ANABOLISME

FOTOSINTESIS
 PHOTOSYNTHETIC
ELECTRON TRANSPORT

Space pictures from

http://visibleearth.nasa.gov
Fotosintesis
terjadi di
Kloroplas sel
daun
 Fase Fotosintesis

Fotosintesis memiliki 2 fase utama:

1. Penyerapan cahaya dan produksi bentuk bentuk energi kimia (reaksi terang)
2. Fiksasi dan reduksi karbon (reaksi gelap)
 Fase-fase Fotosintesis

1. Reaksi cahaya (light harvesting and energy transduction)

- Memerlukan cahaya
- Dikatalisis oleh kompleks protein yang tertanam pada membran tilakoid.

Fase-fase:

a. Penyerapan cahaya

b. Transfer energi ke pusat reaksi fotosistem

c. Pemecahan air dan donor elektron (produksi O2 )

d. Transpor elektron dari PSII ke PSI, dengan proton pumping

e. Transpor elektron dari PSI ke NADP+, membentuk NADPH

f. Sintesis ATP oleh kemiosmosis

 Fase-fase fotosintesis
2. Fiksasi dan Reduksi Karbon (reaksi gelap)

– Terjadi di stroma
– Terjadi dalam keadaan gelap jika ATP dan NADPH ditambahkan
secara artifisial.
– Dikatalisis oleh protein terlarut dalam stroma atau sitosol

Fase-fase:

a. Karboksilasi Ribulosa-1,5 bisphosphate (RuBP) olehRuBP

carboxylase/ oxygenase (Rubisco), membentuk PGA

b. Reduksi PGA menjadi triosa phosphates

c. Regenerasi RuBP dalam the Calvin cycle

d. Produksi tepung dan sukrosa

 Penyerapan
cahaya dan
pemindahan
Ke pusat reaksi
Aliran energi antara Fotosistem dan ke NADP+
 ENERGY FLOW BETWEEN THE PHOTOSYSTEMS AND TO NADP+

High Free Energy

- +1.0
Redox potential (volts)

-0.0

- -1.0

Low Free Energy

This diagram is often called the Z-scheme
 Electron Transport in the Thylakoid Membrane
The red X indicates
that protons do not
directly pass through
the cytochrome
complex.

H+
Protons cross the membrane via
oxidation and reduction of quinones
 Bound Plastoquinone A (Qa)
PHOTOSYSTEM II
Plastoquinone docking site (Qb site)
Inner antennae (chlorophyll protein complex)

Outer antennae (chlorophyll-containing

light harvesting protein complex)

Thylakoid Membrane

The Reaction Center

Pheophytin (like chlorophyll, but no magnesium in

ring-structure)
water-splitting
complex Special pair of chlorophyll a (termed P680)
 Iron-sulfur complex
PHOTOSYSTEM I
phylloquinone
PHOTOSYSTEM II Chlorophyll a

Inner antennae
Outer antennae (chlorophyll-containing
light harvesting protein complex)
The Reaction Center

Special pair of chlorophyll a (termed P700)

 CORE PROTEINS
At the center of each photosystem are two intertwined
proteins to which the special pair chlorophylls, and the initial
electron transport intermediates are bound.

PHOTOSYSTEM II PHOTOSYSTEM I

D1 and D2 A1 and A2
Interchain and Terminal Photosynthetic
Electron Transporters NADP reductase
Mobile Ferridoxin
Plastoquinine

Cytochrome B6f Plastocyanin

 NADP reductase
4 photons
4 photons 2H+ + 2NADP
2 H+ NADPH
PQ Fd

CYT B6f

PSII O2 + 4H+
PSI
PC 4 e-
2H2O

The Path of Electron and Proton Flow in Photosynthetic

Electron Transport
 NADP reductase
photons
photons 2H+ + 2NADP
2 H+ NADPH
PQ Fd

CYT B6f

PSII O2 + 4H+
PSI
PC 4 e-
2H2O

The Path of Electron and Proton Flow in Photosynthetic

Electron Transport
 Under conditions where NADP+ regeneration is slow,
or ATP demand is high, the leaf can cycle electrons
between plastoquinone and PSI, and in doin so pump
protons across the membrane. This is termed cyclic
photophosphrylation.

NADP reductase
2 H+ Fd

PQ
CYT B6f

PSII PC PSI

4 H+

CYCLIC PHOTOPHOSPHORYLATION
 Under conditions where NADP+ regeneration is slow,
or ATP demand is high, the leaf can cycle electrons
between plastoquinone and PSI, and in doin so pump
protons across the membrane. This is termed cyclic
photophosphrylation.

NADP reductase
2 H+ Fd

PQ
CYT B6f

PSII PC PSI

4 H+

CYCLIC PHOTOPHOSPHORYLATION
 H+
PHOTOPHOSPHORYLATION
H+

ATP Synthase Chloroplast stroma – region of high pH

(F-type ATPase) H+ H+
H+
H+ H+ H+ H+
H+
H+
ADP H+
H+ ATP PHOTOSYSTEM I
+ Pi

H+ H+ H+ H+ H+ H+H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+
H+ H+ H+ H+ H+ H+H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+
H+ H+ H+ H+ H+H+
H+ H3+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H
+ H+ H+
H+ H+ H+ H+ H+ H+H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+ H+

Thylakoid Lumen – compartment of low pH

 SINTESIS ATP FOTOSINTETIK
(FOTOFOSFORILASI)

The synthesis of ATP in chloroplasts occurs when the energy of a proton

gradient is coupled to phosphorylate ADP to ATP by an F-type ATPase.

