyl este$s 21+*Es36 the (a#n
o(ponents o!
'#o)#esel 2Fossa#n et
al. 2008 36 th$o%"h
t$ans-este$#!#at#on6 o$
$e!#ne) #nto othe$ !%el
onst#t%ents 2P#en.os
an) Da$,#ns 2009 3.
Total l#p#)s an) othe$
'#o(ass onst#t%ents
an 'e on/e$te) #nto
$%)e o#l alte$nat#/es
th$o%"h the$(o-
he(#al p$oesses
s%h as hy)$othe$(al
l#9%e!at#on 2?a$$e#$o
et al. 201= 3.
*#$oal"al
a$'ohy)$ates an 'e
!e$(ente) #nto ethanol6
an) so(e spe#es an
p$o)%e '#ohy)$o"en
2Ra)a.o/#ts et al.
20103. In a))#t#on to
the#$ )#/e$s#ty o!
p$o)%ts6 (#$oal"ae
a$e att$at#/e as !%el
so%$es 'ea%se (any
spe#es "$o4 $elat#/ely
!ast
o(pa$e) to te$$est$#al
plants an) an 'e "$o4n
on '$a.#sh o$ sal#ne
4ate$6 th%s a/o#)#n" the
%se o! %ns%sta#na'le
9%ant#t#es o! !$esh4ate$6
an #n$eas#n"ly l#(#te)
$eso%$e 2D#s(%.es et
al. 2008 3. Ta'le 1
p$o/#)es an o/e$/#e4 o!
so(e o((e$#al al"al
p$o)%ts an) potent#al
so%$es.
+l"a%lt%$e6 o$ the
!a$(#n" o! al"ae
25a/a"e 2011 36 (e$"es
the $e9%#$e(ents o!
t$a)#t#onal te$$est$#al
plant a"$#%lt%$e s%h
as s%nl#"ht6 4ate$6
<O
2
6 n%t$#ent #np%ts6
an) ha$/est#n" syste(s
4#th a))#t#onal
a9%a%lt%$e
$e9%#$e(ents s%h as
sel!-onta#ne) a9%at#
syste(s6 4ate$ 9%al#ty6
an) 4aste
)#sposal/$eyl#n" 21#".
13. ?ea%se o! the#$
apa'#l#ty to p$o)%e
o((o)#t#es that span
(%lt#ple (a$.ets6
#nl%)#n" those o!
health !oo)6
n%t$ae%t#als6 pha$(ae%t#als6 an#(al !ee)6
he(#als an) ene$"y6 al"ae a$e %n#9%ely /e$sat#le
$ops 2Rosen'e$" et al. 2008 3. These )#/e$se
(eta'ol# apa'#l#t#es a$e )%e6 #n pa$t6 to the
)#/e$s#ty o! st$a#ns !o%n) 4#th#n the al"al l#nea"e.
+l"ae st$a#ns "$o4n !o$ !oo) p%$poses6 s%h as
5p#$%l#na6 ha/e a sta$.ly )#!!e$ent (eta'ol# p$o!#le
!$o( st$a#ns "$o4n !o$ ene$"y6 s%h as
5ene)es(%s. The )#/e$s#ty o! the#$ en) p$o)%ts6
an) the#$ %lt#/at#on %s#n" 'oth a"$#%lt%$e an)
a9%a%lt%$e p$at#es (a.e al"ae %n#9%e a(on"
othe$ a"$#%lt%$al p$o)%ts.
Desp#te s#"n#!#ant o/e$lap 4#th 'oth t$a)#t#onal
a"$#-%lt%$e an) a9%a%lt%$e 24h#h <on"$ess has
)e!#ne) as a"$#%lt%$e6
#nl%)#n" that o! a9%at#
plants3 21oo) an)
+"$#%lt%$e +t o! 19776
197736 al"a%lt%$e has
not yet 'een a!!o$)e) an
o!!##al pos#t#on 4#th#n
T#tle 7 o! the ;.5. <o)e
2;5<3 !o$ +"$#%lt%$e.
