THE Joun~a . OF BIOLOGICAL CHEMISTRY
Vol. 234, NO. 11, No embe 1959
P in ed in
U.S.A.
Pho osyn hesis by Isola ed Chlo oplas s
X. DEPENDENCE OF CARBON DIOXIDE ASSIMILATION ON THE PHOTOCHEMICAL REACTIONS
OF CHLOROPLASTS*
A. V. TREBST, M. LOSADA, AND DANIEL I.
ARNON~
F om he Labo a o y o Plan Physiology, Uni e si y
o Cali o nia,
Be keley, Cali o nia
(Recei ed o publica ion, June 30, 1959)
Un il ecen ly, he only expe imen ally documen ed pho o-
chemical ac i i y o isola ed chlo oplas s was he Hill eac ion
(3) in which illumina ed chlo oplas s e ol e oxygen in acco dance
wi h Equa ion 1:
A + Hz0 + AH2 + $0, (1)
whe e A cp cscn s a nonphysiological elec on o hyd ogen
accep o such as e icyanide o benzoquinone (4).
Recen expe imen s (5-8) ha e shown ha he Hill eac ion
is a agmen o a pho osyn he ic phospho yla ion o he non-
cyclic ype (9). Fo example, wi h e icyanide as he elec on
accep o , oxygen e olu ion is coupled wi h ATP o ma ion, in
acco dance wi h Equa ion 2.*
4Fe3+Cy + 2HzO + 2AnP + 2Pi -+
4Fez+Cy + 02 + 2ATP + 4H+ (‘I
The ecogni ion o he Hill eac ion as an uncoupled pho o-
phospho yla ion was pa alleled (5-S) by he iden i ica ion o he
physiological coun e pa o Reac ion 2, he noncyclic pho o-
phospho yla ion eac ion in which oxygen e olu ion and ATP
o ma ion a c linked wi h TPN educ ion (Equa ion 3).
2TPN + 2HeO + 2ADP + 2P< -- 2TPNHz + 02 + PATP (3)
Isola ed chlo oplas s ha e also been ound o o m ATP by
a cyclic pho ophospho yla ion (5-8) in which ATP is he sole
p oduc o he pho ochemical eac ion (Equa ion 4).
Al1P + Pi -+ ATP
(4)
In addi ion o pho osyn he ic phospho yla ion ecen wo k
has also p o ided di ec expe imen al e idence (10-14) o he
o en asse ed bu ne e p e iously demons a ed capaci y o
isola ed chlo oplas s o assimila e CO2 pho osyn he ically o he
le el o ca bohyd a es. CO2 assimila ion by isola ed chlo o-
plas s was ound o be a da k p ocess (15) dependen on a
pho ochemically gene a ed “assimila o y powe ” comp ising
wo componen s: TPNH2 and ATP o med by he noncyclic
and cyclic pho ophospho yla ion eac ions (Equa ions 3 and 4).
Noncyclic pho ophospho yla ion (Equa ion 3) p o ides all
he h ee expec ed p oduc s o he ligh phase o pho osyn hesis:
02, TPNH2, and ATP. Cyclic pho ophospho yla ion (Equa ion
* P elimina y epo s o his wo k ha e been published p e i-
ously (1) 2).
Aided byg an s om he Na ,ional Ins i u eso Heal h,
IJni ed
S a es Public-Heal h Se ice, and he O ice o Na al Resea ch.
1 The abb e ia ions used a e: FUN, la in mononucleo ide; Pi,
o hophosphn e.
4) supplies only ATP and he pa icipa ion o his eac ion in
CO2 assimila ion would bc needed only i he ATP o med in
Reac ion 3 we e insu icien o COz assimila ion o he le el o
ca bohyd a e.
The pu pose o his a icle is o p esen e idence ha in
pho osyn hesis by isola ed chlo oplas s, assimila ion o CO2 o
he le el o suga phospha es equi es a p ope ly balanced pa ici-
pa ion o bo h ligh eac ions: cyclic and noncyclic pho ophos-
pho yla ion (Equa ions 4 and 3). In he p esen in es iga ion
he balance be ween he wo ligh eac ions was main ained by
egula ing he concen a ion o co ac o s o cyclic pho ophos-
pho yla ion (16, 17).
EXPERIMENTAL
Me hods-B oken chlo oplas s om spinach (16) o suga
bee lea es (18) we e used in all he expe imen s desc ibed he ein.
The b oken chlo oplas s we e p epa ed wi h asco ba e (Cl,
pa icles) as p e iously desc ibed (16). Chlo oplas ex ac
was p epa ed wi h 0.035
M
NaCl and used wi hou dialysis.
