!!!"#$%&'(')*+,+"!'-.

/-011"2'3
u


!"#$#%&'$"(%)%* ,"-$ )% )$.
56'7'1*87601&1 ls a process whereby llghL energy ls converLed Lo chemlcal energy whlch can be used Lo synLhesls large,
organlc molecules from smaller, lnorganlc subsLraLes. lanLs and oLher organlsms whlch are phoLosynLheLlc (for example,
bacLerla) are known as #97'7-'/61, LhaL ls, Lhey make organlc compounds from small lnorganlc precursors. More
speclflcally, Lhose auLoLrophs whlch undergo phoLosynLhesls are known as !"#$#%&$#$'#!"(. Anlmals, and many oLher
organlsms, cannoL synLheslse Lhelr own food, buL lnsLead dlgesL organlc molecules. 1hese are known as 6070-'7-'/61.
carbon dloxlde + waLer glucose + oxygen
6CC
2
+ 6P
2
C C
6
P
12
C
6
+ 6C
2

hoLosynLhesls ls consldered Lo be Lhe mosL fundamenLal blochemlcal process, as all %)'#*)( (organlsms whlch resplre
aeroblcally) requlre Lhe producLs of phoLosynLhesls Lo undergo #0-'%&2 -01/&-#7&'8.
/"(0( 1#(% 2"#$#%&'$"(%)% $-3( 24-5(.
hoLosynLhesls ls a process whlch occurs ln Lwo sLages, boLh enLlrely lnslde Lhe 26('-'/(#17. A chloroplasL ls an organelle
wlLhln phoLosynLheLlc cells. 1he llsL below ouLllnes some of Lhe adapLaLlons chloroplasLs have whlch help Lhem achleve
efflclenL raLes of phoLosynLhesls, and Lhe explanaLlons wlll become clear Lo you as you progress Lhrough Lhe Loplc:
Lhe granal membranes provlde a large surface area for Lhe aLLachmenL of Lhe phoLosynLheLlc plgmenLs (chlorophylls
and caroLenolds), elecLron carrlers and enzymes for Lhe llghL-dependenL reacLlons
a neLwork of proLelns ln Lhe grana hold Lhe plgmenLs ln a very preclse manner LhaL forms Lhe phoLosysLems allowlng
for maxlmum absorpLlon of llghL
Lhe granal membranes have many A1 synLhase enzymes aLLached Lo Lhem whlch, vla 2603&'13'1&1 (see 4.4 LlecLron
LransporL chaln) help Lo manufacLure A1 molecules
Lhe fluld of Lhe sLroma holds all of Lhe enzymes needed Lo carry ouL Lhe llghL-lndependenL reacLlons
Lhe sLroma fluld surrounds Lhe grana, and so Lhe producLs of Lhe llghL-dependenL reacLlons can dlrecLly and readlly
pass lnLo Lhe sLroma for Lhe llghL-lndependenL reacLlons
chloroplasLs conLaln boLh unA and rlbosomes so Lhey can qulckly and easlly manufacLure phoLosynLheLlc proLelns
6$075$70( #8 - 5"4#0#24-%$
ChloroplasLs vary, buL mosL are dlsc-shaped and approxlmaLely 2-10mlong.Eachhasadoublemembrane(calledan
08:0('/0), conslsLlng of Lhe lnner membrane and ouLer membrane.
ChloroplasLs conLaln sLacks of
flaLLened membrane comparL-
menLs. Lach sLack ls called a
)-#893 (plural: ,'%-%) and each
comparLmenL ls a 76*(#;'&..
Small, Lhln membranal
exLenslons connecL dlfferenL
grana, called &870-)-#8#(
(#30((#0. 1he fluld surroundlng
Lhe grana ls called 17-'3#.
SLarch gralns can also be found
ln Lhe sLroma maLrlx, as well as
<=> and -&%'1'301, whlch can
be used Lo make proLelns.
1he Lwo sLages of phoLosynLhesls are Lhe ./,"$01)!)-1)-$ ')%2$/#-( (thefirstorlightstage)andthe./,"$0/-1)!)-1)-$
')%2$/#-( (thesecondordarkstage).1he flrsL sLage Lakes place ln Lhe ,'%-%, and Lhe second sLage ln Lhe ($'#3% 4.&/1 of
Lhe chloroplasL.
?6('-'/(#171 #8. /6'7'1*87601&1
Why organlsms undergo phoLosynLhesls and how chloroplasLs are adapLed for Lhelr funcLlon
@"+
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
"
" "
"
"
"
"
granum
slngle
Lhylakold
rlbosomes
lnLergranal lamella
sLarch graln
sLroma
unA


!!!"#$%&'(')*+,+"!'-./-011"2'3

5*#6)7 $'%-(3/((/#- ).)2$'#- 3/2'#(2#!/2 /3%,) 89:;< #4 % !.%-$ 2).. 2".#'#!.%($ 8=>?@???<
!"#$#%&'$"($)5 2)9:('$% -'1 2"#$#%&%$(:%
Lmbedded ln Lhe Lhylakold membrane among grana are coloured compounds whlch absorb llghL of a shorL range of
wavelengLhs and reflecL llghL of oLher wavelengLhs. 1hese are called /6'7'1*87607&2 /&)30871. A phoLosynLheLlc plgmenL
can absorb some llghL energy of speclflc wavelengLhs. varlous phoLosynLheLlc plgmenLs are arranged lnLo small sLrucLures
called /6'7'1*17031 ln Lhe granal membrane.
1he phoLosysLem shown ln Lhe granal membrane ln Lhe
dlagram ls made up of four phoLosynLheLlc plgmenLs. 1he
plgmenL closesL Lo Lhe ouLer membrane ls 26('-'/6*(( %,
and Lhe Lwo ln Lhe lnLermembrane space are A#876'/6*((
and 2#-'7080. 1hese Lhree plgmenLs are called Lhe
102'8.#-* /&)30871 (or %22)((#'A !/,3)-$().
1he plgmenL on Lhe lnner membrane Lo Lhe organelle ls
called 26('-'/6*(( #, whlch ls Lhe /-&3#-* /&)3087. As llghL
hlLs Lhe secondary plgmenLs whlch absorb Lhe llghL, Lhey
geL exclLed and a palr of elecLrons are passed Lhrough Lhe
plgmenLs and Lhrough Lo Lhe prlmary plgmenL.
1he Lable below shows Lhe dlfferenL phoLosynLheLlc plgmenLs and Lhelr feaLures:
5&)3087 ?'('9- 50#; #%1'-/7&'8 !#:0(08)76 B83C D9827&'8 &8 /6'7'1*87601&1
Chlorophyll a ?ellow-green 430, 662
Absorbs red and blue-purple llghL
Chlorophyll b 8lue-green 433, 642
CaroLene Crange 430
Absorb purple llghL, proLecL chlorophylls
from damage from llghL and oxygen
xanLhophyll ?ellow 430-470