The$e a$e %$$ently a
n%('e$ o! othe$ $ops
that sha$e
o((onal#t#es 4#th
al"ae #n the#$ %lt#/at#on
p$at#es o$ )#/e$s#ty o!
en)-%se (a$.ets6 '%t
these ha/e all 'een
)es#"nate) a plae
4#th#n T#tle 7. 1o$
e7a(ple6 the
o((e$#al %lt#/at#on
o! a9%at# plants6 s%h
as sea"$ass6 #s el#"#'le
!o$ a )#/e$se a$$ay o!
a"$#%lt%$al p$o"$a(s.
5#(#la$ly6 the !a$(#n" o!
te$$est$#al $ops !o$
$ene4a'le ene$"y6 4h#h
sha$es the sa(e en)
(a$.et an)
123
Photosynth Res
1#". 1 +l"a%lt%$e #n the ;.5.
a b
+l"a%lt%$e an ta.e plae #n
lose) photo'#o$eato$s6 l#.e
those o! +l"enol #n 1lo$#)a
2a3 an) 5ol#7 ?#osyste(s #n
<olo$a)o 2'36 o$ #n open pon)s
l#.e those o! 5apph#$e Ene$"y6
In. #n @e4 *e7#o 23. 8#.e
a"$#%lt%$e6 al"ae %lt#/at#on
$e9%#$es "$o4th as 4ell as
ha$/est#n" #n!$ast$%t%$e6 s%h
as that o! 5apph#$e Ene$"y In.
2)3
c d
p%$pose as (any al"al-!a$(#n"
ope$at#ons6 'ene!#ts !$o( #ts
)e!#n#t#on as a"$#%lt%$e.
1%n)#n" !o$ $esea$h an)
)e/elop(ent o! al"al '#o(ass
%lt#/at#on has #n$ease) o/e$
the last )ea)e6 an) has le) to
the e(e$"ene o! $esea$h
p$o"$a(s6 p$#/ate p$oHets6
)e(onst$at#on- an)
o((e$#al-sale !a#l#t#es
a$oss the ;.5. 21#". 2 3. The
#n$ease #s p$#(a$#ly )%e to the
"$o4th o! the al"al '#o!%el
#n)%st$y #n $esponse to the
)e(an) !o$ alte$nat#/e !%el
so%$es )$#/en 'y the
$ene4a'le !%el stan)a$)s 2R153
2Tyne$ 201= 3. Wh#le the %se o!
al"ae as !%nt#onal !oo) o$ !ee)
#n"$e)#ents #s also on the $#se
2I-'anIe, an) <#!%entes 201= 36
the$e a$e %$$ently !e4 !e)e$al
p$o"$a( $eso%$es !o%se) #n
th#s a$ea. The p$o)%t#on o!
al"ae !o$ any en) p$o)%t #s a
t4o-phase p$oess #n/ol/#n"
the !a$(#n" an) %lt#/at#on o!
al"al '#o(ass !ollo4e) 'y
p$oess#n" o! the ha$/este)
'#o(ass. The a'#l#ty o! the
al"al '#o(ass #n)%st$y to
aess !e)e$al
p$o"$a(s that s%ppo$t
the a"$#%lt%$al phase
#s #(pe$at#/e !o$
!%t%$e "$o4th. Th#s
$epo$t analy,es the
plae o! al"ae #n the
%$$ent a"$#%lt%$al
pol#y an) !%n)#n"
lan)sape6 an) the
oppo$t%n#t#es an)
p#t!alls that e7#st !o$
al"ae 4#th#n th#s
pol#y !$a(e4o$..
+"$#%lt%$al
p$o"$a(s
<on"$ess has
le"#slate) a n%('e$ o!
$ene4a'le ene$"y
p$o"$a(s that an 'e
appl#e) to al"ae s%h
as the ?#oen-e$"y
P$o"$a( !o$ +)/ane)
?#o!%els6 the R%$al
Ene$"y !o$ +(e$#a
P$o"$a(6 the ?#o(ass
Resea$h an)
De/elop(ent
In#t#at#/e an) /a$#o%s
"$ants an) loans
esta'l#she) #n the
2008 1a$( ?#ll #n
set#on 900= o! the
;5< 21oo)6 <on-
se$/at#on6 J Ene$"y
+t o! 20086 2008 3.