COz ixa ion was ca ied ou a 20” in ec angula Wa bu g
manome e essels, lushed wi h a gon gas be o e u ning on he
ligh (app oxima ely 23,000 lux). The pe iod o illumina ion
was 30 minu es. The eac ion was s opped by adding o each
essel 0.1 ml o glacial ace ic acid. To al COz ixa ion was
measu ed by pipc ing aliquo s om each ea men on s ainless
s eel planche s, e apo a ing o d yness, and coun ing Cl4 wi h
a hin window Geige -Miillc coun e . Fo he iden i ica ion
o he p oduc s o CO% ixa ion he con en s o he Wa bu g
essels we e cen i uged, and aliquo s o he supc na an liquid
we e subjec ed o wo-dimensional pape ch oma og aphy (on
Wha man No. 41 pape ) using as sol en s (a) 80 phenol-20 wa e
and (b) a mix u e o 52 pa s n-bu anol, 14 pa s glacial ace ic
acid and 35 pa s wa e .
The adioac i i y in he indi idual compounds, loca ed on he
pape s by adioau og aphy, was de e mined by coun ing on he
d ied pape s. The indi idual compounds we e iden i ied by
elu ion and subsequen coch oma og aphy wi h samples o
au hen ic compounds. Suga phospha es we e u he iden i ied
by dephospho yla ion wi h phospha asc (Polidase) and cch oma-
og aphy wi h he co esponding au hen ic suga s.
RESULTS
Expe imen al documen a ion o he pa icipa ion o cyclic
pho ophospho yln ion (Equa ion 4) in COz assimila ion by
chlo oplas s equi ed use o a sys em di e en om he one
desc ibed be o e in which Tl’KI& and ATP we e al eady supplied
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3056 CO2 Assimila ion by Chlo oplas s Vol. 234, No. 11
in excess, and a second eac ion o gene a ing ATP would be
supe luous (cj. T ebs e al. (15), Table I). In he expe imen s
o be desc ibed p esen ly, e idence o he pa icipa ion o cyclic
pho ophospho yla ion in CO2 assimila ion o isola ed chlo o-
plas s was ob ained in a “ca aly ic” sys em, i.e. one in which,
as in an in ac cell, TPNH, and ATP we e o med in ca aly ic
amoun s and CO2 ixa ion was he e o e possible only in he
ligh while TPNHz and ATP we e being con inuously egene a ed
a he expense o abso bed ligh ene gy.
E$ec o FMN
on CO2 Fixa ion-As shown in Table I, when
he exogenous supply o ATP (2 /Imoles) was eplaced by a
“ca aly ic” sys em con aining 0.5 pmole o ADP, o hophos-
pha e, and 0.3 pmole o TPN, CO% ixa ion was sha ply educed
(T ea men B). Howe e , he addi ion o he eac ion mix u e
o an ex emely minu e quan i y o FMN (0.001 pmole) g ea ly
inc eased o al COZ ixa ion (T ea men C). In he p esence o
his minu e amoun o FMN he capaci y o chlo oplas s o
ix COZ was equal o, i no g ea e han,
ha in T ea men A in
which 2 pmoles o exogenous ATP we e supplied.
TABLE I
E ec oj ibojla in phospha e (FMN) on CO, ixa ion dependen on
egene a ion o ATP in ligh
Each essel con ained
in a inal olume o 2.5 ml : b oken chlo o-
plas s (Cl.) con aining 0.5 mg o chlo ophyll; chlo ophyll ex ac
(CE) equi alen o 2 mg o chlo ophyll; and he ollowing in
wmoles: is(hyd oxyme hyl)aminome hane pH 7.5, 80; MgCL,
5; MnC12, 2; sodium asco ba e, 10; sodium phospha e, 5; educed
glu a hione, 5; ibose 5-phospha e, 0.3; sodium ca bona e-W, 10.
In addi ion o he indica ed ATP and ADP supplemen s, T ea -
men A included 2 moles o TPN and each o T ea men s B and
C, 0.3 pmole o TPN.
T ea men To al C”00a ixed
c.p.m.
A. Con ol, 2 amoles o ATP.
232,000
B. 0.5 @mole o ADP.. 109,ooo
C. 0.5 pmole o ADP, 0.001 Hmole o FMN. . 265,000
TABLE II
E$ec
o
FMN
concen a ion
on CO2 $xa ion by illumina ed
chlo oplas agmen s om suga bee s
The eac ion mix u e was he same as ha desc ibed o T ea -
men B in Table I excep ha glu a hione was omi ed and each
essel con ained, in addi ion o he indica ed concen a ion o
FMN, 0.3 mole o glucose l-phospha e ins ead o ibose li-phos-
pha e. Final olume 3.0 ml.