A calorlmeLer ls used Lo measure Lhe absorpLlon of dlfferenL wavelengLhs of llghL by dlfferenL plgmenLs. 1he resulLs of Lhe
calorlmeLer LesL can be ploLLed onLo a graph called Lhe #%1'-/7&'8 1/027-93. Comblnlng Lhe absorpLlon specLra of all
phoLosynLheLlc plgmenLs glves Lhe #27&'8 1/027-93 of overall phoLosynLhesls.
E&)67 E&)67
chlorophyll b
xanLhophyll
caroLene
chlorophyll a

!!!"#$%&'(')*+,+"!'-./-011"2'3









1he graph Lo Lhe lefL above shows Lhe absorpLlon specLra for phoLosynLheLlc plgmenLs, and Lhe dlagram Lo Lhe rlghL
above shows Lhe acLlon specLrum for phoLosynLhesls.
;"0#:-$#90-2")5 %(2-0-$)#' #8 2)9:('$%
A 26-'3#7')-#3 can be used Lo separaLe Lhe dlfferenL phoLosynLheLlc plgmenLs ln a phoLosysLem. ChromaLography ls a
processwherebyamixtureofmaterialsareseparatedbyallowingdifferentparticlestomove(inwhatisknownasthe
3#*/.) !"%()) along a chromaLographlc sLrlp and flx along a cerLaln place (ln whaL ls known as Lhe ($%$/#-%'A !"%()), Lhus
parLlLlonlng Lhe dlfferenL parLlcles.
uslng chromaLography separaLes Lhe plgmenLs lnLo colour order,
accordlng Lo Lhe vlslble llghL specLrum. We can also use Lhe chromaLogram
Lo calculaLe Lhe FG :#(90 (relaLlve fronL) of varlous plgmenLs. 1he 8f value
ls calculaLed by dlvldlng Lhe dlsLance beLween Lhe orlgln and plgmenL fronL
by Lhe dlsLance beLween Lhe orlgln and Lhe solvenL fronL (l.e. where Lhe
solvenL sLops movlng along Lhe chromaLography paper).
lor example, Lhe plgmenL xanLhophyll may have a relaLlve fronL of around
0.4 and chlorophyll b around 0.1. lL ls llkely LhaL caroLene wlll have an 8f
value of beLween 0.9 and 1. 1he caroLenolds (caroLene and xanLhophyll)
wlll Lravel furLher along Lhe paper, and Lherefore have hlgher 8f values
thanthechlorophylls,whichenterthestationaryphaseearlieron.
relaLlve fronL = (orlgln Lo plgmenL fronL) - (orlgln Lo solvenL fronL)
chlorophyll b
caroLenolds
chlorophyll a

a
g
e

a
b
s
o
r
p
L
l
o
n

400 300 600 700
wavelengLh of llghL (nm)
r
a
L
e

o
f

p
h
o
L
o
s
y
n
L
h
e
s
l
s

400 300 600 700
wavelengLh of llghL (nm)
orlgln
plgmenL
fronL
solvenL
fronL
?LLLCW-C8AnCL
(caroLene)
?LLLCW
(xanLhophyll)
?LLLCW-C8LLn
(chlorophyll a)
C8LLn
(chlorophyll b)

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u


1he flrsL sLage, or seL of reacLlons, lnvolved ln phoLosynLhesls ls called Lhe ||ght-dependent sLage. 1he llghL-dependenL
reacLlons occur ln Lhe thy|ako|d membranes lnslde chloroplasLs. lL ls Lhe phoLosysLems (whlch are made up of many
phoLosynLheLlc plgmenLs) embedded ln Lhese membranes whlch are of lmporLance. lor one seL of llghL-dependenL
reacLlons Lo happen, Lwo phoLosysLems are used: photosystem I (Sl), whlch ls found malnly on Lhe lnLergranal lamellae,
and photosystem II (Sll), whlch ls found almosL excluslvely on Lhe granal lamellae. 1he llghL-dependenL sLage ls focused
on converLlng llghL energy (Lrapped by plgmenLs ln Lhe phoLosysLems) lnLo chemlcal energy.
"#$%$&'()( $* +,%-.
1he phoLosysLem whlch ls lnvolved ln Lhe flrsL parL of Lhls sLage ls acLually phoLosysLem ll, as phoLosysLem l ls lnvolved
laLer on. Sll has an enzyme whlch, ln Lhe presence of llghL, spllLs a molecule of waLer lnLo proLons, elecLrons and oxygen.
2P
2
C 4P
+
+ 4e
-
+ C
2

lL ls Lhe proLons and elecLrons produced whlch are of lmporLance here. 1he oxygen ls acLually a wasLe producL for Lhls
reacLlon. Powever, Lhls photo|ys|s (Lhe spllLLlng of a molecule uslng photons, or llghL parLlcles) of waLer from Lhe llghL-
dependenL reacLlons ls seen as one of Lhe mosL lmporLanL reacLlons ln llfe: Lhe oxygen produced ls Lhe source of almosL
all of Lhe oxygen ln Lhe aLmosphere, whlch many organlsms requlre Lo resplre, and Lherefore llve. AlLhough a wasLe
producL ln Lhls reacLlon, planLs use some of Lhe oxygen produced Lo aeroblcally resplre, buL mosL of lL leaves Lhe leaves
Lhrough Lhe stomata (pores ln Lhe leaf surface where gaseous exchange may Lake place ln planLs).
/$.0,%)$1 $* 23"4 5#-0)$(0$()(
1he proLons, or hydrogen lons, have Lwo purposes. llrsLly, Lhey are used Lo generaLe A1 molecules by chem|osmos|s
(see 4.4 LlecLron LransporL chaln for more deLalls). ln summary, proLons dlffuse across Lhe Lhylakold membrane Lhrough
channel proLelns whlch are assoclaLed wlLh Lhe enzyme A1 synthase, drlvlng Lhe roLaLlon of parL of Lhe A1 synLhase
molecule, whlch comblnes a molecule of Au (adenoslne dlphosphaLe) and an lnorganlc phosphaLe Lo form A1.
1he proLons Lhen are accepLed by Lhe coenzyme NAD (nlcoLlnamlde adenlne dlnucleoLlde phosphaLe), alongslde
elecLrons, Lo form reduced nAu, or NADn (also someLlmes wrlLLen nAuP
2
). 8educed nAu ls Lhen used ln Lhe second
sLage of phoLosynLhesls, Lhe !"#$%&"'()*)'()'% ,)-.%"/'0.












