24 Calvin cycle & PPP

The Calvin Cycle and Pentose Phosphate Pathway

Overview of Light Reactions and Carbon Reactions

Light Reactions
  • Definition: The light reactions of photosynthesis require visible light as an energy source.

  • Products: The light reactions produce:

    • Reducing power in the form of NADPH

    • ATP

    • Oxygen (O2)

  • Chemical Equation:
    2 ext{ H}2 ext{O} + 2 ext{ NADP}^+ + 3 ext{ ADP} + 3 ext{ Pi} + ext{ light} ightarrow 2 ext{ NADPH} + 2 ext{ H}^+ + 3 ext{ ATP} + ext{ O}2

Carbon Reactions
  • Function: The carbon reactions, also called dark reactions or light-independent reactions, utilize ATP and NADPH to fix carbon dioxide (CO2) into carbohydrates.

  • Chemical Equation:
    3 ext{ CO}2 + 9 ext{ ATP} + 6 ext{ NADPH} + 6 ext{ H}^+ ightarrow ext{ C}3 ext{H}6 ext{O}3- ext{phosphate} + 9 ext{ ADP} + 8 ext{ Pi} + 6 ext{ NADP}^+ + 3 ext{ H}_2 ext{O}

  • Location of Reactions: Both NADPH and ATP are produced in the stroma of chloroplasts, which is where they are used in the Calvin Cycle.

Calvin Cycle Phases
  • Phases of the Calvin Cycle: The Calvin cycle consists of three main phases:

    1. Carboxylation: Conversion of CO2 to 3-phosphoglycerate (3PG)

    2. Reduction: Transformation of 3PG to glyceraldehyde-3-phosphate (GAP)

    3. Regeneration of Starting Material: Conversion of GAP back to ribulose 1,5-bisphosphate (RuBP)

  • Historical Context: Originally termed "dark reactions" because they do not require light. The Calvin cycle centers around building sugar from CO2 and is a reductive process, necessitating electrons provided by NADPH.

  • Significance: Melvin Calvin was awarded the Nobel Prize in 1961 for elucidating this cycle.

Detailed Phases of the Calvin Cycle

Phase 1: Carboxylation
  • Enzyme Involved: RuBisCO (Ribulose Bisphosphate Carboxylase/Oxygenase)

    • Function: Attaches CO2 to ribulose 1,5-bisphosphate (RuBP) converting the gas into carbohydrate.

    • Characteristics: Considered "slow" with a turnover rate of approximately 3 reactions per second, and can mistakenly attach O2 instead of CO2.

    • Most abundant protein on Earth, making up about 40 million tons or 50% of leaf protein.

    • It is termed the "cornerstone of autotrophy" because it catalyzes the fixation of CO2 into organic molecules.

Phase 2: Reduction
  • Process: Involves the reduction of 3-phosphoglycerate (3PG) to glyceraldehyde-3-phosphate (GAP).

  • Chemical Mechanism: The reduction process represents the reverse of two reactions in glycolysis:

Key Points about Reduction:
  • Cleavage: Cleavage of the acyl-phosphate bond drives reduction.

  • Redox Pair: NADPH/NADP+ are the redox pairs utilized in this reaction, in contrast to glycolysis which uses NADH/NAD+.

    • Cycle Turn: For every 3 turns of the Calvin Cycle using 3 CO2s, one GAP is siphoned off for net carbohydrate synthesis.

Phase 3: Regeneration
  • Purpose: Regenerates the CO2 acceptor, RuBP.

  • Requirements: Conversion of five 3C GAPs into three 5C RuBPs is essential to maintain cycle continuity.

  • In Reactions: Some reactions exchange carbons while others involve hydrolysis or phosphorylation, supported by specific enzymes:

    • Aldolase: Catalyzes the combination of three-carbon aldose and ketose into a six-carbon carbohydrate.

    • Transketolase: Transfers two carbon units, crucial for moving carbons around during the regeneration phase.

    • Bisphosphatases: Catalyze the hydrolysis of phosphates in some of the reactions.

    • Kinase: Adds phosphates to important substrates.

    • Isomerases and Epimerases: Facilitate structural rearrangements within molecules.

Regulated Steps in the Calvin Cycle
  • Key Regulatory Steps: Rubisco and kinase/phosphatases are the points of regulation within the Calvin cycle. The cycle is activated by light, ensuring coordination between light and dark reactions.

  • Reversibility: All regulated enzymatic reactions are irreversible, while other reactions can be reversible.

Energy Requirements
  • During the cycle, a total of nine ATPs and six NADPHs are utilized for each three turns.

  • One GAP is used for net carbohydrate synthesis while five are directed towards regenerating RuBP.

Key Participants in the Pathway
  • Final Products of Calvin Cycle: 2 GAPs from two turns are used to synthesize hexoses, which may then be further processed into starch or sucrose.