UPSC MainsBOTANY-PAPER-II202015 Marks
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Q21.

Carboxylation and oxygenation compete to decrease the efficiency of photosynthesis. Discuss.

How to Approach

This question requires a detailed understanding of the C3 and C4 photosynthetic pathways, and photorespiration. The answer should explain the biochemical processes of carboxylation and oxygenation, highlighting why they compete and how this competition impacts photosynthetic efficiency. Structure the answer by first defining the processes, then explaining the competition, its consequences, and finally, adaptations plants have evolved to mitigate this issue. Focus on the enzyme RuBisCO and its dual affinity for CO2 and O2.

Model Answer

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Introduction

Photosynthesis, the cornerstone of life on Earth, is the process by which plants convert light energy into chemical energy. However, the efficiency of this process isn’t absolute. A significant limitation arises from the inherent competition between carboxylation and oxygenation reactions within the Calvin cycle. This competition, mediated by the enzyme RuBisCO, reduces the overall efficiency of carbon fixation, particularly under certain environmental conditions. Understanding this interplay is crucial for comprehending plant productivity and adaptation strategies.

Understanding Carboxylation and Oxygenation

Carboxylation is the initial step of the Calvin cycle, where carbon dioxide (CO2) is fixed by the enzyme Ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBisCO) to Ribulose-1,5-bisphosphate (RuBP), forming two molecules of 3-phosphoglycerate (3-PGA). This is the desired reaction, leading to sugar production.

Oxygenation, on the other hand, occurs when RuBisCO binds to oxygen (O2) instead of CO2. This results in the formation of one molecule of 3-PGA and one molecule of phosphoglycolate. Phosphoglycolate is a toxic compound that needs to be salvaged through a costly process called photorespiration.

The Competition: RuBisCO’s Dual Affinity

The competition between carboxylation and oxygenation stems from RuBisCO’s inherent properties. RuBisCO isn’t perfectly specific for CO2; it also has an affinity for O2. The relative rates of carboxylation and oxygenation depend on the concentrations of CO2 and O2, as well as temperature and light intensity.

Specifically, the ratio of carboxylation to oxygenation is influenced by:

  • CO2 Concentration: Higher CO2 concentrations favor carboxylation.
  • O2 Concentration: Higher O2 concentrations favor oxygenation.
  • Temperature: Higher temperatures increase the rate of oxygenation more than carboxylation, as the affinity of RuBisCO for O2 increases with temperature.
  • Light Intensity: High light intensity can lead to increased O2 production during the light-dependent reactions of photosynthesis, potentially increasing oxygenation.

Consequences of Oxygenation: Photorespiration

When oxygenation occurs, the resulting phosphoglycolate is metabolized through a complex pathway called photorespiration. This process involves the chloroplast, peroxisome, and mitochondria, and it consumes ATP and NADPH without producing any sugar. In fact, it releases CO2, effectively reversing some of the carbon fixation achieved during carboxylation.

Photorespiration can reduce photosynthetic efficiency by as much as 25-50% in C3 plants under hot, dry conditions. This is because stomata close to conserve water, reducing CO2 entry and increasing O2 concentration inside the leaves.

Adaptations to Minimize Photorespiration

Plants have evolved various adaptations to minimize photorespiration and enhance photosynthetic efficiency:

  • C4 Photosynthesis: C4 plants (e.g., maize, sugarcane) spatially separate initial CO2 fixation from the Calvin cycle. They use PEP carboxylase to fix CO2 into a four-carbon compound in mesophyll cells, which is then transported to bundle sheath cells where the Calvin cycle occurs. This concentrates CO2 around RuBisCO, minimizing oxygenation.
  • CAM Photosynthesis: CAM plants (e.g., cacti, succulents) temporally separate CO2 fixation from the Calvin cycle. They open their stomata at night to fix CO2 into organic acids, which are stored in vacuoles. During the day, stomata close to conserve water, and the organic acids release CO2 for use in the Calvin cycle.
  • RuBisCO Modifications: Research is ongoing to engineer RuBisCO with higher specificity for CO2 and lower affinity for O2.
Feature C3 Plants C4 Plants CAM Plants
Initial CO2 Fixation RuBisCO PEP Carboxylase PEP Carboxylase (night)
Spatial Separation No Yes (Mesophyll & Bundle Sheath) No
Temporal Separation No No Yes (Night & Day)
Photorespiration High Low Very Low

Conclusion

The competition between carboxylation and oxygenation, driven by RuBisCO’s dual affinity, represents a fundamental limitation to photosynthetic efficiency. Photorespiration, a consequence of oxygenation, reduces carbon fixation and plant productivity. However, plants have evolved remarkable adaptations, such as C4 and CAM photosynthesis, to overcome this challenge and thrive in diverse environments. Ongoing research into RuBisCO engineering holds promise for further enhancing photosynthetic efficiency and improving crop yields in the future.

Answer Length

This is a comprehensive model answer for learning purposes and may exceed the word limit. In the exam, always adhere to the prescribed word count.

Additional Resources

Key Definitions

RuBisCO
Ribulose-1,5-bisphosphate carboxylase/oxygenase, an enzyme responsible for the initial fixation of carbon dioxide in the Calvin cycle. It is the most abundant protein on Earth.

Key Statistics

Globally, photorespiration is estimated to reduce potential crop yields by up to 37% (estimated based on knowledge cutoff 2023).

Source: Long, S. P. (2013). Intercropping and the synergy of ecological processes. Frontiers in Ecology and the Environment, 11(1), 45–54.

C4 plants account for approximately 3-5% of plant species globally, but contribute over 20% of global net primary production (estimated based on knowledge cutoff 2023).

Source: Sage, R. F. (2004). The evolution of C4 photosynthesis. New Phytologist, 161(1), 35–59.

Examples

Maize (C4 Plant)

Maize, a C4 plant, exhibits higher photosynthetic rates and water-use efficiency compared to C3 plants like wheat, especially in hot and dry climates. This is due to its ability to concentrate CO2 around RuBisCO, minimizing photorespiration.

Frequently Asked Questions

Can photorespiration be completely eliminated?

Completely eliminating photorespiration is challenging due to RuBisCO’s inherent properties. However, genetic engineering efforts are focused on reducing its occurrence and improving RuBisCO’s specificity for CO2.

Topics Covered

BotanyPlant SciencePhysiologyPhotosynthesisPlant MetabolismC3 C4 Plants