1he dlagram above shows Lhe phoLosysLems l and ll and Lhe enzyme A1 synLhase embedded ln a Lhylakold membrane.
1he blue area shows Lhe phoLolysls of waLer, and Lhe paLhways of Lhe elecLrons and proLons produced. 1he oxygen may
be used for aeroblc resplraLlon, buL wlll usually leave Lhe leaf Lhrough Lhe sLomaLa.
L|ght-dependent react|ons
1he llghL sLage of phoLosynLhesls: phoLolysls of waLer and phoLophosphorylaLlon
3.2
P
2
C
xC
2

2P
+

2e
-

SII SI
2e
-

2e
-

2P
+

2P
+

+2P
+

nAu nAuP
Au + l
A1
llghL llghL
thy|ako|d space
stroma
A1 synLhase
elecLron
carrlers

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3#- 678,%#+,'
1he serles of reacLlons lnvolved ln Lhe llghL-dependenL sLage are known collecLlvely as Lhe Z-paLhway because of Lhe
movemenL of elecLrons across Lhe phoLosysLems and Lhylakold membrane.












1 ln Lhe presence of llghL, a molecule of waLer ls spllL uslng an enzyme found wlLhln phoLosysLem ll, wlLh a peak
absorpLlon of llghL wavelengLh 680nm (wrlLLen
680
), lnLo an aLom of oxygen, as well as Lwo elecLrons and Lwo
proLons. 1he oxygen ls a by-producL whlch ls noL used ln Lhls reacLlon
2 1he Lwo hydrogen lons (proLons) remaln lnslde Lhe Lhylakold space, buL Lhe elecLrons are accepLed by phoLosysLem ll
when llghL ls presenL, as Lhe phoLons of llghL exclLe Lhe elecLrons, so LhaL Lhey are Lhen accepLed by an e|ectron
acceptor and passed along varlous e|ectron carr|ers (or cytochromes)
3 1he movemenL of elecLrons along Lhe Lhylakold membrane beLween dlfferenL cyLochromes releases energy, whlch
pumps proLons across Lhe membrane from Lhe sLroma lnLo Lhe Lhylakold lnner space
4 1hls creaLes a concenLraLlon gradlenL of proLons over Lhe membrane as Lhey begln Lo accumulaLe, and so proLons
begln Lo dlffuse back Lhrough Lo Lhe sLroma Lhrough channel proLelns, whlch are assoclaLed wlLh Lhe enzyme A1
synLhase. As proLelns flow Lhrough Lhe channel, Lhe enzyme ls acLlvaLed, as Lhe LransporL of Lhe proLon drlves Lhe
roLaLlon of Lhe enzyme so LhaL one Au molecule and one phosphaLe group form a molecule of A1 Lhls ls
photophosphory|at|on (producLlon of A1 uslng llghL energy)
1here are Lwo Lypes of phoLophosphorylaLlon whlch Lake place durlng Lhe llghL-dependenL reacLlons. Cne Lype lnvolves
only phoLosysLem l (whlch has a peak absorpLlon of 700, so ls also called
700
), and Lhe oLher boLh Lhe phoLosysLems.
!"#$%&%'(% *+","*+"-*+".&'/,("#
LssenLlally, Lhe enLlre llghL-dependenL sLage ls Lhe process of non-cyc||c photophosphory|at|on. 1hls lnvolves Lhe flow of
elecLrons from Lhe phoLolysed waLer (Lhe orlglnal elecLron donor) Lhrough Lhe phoLosysLems and cyLochromes Lo Lhe
reducLlon of nAu. non-cycllc phoLophosphorylaLlon lnvolves Lhe sLeps 1-4 as seen above. ln Lhe dlagram above of Lhe Z-
paLhway, non-cycllc phoLophosphorylaLlon can be seen produclng A1 on Lhe dlagonal sLroke of Lhe Z-shape Lravelllng
Lowards phoLosysLem l.
1he elecLrons losL by phoLosysLem l are replaced by Lhe elecLrons from oxldlsed phoLosysLem ll, and Lhe elecLrons
donaLed from phoLolysed waLer replace Lhose Lhen losL by Sll. 1hls flow of elecLrons ls whaL makes Lhe Z-paLhway.
0&%'(% *+","*+"-*+".&'/,("#
AnoLher Lype, cyc||c photophosphory|at|on also occurs, buL only aL phoLosysLem l. LlecLrons are losL from Sl as phoLons
of llghL exclLe Lhem, and passed around elecLron carrlers and Lhen back Lo Lhe chlorophyll of Sl where Lhey flrsL came
from, ln a cycle. 1hls movemenL of elecLrons does generaLe small amounLs of A1, buL does noL lnvolve phoLolysls of
waLer or reduce any nAu. 1he A1 produced may be used for oLher funcLlons where energy ls requlred.
2e
-
elecLron
accepLor

2e
-
elecLron
accepLor

Au + l
A1
A1
2e
-
nAu reducLase

n
2
C
2e
-
2P
+
2n
+
nAu

nAuP

SII
!
"#$
SI
!
&$$
key:

llghL

cycllc phosphorylaLlon

non-cycllc phosphorylaLlon


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1he Lable below compares Lhe dlfferenL Lypes of phoLophosphorylaLlon Lo provlde a summary:
non-cyc||c photophosphory|at|on cyc||c photophosphory|at|on
Whlch phoLosysLems are lnvolved? Sl and Sll Sl only
uoes Lhe reacLlon lnvolve phoLolysls of waLer? yes no
Whlch molecule ls Lhe elecLron donor? waLer chlorophyll a (
700
) ln Sl
Whlch molecule ls Lhe flnal elecLron accepLor? nAu chlorophyll a (
700
) ln Sl
WhaL are Lhe producLs? reduced nAu, A1 and oxygen A1 only