These p$o"$a(s6
ho4e/e$6 !o%s on
$esea$h an)
)e/elop(ent o! al"ae
!o$ !%els at s(alle$
sales. Wh#le th#s
#n#t#al #n/est(ent #n
$esea$h J
)e/elop(ent 2RJD3
#s essent#al to '%#l)
.no4le)"e6 e7pe$t#se6
an) tehnolo"y
a$o%n) al"ae6 the
#n)%st$y #s no4
ente$#n" the !o$(at#/e
sta"e o! la$"e-sale
o((e$#al#,at#on6
4h#h $e9%#$es
'$oa)e$ oo$)#nat#on
a(on" !e)e$al
a"en#es an) s%ppo$t
#n!$ast$%t%$e to "a#n
p$ope$ al#"n(ent at
the !e)e$al an) state
le/el $e9%#$e) !o$ a
s%ess!%l #n)%st$y.
?#o(ass $op ass#stane
p$o"$a(
The ?#o(ass <$op +ss#stane
P$o"$a( 2?<+P3 4as
esta'l#she) #n the 2008 !a$(
'#ll 21oo) J <onse$/at#on +t
o! 20086 2008 3 to !#nan#ally
ass#st !a$(e$s 4#sh#n" to
esta'l#sh6 p$o)%e6 an) )el#/e$
'#o(ass !ee)sto.s. ?<+PEs
p%$pose #s to p$o(ote !a$(#n"
o! '#oene$"y $ops. The
p$o"$a( p$o/#)es e#the$ one-
t#(e esta'l#sh(ent
pay-(ents6 ann%al
pay(ents6 o$
(ath#n" pay(ents to
help 4#th ha$/est6
sto$a"e6 an)
t$anspo$tat#on o!
'#o(ass. P$o-posals
!o$ ?<+P !%n)#n" a$e
s%'(#tte) to the 15+
an) an o(e !$o(
e#the$ p$o)%e$s o$
on/e$s#on !a#l#t#es
25h-nep! 2011 3.
Wh#le (any
t$a)#t#onal '#o!%el
$ops a$e %$-$ently
el#"#'le !o$ ?<+P
!%n)#n"6 s%h as
s4#th"$ass an) (ost
non-!oo) '#o(ass6 the
2008 !a$( '#ll
spe#!#ally e7l%)e)
al"ae !$o(
pa$t##pat#on #n the
(ath#n" pay(ent
s#)e o! ?<+P '%t
9%al#!#es al"ae !o$
esta'l#sh(ent
123
Photosynth Res
1#". 2 +l"ae p$oHets #n the ;.5. +l"al '#o(ass p$oHets e7#st #n al(ost e/e$y state #n the ;.5. ?l%e p#ns )enote a $esea$h #nst#t%t#on6 "$een
)enote a p$#/ate p$oHet o$ o(pany
pay(ents th$o%"h ?<+P
21oo) J <onse$/at#on +t
o! 20086 2008 3.
5%ppo$t p$o"$a(s
<on"$ess has app$op$#ate) n%(e$o%s !e)e$al a"en#es6 s%h as
the ;5D+ an) DOE6 !%n)s an) a%tho$#,at#on to #(ple(ent
p$o"$a(s that a#) an) s%ppo$t )e/elop(ent o! a"$#%lt%$e an)
a9%a%lt%$e $eso%$es 2Ta'le 2 3. 5#ne the passa"e o! the
o$#"#nal +"$#%lt%$al +)H%st(ent +t o! 19==6 eah s%'se9%ent
!a$( '#ll has e/ol/e) to a))$ess $#s#n" $ele/ant #ss%es #n
a"$#%lt%$e. Th#s !$e9%ently #n/ol/es )$a!t#n" ne4 p$o"$a(s o$
e7pan)#n" e7#st#n" p$o"$a(s to the ne4 )e/elop#n"
tehnolo"#es. The 1977 !a$( '#ll 21oo) J +"$#%lt%$e +t o!
19776 1977 3 e7pan)e) the )e!#n#t#on o! a"$#%lt%$e to #nl%)e
a9%a%lt%$e6 th%s sp%$$#n" the )e/elop(ent o! #n)%st$y #n the
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