FOTOSINTESIS: FIKSASI

KARBONDIOKSIDA DAN SINTESIS

KARBOHIDRAT

Y. Sri Wulan Manuhara

Setelah energi asimilasi yang berupa ATP dan NADPH2
diperoleh dari reaksi terang, energi ini digunakan untuk
mengubah CO2 menjadi karbohidrat

Untuk mengetahui tahap-tahap reaksi kimia dalam reaksi

gelap para ilmuwan menggunakan cara autoradiografi dan
kromatografi dua dimensi. Dengan menggunakan isotop
14
C dan dengan mengubah-ubah waktu antara awal
pemberian cahaya dan pematian daun, kemudian senyawa
yang diperoleh dianalisis dengan kromatografi dapatlah
diketahui senyawa-senyawa apa yang dihasilkan.

(Benson dan Calvin, 1946-1953)

Reaksi Penambatan CO2

• Sekitar tahun 1954 ditemukan sebuah enzim yang

mengkatalisis secara irreversible reaksi penggabungan
CO2 dan RuBP (ribulosa bifosfat) untuk membentuk 2
molekul 3-PGA (fosfogliseraldehida).

• Enzim yang mengkatalisis reaksi ini dinamakan ribulosa

bisfosfat karboksilase (rubisko)

• Enzim ini penting bukan hanya karena reaksi yang

dikatalisisnya itu penting, tapi karena sejauh ini juga
merupakan protein yang terbanyak dibumi (Ellis, 1979)
1/8-1/4 dari protein daun adalah enzim rubisco.
1. Dari hasil percobaan diketahui bahwa pada berbagai
jenis tumbuhan fiksasi CO2 berlangsung melalui
beberapa jalur, yaitu:

1. Jalur/Daur Calvin (Jalur C-3)

2. Jalur Hatch – Slack (Jalur C-4)
3. Jalur Crasulacean Acid Metabolism (CAM)
DAUR CALVIN/JALUR C3/CALVIN CYCLE

• The Calvin cycle can be subdivided into the 3 basic

steps:
1. The carboxylation of the C5 sugar, rubilose 1.5-
bisphosphate, leading to the formation of two
molecules of 3-phosphoglycerate; or the
incorporation of CO2 in the RuBP
2. The reduction of 3-phosphoglycerate to triose
phosphate
3. The regeneration of the CO2 acceptor rubilose 1.5-
bisphosphate from triose phosphate.
 -In phase 1 (Carbon Fixation), CO2 is
incorporated into a five-carbon sugar named
ribulose bisphosphate (RuBP). The enzyme
which catalyzes this first step is RuBP
carboxylase or rubisco. It is the most
abundant protein in chloroplasts and
probably the most abundant protein on Earth.
The product of the reaction is a six-carbon
intermediate which immediately splits in half
to form two molecules of 3-phosphoglycerate

- In phase 2 ( Reduction), ATP and NADPH 2

from the light reactions are used to convert 3-
phosphoglycerate to glyceraldehyde 3-
phosphate, the three-carbon carbohydrate
precursor to glucose and other sugars.
- In phase 3 (Regeneration), more ATP is used to
convert some of the of the pool of glyceraldehyde 3-
phosphate back to RuBP, the acceptor for CO2,
thereby completing the cycle. For every three
molecules of CO2 that enter the cycle, the net output
is one molecule of glyceraldehyde 3-phosphate
(G3P). For each G3P synthesized, the cycle spends
nine molecules of ATP and six molecules of
NADPH2. The light reactions sustain the Calvin
cycle by regenerating the ATP and NADPH 2.
JALUR C4 (JALUR HATCH-SLACK)

• Pada beberapa jenis tumbuhan diketahui bahwa

senyawa pertama yang dihasilkan bukan PGA,
tetapi asam dikarboksilat dengan rantai C-4.
Diketemukan bahwa CO2 diikat oleh fosfoenol
piruvat (PEP) menjadi asam oksaloasetat.