".$9:5%( $* %#- &);#%79-8-19-1% (%,;-
8oLh cycllc and non-cycllc phoLophosphorylaLlon drlve Lhe synLhesls of A1, so we know Lhls ls one producL of Lhe llghL-
dependenL sLage. 1he oLher useful producL for phoLosynLhesls ls nAuP (reduced nAu), whlch ls an lmporLanL molecule
for Lhe followlng sLage (see 3.3 LlghL-lndependenL reacLlons). Cxygen ls a by-producL whlch leaves Lhe chloroplasL and Lhe
leaf Lhrough Lhe sLomaLa, alLhough some of Lhe oxygen produced may be used for aeroblc resplraLlon.

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u


1he ||ght-|ndependent sLage of phoLosynLhesls ls Lhe second and flnal seL of reacLlons. lL ls named so because Lhe
reacLlons lnvolved do !"# need llghL Lo occur, and so Lechnlcally can Lake place wlLhouL llghL. Powever, Lhe producLs (A1
and reduced nAu) of Lhe llghL-dependenL reacLlons are requlred, and so ln pracLlce, Lhese reacLlons wlll noL conLlnue for
very long wlLhouL llghL, as Lhe llghL-dependenL sLage wlll sLop produclng A1 and nAuP.
1he llghL-lndependenL reacLlons occur ln Lhe stroma of Lhe chloroplasL (fluld maLrlx surroundlng Lhe grana). lL ls also
called Lhe Ca|v|n cyc|e, named afLer Melvln Calvln eL al. (1946-33). WhllsL Lhe producLs of Lhe llghL-dependenL sLage (A1
and nAuP) are needed for Lhls sLage, carbon dloxlde ls also requlred. 1hls comes from Lhe aLmosphere. Carbon dloxlde
dlffuses Lhrough Lhe sLomaLa of leaves, and enLers leaf cells and Lhen Lhe chloroplasLs.
!"# %&'()* +,+'#
1he Calvln cycle cannoL Lake place wlLhouL carbon dloxlde. When lL enLers a leaf, %&''()&!* Lhrough sLomaLa (almosL
excluslvely on Lhe underslde of Lhe leaf), lL dlffuses Lhrough Llny alr spaces ln Lhe spongy mesophy|| layer, unLll lL reaches
Lhe pa||sade mesophyll layer. lL Lhen dlffuses Lhrough a Lhln cellulose cell wall, Lhe plasma membrane, Lhe cyLoplasm and
Lhrough Lhe chloroplasL envelope (double membrane), lnLo Lhe sLroma.


















1 1he carbon dloxlde comblnes wlLh a molecule called r|bu|ose b|phosphate (ku8), a flve-carbon compound and a
carbon dloxlde accepLor. 1hls reacLlon ls caLalysed by ku8|sCC (rlbulose blphosphaLe carboxylase-oxygenase), an
enzyme whlch adds a carboxyl group Lo Lhe 8u8 molecule
2 1hls comblnaLlon forms an unsLable slx-carbon compound, before spllLLlng lnLo Lhe producLs of Lhe reacLlon, whlch are
Lwo molecules of g|ycerate 3-phosphate (G), a Lhree-carbon compound, and aL Lhls sLage Lhe carbon dloxlde ls flxed
L|ght-|ndependent react|ons
SLage Lwo of phoLosynLhesls: Lhe Calvln cycle
3.3

8u8lsCC

x2 glyceraLe
3-phosphaLe
(3C)


(2C)
x2 Lrlose
phosphaLe
(3C)


(21)
rlbulose
blphosphaLe
(3C)


(8u8)
nAuP

nAu

CC
2

A1

Au + l

Au + l

A1

hexose sugars
(e.g. glucose)
glycerol
sLarch
cellulose
polysaccharldes
(e.g. glycogen)
faLLy aclds
amlno aclds
llplds
llplds