T ea -
men FMN added
jmlle/3 ml c.p.m. %
%
A 0 110,000 93 3
0.0002 105,000 32 65
0.0005 220,000 34 63
B 0.001 208,000 20 80
0.005 231,000 10 85
0.01 208,000 14 77
0.1 275,000 60 35
C 0.5 164,000 75 24
* Suga mono- and diphospha es and dihyd oxyace one phos-
pha e.
To al C” ixed as
To al Co? ixed Phospho&e i Suga
phospha es*
The e ec o FMN on COz ixa ion was in es iga ed in g ea e
de ail by a ying he concen a ion o his co ac o o cyclic
pho ophospho yla ion. In addi ion o an inc ease in o al COz
ixa ion, a s iking co ela ion was obse ed be ween he con-
cen a ion o added FMN and he pa e n o ca bon compounds
o med. Table II shows ha , depending on he concen a ion
o FMN, he p oduc s o CO2 assimila ion we e ei he p edomi-
nan ly suga phospha es, which a e aken he e as a measu e o a
educ i e (pho osyn he ic) assimila ion pa e n, o phospho-
glyce ic acid. Phosphoglyce ic acid was he chie p oduc o
CO2 assimila ion when ei he no FMN (T ea men A) o a
ela i ely la ge amoun o FMN (0.5 pmole) (T ea men C)
was added o he eac ion mix u e. A a ange o low FMN
concen a ion, om abou 0.001 o 0.01 c mole pe 3 ml, suga
phospha es we e he p edominan p oduc s o CO2 assimila ion
(T ea men B). These e ec s o FMN concen a ion on he
pa e n o CO2 assimila ion a e illus a ed in Figs. 1 o 3.
E ec o Vi amin KS
and Phenazine
Me hosulja e on CO2
Fixa ion-Simila e ec s on o al CO2 ixa ion and he pa e n
o compounds o med we e also obse ed by adding wo o he
co ac o s o cyclic pho ophospho yla ion: i amin K and
phenasine me hosul a e. Typical esul s a e shown in Table
III. A he same mola concen a ion, FMN, i amin K3, o
phenasine me hosul a e p oduced compa able e ec s on o al
CO2 ixa ion and on he ela ion be ween phosphoglyce a e and
suga phospha es. The addi ion o small amoun s o any one
o he h ee co ac o s o cyclic phospho yla ion (0.01 pmole pe
3 ml) inc eased o al CO2 ixa ion se e al imes and ga e suga
phospha es as he main p oduc s o pho osyn hesis. A a
concen a ion o 0.3 pmoles pe 3 ml he o ma ion o suga
phospha e was ma kedly dec eased and phosphoglyce ic acid
appea ed as he p incipal p oduc o CO2 ixa ion.
DISCUSSION
The esul s o his in es iga ion show ha he addi ion o one
o he co ac o s o cyclic pho ophospho yla ion, FMN, i amin
FIG.
1. Radioau og aph o a ch oma og am showing p oduc s
o pho osyn he ic CWn assimila ion by illumina ed chlo oplas s
in he absence o added ibo la in phospha e
(FMN). O he
con-
di ions as gi en in Table 2.
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No embe 1959
A. V. T ebs , M. Losada, and D. I. A wn 3057
FIG. 2. Radioau og aph o a ch oma og am showing p oduc s
o pho osyn he ic CY402 assimila ion by illumina ed chlo oplas s FIG. 3. Radioau og aph o a ch oma og am showing p oduc s
supplied wi h 0.001
PM
ibo la in phospha e (FMN). O he con- o pho osyn he ic CIQ assimila ion by illumina ed chlo oplas s
di ions as gi en in Table 2. supplied wi h 0.5 PM ibo la in phospha e (FMN). O he condi-
ions as gi en in Table 2.