• Tumbuhan yang melakukan karboksilasi jalur C-

4 ini mempunyai struktur anatomi daun yang
khusus yang disebut tipe Kranz, yaitu berkas
pengangkut pada daun dibungkus oleh selubung
berkas pengangkut (bundle sheat) yang terdiri
dari sel-sel parenkim besar berisi kloroplas.
C4 carbon fixation
The first step in the pathway is the conversion of pyruvate
to PEP by the enzyme pyruvate-phosphate dikinase (
pyruvate, orthophosphate dikinase). This reaction requires
inorganic phosphate and ATP plus pyruvate, producing
phosphoenolpyruvate, AMP, and inorganic pyrophosphate

(PPi).
The next step is the fixation of CO2 into PEP by
the enzyme PEP carboxylase. Both of these steps
occur in the mesophyll cells:

pyruvate + Pi + ATP → PEP + AMP + PPi

PEP carboxylase + PEP + CO2 → oxaloacetate

The product is usually converted to malate, a
simple organic compound, which is transported to
the bundle-sheath cells surrounding a nearby vein.
Here, it is decarboxylated to produce CO2 and
pyruvate. The CO2 now enters the Calvin cycle
and the pyruvate is transported back to the
mesophyll cell.
Since every CO2 molecule has to be fixed twice,
first by 4-carbon organic acid and second by
RuBisCO, the C4 pathway uses more energy than
the C3 pathway. The C3 pathway requires 18
molecules of ATP for the synthesis of one
molecule of glucose, whereas the C4 pathway
requires 30 molecules of ATP. This energy debt is
more than paid for by avoiding losing more than
half of photosynthetic carbon in photorespiration
as occurs in some tropical plants,[citation needed]
making it an adaptive mechanism for minimizing
the loss.
C4 Leaf Anatomy

• The C4 plants possess a characteristic leaf anatomy.

Their vascular bundles are surrounded by two rings of
cells, the inner ring, called bundle sheath cells, contain
starch-rich chloroplasts lacking grana, which differ from
those in mesophyll cells present as the outer ring. Hence,
the chloroplasts are called dimorphic. This peculiar
anatomy is called kranz anatomy

• The primary function of kranz anatomy is to provide a site

in which CO2 can be concentrated around RuBisCO,
thereby reducing photorespiration
Plants that use C4 carbon fixation
• About 7600 species of plants use C4 carbon fixation,
which represents about 3% of all terresterial species of
plants. All these 7600 species are angiosperms. C4
carbon fixation is less common in dicots than in
monocots, with only 4.5% of dicots using the C4
pathway, compared to 40% of monocots

• Forty-six percent of grasses are C4 and together

account for 61% of C4 species. These include the food
crops maize, sugar cane, millet, and sorghum
CRASULACEAN ACID METABOLISM

Crassulacean acid metabolism, also

known as CAM photosynthesis, is a
carbon fixation pathway present
in some plants. These plants fix carbon
dioxide (CO2) during the night, storing it
as the four-carbon acid malate.

The CO2 is released during the day,

where it is concentrated around the
enzyme RuBisCO, increasing
the efficiency of photosynthesis

Pineapple is the CAM plant

The CAM pathway allows stomata
to remain shut during the day,
reducing evapotranspiration;
therefore, it is especially common
in plants adapted to arid conditions.
During the night
During the night, the CAM plant's stomata are open,
allowing CO2 to enter and be fixated as organic acids that
are stored in vacuoles. During the day the stomata are
closed (thus preventing water loss), and the carbon is
released to the Calvin cycle so that photosynthesis may
take place.

The carbon dioxide is fixed in the mesophyll cell's

cytoplasm by a PEP reaction similar to that of C4 plants.
But, unlike C4 plants, the resulting organic acids are stored
in vacuoles for later use; that is, they are not immediately
passed on to the Calvin cycle. Of course, the latter cannot
operate during night because the light reactions that
provide it with ATP and NADPH cannot take place without
light
During the day

The carbon in the organic acids is freed from the

mesophyll cell's vacuoles and enters the
chloroplast's stroma and, thus, into the Calvin
cycle
 Karbohidrat cadangan

CO2 PEP (phosphoenol piruvat)

Oksaloasetat
Daur Calvin
Malat
Cadangan Malat CO2
KLOROPLAS
Piruvat
VAKUOLA

Daur Calvin
MITOKONDRIA