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3 1he C molecules are reduced (uslng Lwo hydrogen aLoms donaLed by reduced nAu from Lhe llghL-dependenL sLage)
and Lhen phosphory|ated (uslng an lnorganlc phosphaLe group from one molecule of A1, also from Lhe llghL-
dependenL sLage) Lo form anoLher Lhree-carbon compound called tr|ose phosphate (elLher 1 or GAL)
4 llve ouL of every slx molecules of 1/CAL are recycled vla phosphorylaLlon (uslng anoLher A1 molecule from Lhe
llghL-dependenL sLage of phoLosynLhesls) lnLo Lhree more molecules of 8u8 (flve-carbon compound), Lhe remalnlng
molecule of 1 produced (per molecule of 8u8) can be used Lo form oLher blologlcal molecules (see below)
-./0&1).* .2 3/.45+16
lalrly obvlously, Lhe Calvln cycle would be of no use lf all of Lhese producLs wenL around produclng no oLher producLs.
lorLunaLely, some molecules whlch are found ln Lhese reacLlons can be used Lo synLheslse oLher molecules whlch are of
use Lo organlsms:
one of every slx molecules of 1/CAL produced are !"# recycled as 8u8, buL are acLually used Lo make oLher
blologlcal molecules such as saccharldes Lwo molecules of 1 can comblne Lo form a hexose sugar, e.g. g|ucose and
such sLrucLures can polymerlse Lo form sLarches, sugars and cellulose
molecules of C can be used Lo form amlno aclds and faLLy aclds
1 can also be used Lo form glycerol, whlch can comblne wlLh Lhe faLLy aclds made by C Lo form varlous ||p|ds
758)6%9
1he enzyme 8u8lsCC (shorL for +&,(-"). ,&/0")/01#. 21+,"34-1).5"34*.!1).) ls Lhe mosL abundanL enzyme on earLh, as lL
makes approxlmaLely 30 of leaf proLeln. lL ls of upmosL lmporLance Lo llfe. AlLhough you can see LhaL Lhe Calvln cycle
uses 8u8lsCC Lo comblne a molecule of 8u8 and carbon dloxlde, as Lhe name of 8u8lsCC suggesLs, oxygen can also flL
lnLo Lhe enzyme complex. 1hls resulLs ln a reacLlon called photoresp|rat|on.
hoLoresplraLlon ls a process whereby oxygen comblnes wlLh 8u8 ln Lhe place of carbon dloxlde. 1hls lowers Lhe
efflclency of phoLosynLhesls ln planLs, as Lhls undoes a loL of Lhe work of Lhe process so far and also leads Lo Lhe
formaLlon of Lhe Loxlc hydrogen peroxlde.
When phoLoresplraLlon beglns Lo occur, l.e. when Lhere ls a hlgh concenLraLlon of oxygen ln Lhe aLmosphere ln relaLlon Lo
Lhe concenLraLlon of carbon dloxlde, Lhe reacLlons caLalysed by 8u8lsCC produce Lwo producLs: phosphog|ycerate (CA)
and phosphog|yco|ate (C). CA re-enLers Lhe Calvln cycle, belng converLed back lnLo 8u8, and so ls noL Loo
problemaLlc, buL Lhls does slow down raLe of phoLosynLhesls somewhaL. C, however, ls much more dlfflculL Lo geL rld
of, as lL has Lo leave Lhe chloroplasL and enLer mlLochondrla (among oLher organelles), undergolng a long serles of
reacLlons before Lhe end producLs can re-enLer Lhe Calvln cycle. 1hls obvlously lowers phoLosynLheLlc efflclency.
So why does 8u8lsCC have oxygenase funcLlons? uue Lo Lhe conLroverslal funcLlons of phoLoresplraLlon, Lhere ls no exacL
answer aL Lhe momenL, alLhough many sclenLlsLs have speculaLed. CerLaln Lheorles have suggesLed LhaL, slnce Lhe
comblnaLlon of 8u8 and oxygen leads Lo Lhe formaLlon of hydrogen peroxlde (P
2
C
2
), lL ls Lo ald homeosLaLlc mechanlsms
as Lhls subsLance makes a key conLrlbuLlon Lo cellular redox homeosLasls. lL has also been suggesLed LhaL 8u8lsCC allows
oxygen Lo comblne wlLh 8u8 Lo sLop free A1 and nAu from mlxlng wlLh oxygen and formlng rad|ca|s (compounds wlLh
unpalred elecLrons and lncompleLe ouLer energy levels) whlch can damage Lhe cells meLabollc funcLlon.

!!!"#$%&'(')*+,+"!'-./-011"2'3
u


!"#"$"%& ()*$+,-
When Lhere are a number of facLors whlch conLrlbuLe Lowards a process and conLrol lLs efflclency, Lhe facLor whlch ls
worklng aL Lhe leasL favourable level wlll be Lhe (&3&5&6) 7#25'- of Lhe process. ln phoLosynLhesls, Lhe llmlLlng facLor ls Lhe
quallLy or condlLlon of Lhe envlronmenL whlch ls leasL favourable and Lherefore acLs as a consLralnL on Lhe raLe of
phoLosynLhesls. lor example:
on a warm nlghL, Lhe llghL lnLenslLy ls Lhe llmlLlng facLor (Lhe LemperaLure ls sufflclenL buL Lhere ls no llghL)
on a cold, frosLy mornlng, Lhe LemperaLure ls Lhe llmlLlng facLor (Lhe LemperaLure lmpalrs Lhe phoLosynLheLlc raLe)
on a normal, sunny day Lhe carbon dloxlde concenLraLlon ln Lhe aLmosphere may be Lhe llmlLlng facLor
1he maxlma Lend Lo be around 30
o
C (LemperaLure), 0.2 - 0.3 carbon dloxlde concenLraLlon and brlghL sunllghL durlng
Lhe day.
!""#$% '" %#()#*+%,*# '- ).'%'/0-%.#/1/
WhllsL Lhe llghL-dependenL sLage of phoLosynLhesls ls noL very much aL all affecLed by Lhe LemperaLure of Lhe surroundlng
envlronmenL, Lhls ls cruclal durlng Lhe Calvln cycle, where Lhere ls fundamenLal lmporLance on enzyme acLlon. As you well
know, LemperaLures whlch flucLuaLe cause poor enzyme performance and Lhose whlch are Loo hlgh wlll denaLure
enzymes beyond repalr.
1he oLher problem, however, relaLed wlLh LemperaLure, ls LhaL an lncrease ln LemperaLure wlll cause a planL Lo lose more
waLer Lhrough Lhe sLomaLa. 1hls leads Lo a sLress response whereby Lhe sLomaLa close, whlch llmlLs Lhe avallablllLy of
carbon dloxlde as well as loslng waLer.
!""#$% '" $+*2'- 31'413# '- ).'%'/0-%.#/1/
Cenerally speaklng, an lncrease ln carbon dloxlde
avallablllLy means an lncrease ln Lhe raLe of
phoLosynLhesls. Cur aLmosphere has a CC
2

concenLraLlon of approxlmaLely 0.03 - 0.06 and Lhe
raLe of phoLosynLhesls wlll conLlnue Lo lncrease wlLh
carbon dloxlde concenLraLlon, buL only up unLll a
cerLaln polnL. AL around 0.3 concenLraLed, Lhe raLe
sLops lncreaslng because lL cannoL physlcally lncrease
any furLher as Lhe oLher condlLlons (llghL and
LemperaLure) become llmlLlng facLors Lhemselves.
1herefore, Lhe llmlLlng facLor graph looks as shown.
!""#$% '" 516.% 1-%#-/1%0 '- ).'%'/0-%.#/1/
When llghL lnLenslLy ls Lhe llmlLlng facLor on Lhe raLe of phoLosynLhesls, Lhe raLe ls dlrecLly proporLlonal Lo an lncrease or
decrease ln llghL lnLenslLy. As Lhe llghL lncreases, so does Lhe raLe of phoLosynLhesls. 1hls ls because more llghL means:
more sLomaLa are opened, so more carbon dloxlde can enLer
more llghL can be Lrapped by chlorophyll Lo exclLe elecLrons durlng Lhe llghL-dependenL sLage
more efflclenL phoLolysls of waLer durlng Lhe llghL-dependenL sLage
8&3&5&6) 7#25'-1
CuallLles whlch lmpalr phoLosynLheLlc performance
9":
LlghL lnLenslLy
8
a
L
e