K, o phenazine me hosul a e, has a ma ked e ec on COZ
TABLE
III
assimila ion in a “ca aly ic” sys em in which CO2 assimila ion
E$ec o concen a ion o co ac o s o cyclic pho ophospho yla ion
by chlo oplas s depends on egene a ion by ligh o assimila o y
on CO2 $xa ion by illumina ed spinach chlo oplas s
powe , i.e. on TPNHz and ATP. Since he e is no e idence o Expe imen al condi ions as in Table II, excep o he indica ed
indica e ha minu e amoun s o FMN, i amin K, o phenazine addi ions o FMN, i amin K3, and phenazine me hosul a e.
me hosul a e in luence di ec ly he enzyma ic eac ions esponsi-
ble o COZ assimila ion, i seems easonable o seek an explana-
ion o he obse ed esul s in he p e iously obse ed e ec s o
hese co ac o s o cyclic pho ophospho yla ion on he cou se o
he ligh eac ions (5, 17).
Co ac o added To al C’“Oz ixed
pmole C.). n.
79,000
318,066
415,060
530,000
390,090
310,660
267,666
Wi hou he addi ion o one o he co ac o s o cyclic pho o- None
phospho yla ion, he ligh eac ions o isola ed chlo oplas s a e FMN
limi ed o noncyclic pho ophospho yla ion (Equa ion 3). The
a io o TPNHz o ATP o med is 1: 1. Adding a minu e amoun Vi amin K1
o FMN, i amin KS, o phenazine me hosul a e o a noncyclic
pho ophospho yla ion sys em inc eases ATP o ma ion wi hou Phenazine
app eciably dep essing oxygen e olu ion and he co esponding me hosul a e
TPNHz accumula ion (c . A non e al. (17), Tables 3 and 4).
The a io o ATP o TPNH o med becomes g ea e han 1.
I appea s likely ha unde hese condi ions cyclic pho ophos-
pho yla ion (Equa ion 4) is supe imposed on he noncyclic
p ocess (Equa ion 3) and con ibu es addi ional ATP.
The shi om phosphoglyce a e o suga phospha es as he
main p oduc s o CO2 assimila ion (compa e Fig. 1 and Fig. 2)
is explained by he addi ional ATP o med in he ligh by cyclic
pho ophospho yla ion as a esul o adding one o i s co ac o s.
Wi hou his addi ion he ligh eac ion is limi ed o noncyclic
pho ophospho yla ion (Equa ion 3) and ails o p o ide su i-
cien ATP o he educ ion o CO2 o suga s. I is concluded
ha he 1:l a io o ATP o TPNHs which cha ac e izes Reac-
ion 3 is insu icien o he educ ion o COZ o he le el o
ca bohyd a e.
The need o mo e ATP han TPNHz in CO2 assimila ion is
consis en wi h he iew ha he e a e wo si es o phospho yla-
ion bu only one si e o educ ion in he o ma ion o ca bo-
hyd a es, The wo phospho yla ion eac ions a e: he
phospho ibulokinase eac ion (Equa ion 5) (19-22) and he phos-
-
I
% %
0.01
0.3
0.01
0.3
0.01
0.3
15
69
65 32
16 77
92 6
8 86
70 27
* Suga mono- and diphospha es and dihyd oxyace one phos-
pha e.
To al C”Oz ixed as
Phospho-
glyce a e Suga
phospha es*
phoglyce a e kinase eac ion (Equa ion 6). TPNHz is used only
in he iosephospha e dehyd ogenase eac ion (Equa ion 7). All
h ee enzymes conce ned ha e been ound in he chlo oplas
p epa a ions used in hese expe imen s.2
Ribulose 5-phospha e + ATP phospho ibulokinase , (5)
ibulose 1,5-diphospha e + ADP
3-phosphoglyce ic acid + ATP phosphoglyce a e kinase b
(6)
1,3-diphosphoglyce ic acid + ADP
1,3-diphosphoglyce ic acid + TPNHz
iosephospha e dehyd ogenase > (7)
glyce aldehyde 3-phospha e + H3POd + TPN
2 M. Losada, A. V. T ebs , and D. I. A non, manusc ip sub-
mi ed o publica ion.
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3058
CO2 Assimila ion
by Chlo oplas s
Vol. 234, No. 11
When, in he absence o added co ac o s o cyclic pho ophos-
pho yla ion, he ligh eac ions o chlo oplas s a e limi ed o
noncyclic pho ophospho yla ion (Equa ion 3), he ATP o med
is insu iicicn o bo h phospho yla ing si es. As he ATP
o med by Reac ion 3 is used in he phospho yla ion o ibulose
5.phospha e (Equa ion 5), he 1: 1 a io be ween ATP andTPNH:!
ncedcd o he educ ion o phosphoglycc a c (Equa ions 6 and
7) begins o dcc msc. E en ually, in a “ca aly ic” sys em, his
would b ing he educ ion o phosphoglycc a c o a hal . Phos-
phoglycc a c would hen end o accumula e (Fig. 1) as he
ibulosc diphospha e o med in Reac ion 5 ac s as a CO2 accep o
and is subsequen ly clea ed by he ca boxylase eac ion (21-24).