Protons are accumulated on the inside of the thylakoid lumen and depleted in
the stroma compartment

pH of chloroplast compartments

Thylakoid lumen Stroma

Dark 7.0 7.0

Light 5.2 8.2
 FIKSASI KARBON FOTOSINTETIK
EMPAT FASE:

A) Karboksilasi Ribulose Bisphosphate (RuBP)

menjadi Phosphoglycerate (PGA), dikatalisis
oleh enzyme RuBP carboxylase/oxygenase
(Rubisco)

B) Reduksi PGA menjadi triosa fosfat

C) Regenerasi RuBP
D) Sintesis produks akhir (sintesis tepung dan
sukrosa)
 CALVIN CYCLE
• Molekul berenergi
tinggi dihasilkan
selama reaksi
terang digunakan
untuk mereduksi
CO2 menjadi
glukosa
– Glukosa ini
dapat
dikonversi
menjadi
senyawa lain
(asam amino,
dsb.)
– Glukosa ini
dapat
dioksidasi
menjadi
penyimpan
energi
 Fase 1: Karboksilasi

RuBP (5 carbon) + CO2 RUBISCO 2 PGA (3 carbon)

Note: The oxygen in

CO2 is incorporated
into one of the PGA
molecules. It is not
released as O2.
 FASE 2: REDUKSI PGA
ADP + Pi NADP+ Triose-phosphate
isomerase
PGA 1,3 bisPGA G3P DHAP
PGA NADP
kinase
ATP NADPH dehydrogenase

These two molecules

are collectively termed
triose phosphates.
PGA - 3-phosphoglyceric acid
G3P -glyceraldehyde 3 phosphate
DHAP -dihydroxyacetone phosphate
TP - triose phosphates (G3P and DHAP)
 FASE 3: REGENERASI RUBP
•Triosa fosfat digabung dan dikombinasi melalu
kompleks reaksi berseri menjadi Ru5P (Ribulose
5-P)

•Tahap akhir regenerasi RuBP:

Ru5P + ATP RuBP + ADP + Pi

Karboksilasi, reduksi dan regenerasi secara

kolektif menjadi Siklus Calvin-Benson.
 FASE 4:SINTESIS TEPUNG DAN SUKROSA

Konversi triosa fosfat menjadi tepung dan

sukrosa, menggunakan energi dari ATP.

A. Sintesis tepung terjadi di stroma.

- tersimpan sebagai cadangan makanan
pada tempat penimbunan
B. Sintesis sukrosa terjadi di sitosol
- untuk pengangkutan ketempat penimbunan
 The Phases of Photosynthesis

Carboxylation

The Calvin
Cycle

PGA Reduction
RuBP
Regeneration

End Product
Synthesis
 Bagaimana terjadinya reaksi gelap?

• CO2 diikat oleh RuBP menjadi senyawa 6 C yang labil.

Kemudian dipecah menjadi 2 PGA
• Masing-masing PGA menerima gugus fosfat dari ATP
dan H serta elektron dari NADPH, menghasilkan PGAL
• Untuk 6 molekul CO2 akan dihasilkan 12 PGAL
• Dari 12 PGAL, 10 molekul kembali ke tahap awal
menjadi RuBP, dan seterusnya RuBP akan mengikat
CO2 yang baru.
• Dua PGAL lainnya berkondensasi menjadi glukosa 6
fosfat yang merupakan prekursor produk akhir yaitu
tepung dan sukrosa
 Starch synthesis
(occurs in the chloroplast and plastids)

Glucose ADP Glucose Glucose Glucose Glucose Gluco

ADP-glucose Starch polymer

Starch
synthase

ADP

Glucose Glucose Glucose Glucose Glucose Gluco

Starch polymer plus one more glucose unit

 Sucrose synthesis
(occurs in the cytosol)

Glucose UDP Fructose P

UDP-Glucose Fructose-6-phosphate
(UDP is uridine diphosphate)

Glucose
Sucrose
synthase

Sucrose
UDP phosphate

Fructose
synthase
Glucose Fructose P

Sucrose 6-phosphate Sucrose

 This diagram from your text
shows the path of carbon
flow in the “Calvin” cycle
Three turns for one and to the end products,
triose phosphate starch and sucrose. It is set
up to show the number of
turns needed to produce one
triose phosphate.

The Calvin Triose phosphates also serve as

the precursors for many of the
Cycle
complex molecules of life such as
fats, lipids, nucleic acids, amino
acids and proteins. Some consider
them the real end products of
photosynthesis. However, because
starch and sucrose synthesis are
closely linked with photosynthesis
through Pi release, photosynthesis
experts usually consider starch and
sucrose as the end products of
photosynthetic metabolism.
 The Phases of Photosynthesis

Carboxylation

The Calvin
Cycle

PGA Reduction
RuBP
Regeneration

End Product
Synthesis
 The Phases of Photosynthesis
There are four
Carboxylation steps where
energy in the form
of ATP or NADPH
3. At are consumed.
Ru5P
1. At PGA
kinase
kinase
The Calvin
Cycle
PGA Reduction

PGA Reduction
RuBP
Regeneration
2. At NADP
dehydrogenase

4. In starch and sucrose End Product

synthesis Synthesis
 STOICHIOMETRY OF CARBON FIXATION
For 6 RuBP carboxylated (30 carbon atoms) there are:

12 PGA produced (36 carbons)

18 ATP consumed (12 for PGA reduction and 6 for Ru5P phosphorylation)
12 NADPH consumed in PGA reduction to triose phosphate

10 triose phosphates (30 carbon atoms) must be diverted back into RuBP
regeneration to replace the 6 RuBP consumed.

This leaves 2 triose phosphates as net product which can be used for sugars,
starches, lipids, amino acids and proteins.

Simplifying - For each CO2 fixed there are:

3 ATP consumed (from 9 protons, 8 photons)
2 NADPH consumed (from 8 photons)
1 O2 released