o
f

p
h
o
L
o
s
y
n
L
h
e
s
l
s

13
o
C, 0.3 CC
2

13
o
C, 0.03 CC
2










Commun|cat|on, homeostas|s and energy
Un|t 3: hoLosynLhesls
I214
CnLCkCLAS1S
1he process of photosynthes|s ls one whlch converLs llghL energy lnLo chemlcal energy Lo synLhesls large organlc
molecules from smaller, lnorganlc subsLraLes. lanLs and oLher phoLosynLheLlc organlsms are Lherefore autotroph|c
(Lhey make Lhelr own food). 1here are Lwo maln sLages of phoLosynLhesls, whlch boLh occur lnslde Lhe ch|orop|ast
lnslde chloroplasLs are sLacks
of flaLLened membrane
comparLmenLs called grana
(each slngle comparLmenL
called a thy|ako|d). 1hln
membranal exLenslons connecL
one granum Lo anoLher, Lhese
are called |ntergrana| |ame||ae.
1he fluld surroundlng Lhe
grana ls called Lhe stroma. 1he
chloroplasL conLalns lLs own
unA and many rlbosomes, so
LhaL ls can make proLelns
AdapLaLlons of Lhe organelle for lLs funcLlon durlng phoLosynLhesls lnclude:
Lhe granal membranes provlde a large surface area for Lhe aLLachmenL of Lhe phoLosynLheLlc plgmenLs
(chlorophylls and caroLenolds), elecLron carrlers and enzymes for Lhe llghL-dependenL reacLlons
a neLwork of proLelns ln Lhe grana hold Lhe plgmenLs ln a very preclse manner LhaL forms Lhe phoLosysLems
allowlng for maxlmum absorpLlon of llghL
Lhe membranes have A1 synLhase aLLached Lo Lhem whlch vla chemlosmosls help Lo manufacLure A1
Lhe fluld of Lhe sLroma holds all of Lhe enzymes needed Lo carry ouL Lhe llghL-lndependenL reacLlons
.
.
.
.
.
.
.
.
. .
.
.
.
.
.
.
.
.
.
.
. .
.
.
.
.
granum
slngle
Lhylakold
rlbosomes
lnLergranal lamella
sLarch
graln
sLroma
unA
nC1CSN1nL1IC IGMLN1S AND nC1CSS1LMS
Lmbedded ln Lhe Lhylakold membranes are coloured
compounds called photosynthet|c p|gments whlch absorb
llghL of cerLaln wavelengLhs and reflecL oLher llghL. lgmenLs
are found ln large numbers arranged lnLo photosystems
1here are Lwo Lypes of plgmenL found ln phoLosysLems. 1he
chlorophylls are ch|orophy|| b and ch|orophy|| a. 1he
caroLenolds are xanthophy|| and carotene. Chlorophyll b,
xanLhophyll and caroLene are known as secondary plgmenLs,
closesL Lo Lhe sLroma slde of Lhe membrane, and absorb Lhe
llghL, exclLlng a palr of elecLrons Lo be passed Lo chlorophyll a
Lhe prlmary plgmenL
|gment Co|our
eak absorpt|on wave|ength
(nm)
Iunct|on |n photosynthes|s
Chlorophyll a ?ellow-green 430, 662
Absorbs red and blue-purple llghL
Chlorophyll b 8lue-green 433, 642
CaroLene Crange 430
Absorb purple llghL, proLecL chlorophylls
from damage from llghL and oxygen
xanLhophyll ?ellow 430-470

L|ght L|ght
chlorophyll b
xanLhophyll
caroLene
chlorophyll a
A8SCk1ICN SLC1kA AND AC1ICN SLC1kUM
A calorlmeLer can be used Lo
measure Lhe llghL absorpLlon
of dlfferenL phoLosynLheLlc
plgmenLs. llndlngs are ploLLed
onLo graphs called absorpt|on
spectra and Lhe effecLlveness
of plgmenLs ls shown on Lhe
act|on spectrum whlch shows
raLe of phoLosynLhesls aL
dlfferenL wavelengLhs of llghL.
1he Lable aL Lhe boLLom of Lhe
page summarlses Lhe
absorpLlon specLra by sLaLlng
Lhe colours/wavelengLhs each
plgmenL besL absorbs
chlorophyll b
caroLenolds
chlorophyll a