The p edominance o phosphoglycc ic acid among he p oduc s
o COZ assimila ion when a la ge amoun o one o he ac o s
o cyclic pho ophospho yla ion is added o he eac ion mix u e
(Fig. 3) is also explained by he p e iously obsc cd e ec s o
FMN, i amin K, o phenazinc mc hosul a e on noncyclic
pho ophospho yla ion (5, 17). When one o hese co ac o s is
added o a noncyclic pho ophospho yla ion sys em a he highe
concen a ions shown in Fig. 3 (also in Tables II and III), he
noncyclic pho ophospho yla ion is con e ed o he cyclic ype
(5, 17). Oxygen e olu ion and he accumula ion o educed
TPN a e supp essed, phospho yla ion is sha ply inc eased, and
he p incipal p oduc o he ligh eac ion is ATI’ (cj. A non e al.
(17), Figs. 7 and 8). Unde hese condi ions he educ ion o
phosphoglycc a c acid could no occu , since he ioscphospha e
dchyd ogenasc eac ion (Equa ion 7) would be blocked by a lack
o he educ an TPNH2.
I appea s, he e o e, ha he nonoccu ence o a educ i e
(pho osyn he ic) COz assimila ion esul ing in he o ma ion o
suga phospha e can be caused ei he by a sho age o ATP
(Fig. 1) o o TPNHz (Fig. 3). In bo h cases phosphoglyce a e
would appea as he p edominan p oduc o CO* assimila ion
because i s u he assimila ion would be blocked.
Al hough on he basis o p esen e idence he wo si es o
ATP ac ion in COZ assimila ion appea o be he phospho-
ibulosc and he phosphoglyce a c kinase eac ions (Equa ions
5 and 6), he e is a possibili y ha he second si e o ATP
ac ion in he phospho yla ion may be no he phospho yla ion
o phosphoglyce a c bu ha o some uns able 6 ca bon com-
pound, which, in ai o, unde goes phospho yla ion be o e educ-
ion by TPSHz wi hou b eaking up in o 2 moles o phospho-
glyce a c (25, 26).
In he expe imen s epo ed he e he o ma ion o suga,
phospha es which is aken as a mcasu c o pho osyn he ic CO2
assimila ion occu ed only when a p ope balance was main ained
bc wccn cyclic and noncyclic pho ophospho yla ion. In isola ed
chlo oplas s his balance was main ained by adding di e en
amoun s o one o he ca alys s o cyclic pho ophospho yla ion.
I is assumed ha he in ac ccl1 has sui able physiological
cguln o y mechanisms o keeping he wo cnc ions in balance.
SUMhl.QRy
CO? assimila ion in isola ed chlo oplas s was inycs iga. cd in a
“ca aly ic” sys em unde h ee condi ions: (I) when he pho o-
chemical phase was limi ed o noncyclic pho ophospho yla ion,
(2) when he pho ochcmical phase was limi ed o cyclic pho o-
phospho ~ln iol1, and (3) when he pho ochemical phase included
bo h cyclic and noncyclic pl~o ophospho yla ions.
Unde Condi ions 1 o 2, CO2 assimila ion was limi ed almos
en i ely o he o ma ion o phosphoglyce a e. Suga phos-
pha es we e he p edominan p oduc s o CO2 assimila ion only
in Condi ion 3. These esul s a e in e p e ed as ha ing been
caused by a sho age o adenosine iphospha e (ATP) in Condi-
ion 1 and o educed iphosphopy idine nuclco ide (TPNH2)
in Condi ion 2; only in Condi ion 3 was a p ope balance es ab-
lished be ween ATP and TPNHz o med a he expense o ligh
ene gy, o make he o ma ion o suga phospha es possible.
The balance be ween he noncyclic and cyclic pho ophos-
pho yla ion necessa y o b ing abou a pho osyn he ic ( educ-
i e) ype o CO2 assimila ion was main ained by adding o he
econs i u ed chlo oplas sys em minu e amoun s o one o he
ca alys s o cyclic pho ophospho yla ion, ibo la in phospha e,
i amin K, o phenazine me hosul a e.
1.
2.
3.
4.
5.
6.
7.
a.
9.
10.
11.
12.
13.
14.
15.
16.
17.
18.
19.
20.
21.
22.
23.
24.
25.
26.
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