a
g
e

a
b
s
o
r
p
L
l
o
n

400 300 600 700
wavelengLh of llghL (nm)
r
a
L
e

o
f

p
h
o
L
o
s
y
n
L
h
e
s
l
s

400 300 600 700
wavelengLh of llghL (nm)
LIGn1-DLLNDLN1 kLAC1ICNS
1he flrsL of Lwo sLages ln phoLosynLhesls ls Lhe ||ght-dependent sLage, whlch Lakes place ln Lhe Lhylakold membranes. 1wo phoLosysLems are used ln
Lhls sLage: photosystem II and photosystem I, as well as Lhe many A1 synthase enzymes embedded ln Lhe membranes Lo manufacLure A1
llrsLly, Lhere ls photo|ys|s of waLer (spllLLlng uslng llghL phoLons). 1wo molecules of waLer 2P2C are spllL lnLo four elecLrons, four proLons and one
palr of oxygen aLoms. 1hls occurs aL phoLosysLem ll (whlch ls Lhe flrsL phoLosysLem Lo be used). 1he oxygen ls acLually a wasLe producL and mosLly
leaves Lhe leaf Lhrough Lhe sLomaLa, alLhough some oxygen remalns ln Lhe planL Lo be used for aeroblc resplraLlon
1he elecLrons however, are
accepLed by Lhe Sll, and Lhe
llghL energy exclLes Lhe elecLrons
causlng Lhem Lo move beLween
Lhe plgmenLs unLll Lhey are
accepLed by an e|ectron carr|er
whlch passes Lhe elecLrons
Lhrough Lhe membrane, carrler-
Lo-carrler, unLll one passes Lhe
elecLrons Lo phoLosysLem l (Sl)
1he movemenL of elecLrons
beLween elecLron carrlers
(cytochromes) releases energy,
whlch ls used Lo acLlvely pump
proLons across Lhe membrane
agalnsL Lhe gradlenL
1he elecLrons are accepLed by more carrlers from Sl and are evenLually Laken Lo an enzyme called NAD reductase whlch reduces a coenzyme called
NAD (nlcoLlnamlde adenlne dlnucleoLlde phosphaLe). 1he hydrogen lons (proLons) produced from Lhe phoLolysls of waLer remaln ln Lhe Lhylakold
space, ln order Lo bulld up a heavy concenLraLlon wlLhln Lhe Lhylakold. 1hls malnLalns a consLanL concenLraLlon gradlenL down from Lhe Lhylakold Lo
Lhe sLroma (where Lhere ls a weaker concenLraLlon). 1hls means LhaL when proLons are pumped lnLo Lhe Lhylakold space beLween phoLosysLems due
Lo Lhe release of energy from elecLron carrlers (Lhls release of energy and movemenL of proLons ls called chem|osmos|s), proLons always flow back
over Lhe membrane down Lhe gradlenL Lhrough speclallsed channel proLelns assoclaLed wlLh A1 synLhase. As proLons flow Lhrough Lhese channels,
A1 synLhase ls acLlvaLed and adds one phosphaLe group Lo a molecule of Au Lo produce A1 (Lhls ls photophosphory|at|on)
As hydrogen lons flow back across Lhe membrane lnLo Lhe sLroma, Lhey meeL up wlLh Lhe coenzyme nAu and Lhe elecLrons recelved from Lhe
elecLron carrlers. 1he elecLrons and proLons (Lwo of each) comblne Lo form Lwo hydrogen aLoms, and Lhen Lhe enzyme nAu reducLase caLalyses Lhe
reacLlon beLween nAu and Lhe hydrogen Lo produce reduced nAu (NADn or nAuP2). nAuP ls one of Lhe producLs of Lhls sLage and ls used as
Lhe maln subsLraLe for Lhe second sLage of phoLosynLhesls
P2C
xC2
2P
+

2e
-

SII SI
2e
-

2e
-

2P
+

2P
+

+2P
+

nAu nAuP
Au + l
A1
llghL llghL
thy|ako|d space
stroma
A1 synLhase
elecLron
carrlers

CCLIC AND NCN-CCLIC nC1CnCSnCkLA1ICN
uurlng Lhe llghL-dependenL reacLlons, phoLophosphorylaLlon occurs Lhls ls Lhe addlLlon of an lnorganlc phosphaLe group (l) Lo a molecule of Au Lo
produce A1 ln Lhe presence of llghL. 1here are Lwo Lypes of phoLophosphorylaLlon found ln Lhls serles of processes
1here ls cyc||c photophosphory|at|on whlch occurs
only uslng phoLosysLem l. When Sl absorbs a phoLon
of llghL, chlorophyll a ends up emlLLlng an exclLed
elecLron whlch ls passed around a chaln of carrlers
before belng reLurned Lo Sl agaln. 1he energy
released by Lhe movemenL of Lhe elecLrons here ls
used Lo generaLe A1
Non-cyc||c photophosphory|at|on uses boLh Sll and
Sl. As llghL exclLes Sll an elecLron ls passed onLo a
chaln of elecLron carrlers whlch pass lL Lo Sl. Sll
regalns lLs losL elecLron from Lhe elecLrons produced
by Lhe phoLolysls of waLer. 1he elecLron Lhen moves
from Sl Lhrough carrlers agaln and onLo nAu
reducLase. 1he movemenL of elecLrons generaLes a
proton mot|ve force whlch pumps proLons Lhrough
Lhe membrane, and as Lhey dlffuse back Lhrough
channels connecLed Lo A1 synLhase, A1 ls made
2e
-
elecLron
accepLor

2e
-
elecLron
accepLor

Au + l
A1
A1
2e
-
nAu reducLase

n2C
2e
-
2P
+
2n
+
nAu

nAuP

SII
!"#$
SI
!&$$
key:
llghL

cycllc phosphorylaLlon

non-cycllc
phosphorylaLlon

LIGn1-INDLLNDLN1 kLAC1ICNS
1he second, and flnal, sLage of phoLosynLhesls ls known as Lhe ||ght-|ndependent sLage and does noL need llghL Lo Lake place, alLhough wlLhouL Lhe
producLs of Lhe llghL-dependenL reacLlons (nAuP and A1) Lhese reacLlons cannoL occur, so wlLhouL llghL Lhls sLage also qulckly sLops worklng
1hese reacLlons Lake place ln Lhe sLroma of Lhe chloroplasL. 1he nAuP and A1 produced are already ln Lhe sLroma, and all else LhaL ls needed ls
carbon dloxlde, whlch enLers Lhe leaf by dlffuslng Lhrough Lhe sLomaLa. CC2 enLers Lhe leaf and dlffuses Lhrough Lhe alr spaces ln Lhe spongy
mesophyll layer, and Lhen Lhrough Lhe pallsade layer. 1hen lL dlffuses Lhrough Lhe cellulose cell wall, Lhe cell surface membrane, Lhey cellular
cyLoplasm and Lhen Lhe chloroplasL double membrane lnLo Lhe sLroma. 1he serles of reacLlons here ls collecLlvely known as Lhe Ca|v|n cyc|e
1 A molecule of r|bu|ose b|sphosphate (8u8), a flve-
carbon compound, has carbon dloxlde added Lo lL by Lhe
enzyme ku8|sCC
2 1hls forms an unsLable slx-carbon compound whlch spllLs
lnLo Lwo molecules of g|ycerate phosphate (C)
3 1he Lwo molecules of C are phosphorylaLed (uslng a
phosphaLe from A1) and reduced (uslng hydrogens from
nAuP) Lo form Lwo molecules of tr|ose phosphate (1),
also a Lhree-carbon compound
4 1en of every Lwelve molecules of 1 produced are
phosphorylaLed agaln and recycled lnLo slx molecules of
8u8, and Lhe oLher Lwo 1 go on Lo be made lnLo oLher
blologlcal molecules (e.g. glucose)
1here are a number of opporLunlLles for Lhe Calvln cycle Lo produce oLher
blologlcal molecules. Cne slxLh of 1 produced ls noL recycled Lo go back
around Lhe cycle, buL ls modlfled mosL ofLen lnLo g|ucose (Lwo 1 comblne Lo
produce one molecule of glucose) and oLher hexose sugars. 1 can also be
converLed lnLo glycerol, whlch comblned wlLh faLLy aclds can form llplds,
anoLher resplraLory subsLraLe
Also, someLlmes molecules of C are used Lo form amlno aclds and faLLy aclds. Amlno aclds Lhemselves mlghL be resplred Lo produce energy, and
faLLy aclds become llplds when comblned wlLh glycerol from 1. ln summary Lhe evenLs of boLh sLages of phoLosynLhesls can be expressed as:
6CC2 + 6P2C C6P12C6 + 6C2

Ca|v|n
cyc|e

8u8lsCC

x2 glyceraLe
3-phosphaLe
(3C)


(2C)
x2 Lrlose
phosphaLe
(3C)


(21)
rlbulose
blphosphaLe
(3C)


(8u8)
nAuP

nAu

CC2
A1

Au + l

Au + l

A1

hexose sugars
(e.g. glucose)
glycerol
sLarch
cellulose
polysaccharldes
(e.g. glycogen)
faLLy aclds
amlno aclds
llplds
llplds
kU8ISCC
1he enzyme r|bu|ose b|sphosphate carboxy|ase oxygenase (8u8lsCC) ls used ln Lhe Calvln cycle Lo add carbon
dloxlde Lo rlbulose blsphosphaLe, buL as 8u8lsCCs name suggesLs, lL ls also capable of comblnlng 8u8 wlLh oxygen.
When Lhls occurs, a process called photoresp|rat|on Lakes place. hoLoresplraLlon acLually undoes a loL of Lhe work
LhaL has Laken place ln phoLosynLhesls so far, so severely lowers phoLosynLhesls efflclency. lL also leads Lo Lhe
formaLlon of Loxlc hydrogen peroxlde
When 8u8 and oxygen come LogeLher, Lwo producLs are formed: phosphog|ycerate (CA) whlch slmply geLs
converLed back lnLo 8u8 and re-enLers Lhe Calvln cycle so ls noL Loo problemaLlc, and phosphog|yco|ate (C)
whlch Lakes a long serles of reacLlons Lo be reusable agaln. C musL leave Lhe chloroplasL and be converLed several
Llmes unLll lL reaches a compound whlch can re-enLer Lhe sLroma and enLer Lhe Calvln cycle once more. So evldenLly,
Lhls funcLlon of 8u8lsCC ls noL very efflclenL and slows down phoLosynLhesls slgnlflcanLly when Lhere ls Loo low a
concenLraLlon of carbon dloxlde ln relaLlon Lo normal aLmospherlc concenLraLlon (allowlng oxygen Lo flL lnLo Lhe
enzymes complex more ofLen)
LIMI1ING IAC1CkS
A ||m|t|ng factor ls any quallLy of Lhe envlronmenL whlch ls Lhe leasL favourable varlable and so lmpalrs Lhe raLe of
phoLosynLhesls or prevenLs opLlmal raLe of phoLosynLhesls
AlLhough LemperaLure does noL really affecL Lhe llghL-dependenL sLage, a sLable LemperaLure ls cruclal durlng Lhe
llghL-lndependenL sLage where Lhe enzyme acLlon of 8u8lsCC ls fundamenLal. 1emperaLures whlch flucLuaLe alLer
enzyme acLlon, and Lhose whlch are Loo hlgh or Loo low can permanenLly denature enzymes. Powever, sllghL
lncreases ln LemperaLure do lncrease enzyme acLlvlLy. 1he lncrease ln phoLosynLhesls due Lo LemperaLure lncrease
depends on Lhe llghL lnLenslLy (when Lhere ls more llghL, lncreaslng LemperaLure has a blgger effecL)
lncreaslng Lhe carbon dloxlde concenLraLlon does
lncrease Lhe raLe of phoLosynLhesls, up unLll a polnL
where elLher llghL lnLenslLy or Lhe LemperaLure
become Lhe llmlLlng facLor
lncreaslng llghL lnLenslLy generally lncreases Lhe
raLe of phoLosynLhesls because more sLomaLa open
so more carbon dloxlde enLers Lhe leaf, more llghL
can be Lrapped by chlorophyll Lo exclLe elecLrons
durlng Lhe llghL-dependenL reacLlons, and Lhere ls
more efflclenL phoLolysls of waLer, all of whlch wlll
conLrlbuLe Lo lncreaslng Lhe raLe
LlghL lnLenslLy
8
a
L
e

o
f

p
h
o
L
o
s
y
n
L
h
e
s
l
s

13
o
C, 0.3 CC2
13
o
C, 0.03 CC2
INVLS1IGA1ING 1nL kA1L CI nC1CSN1nLSIS
1he raLe of phoLosynLhesls ls measured uslng a photosynthometer whlch measures Lhe producLlon of oxygen ln a
planL. CerLaln llmlLlng facLors (usually llghL lnLenslLy, buL also LemperaLure) are ad[usLed Lo measure Lhe raLe of
oxygen producLlon under Lhe dlfferenL condlLlons
8ubbles of oxygen collecL ln a caplllary Lube one end of Lhe phoLosynLhomeLer, and when an agreed amounL of
oxygen has been produced over a conLrolled Llme perlod, a polnL can be markeL on a scale where Lhe bubbles
reached and Lhe experlmenL repeaLed under a dlfferenL condlLlon
CnkCMA1CGkAnIC SLAkA1ICN CI nC1CSN1nL1IC IGMLN1S
A chromatogram can be used Lo separaLe Lhe dlfferenL
phoLosynLheLlc plgmenLs ln a phoLosysLem. ChromaLography
ls a process whereby a mlxLure of maLerlals are separaLed by
allowingdifferentparticlestomove(inwhatisknownas
Lhe moblle phase) along a chromaLographlc sLrlp and flx
along a cerLaln place (ln whaL ls known as Lhe sLaLlonary
phase), Lhus parLlLlonlng Lhe dlfferenL parLlcles
orlgln
?LLLCW-C8AnCL
(caroLene)
?LLLCW
(xanLhophyll)
?LLLCW-C8LLn
(chlorophyll a)
C8LLn
(chlorophyll b)
plgmenL
fronL
solvenL
fronL