UPSC MainsBOTANY-PAPER-II202420 Marks
Q25.

Explain the compartmentation of biochemical reactions in photorespiration. Comment upon the significance of the process.

How to Approach

This question requires a detailed understanding of photorespiration, focusing not just on the process itself, but specifically *how* it's spatially organized within the plant cell. The answer should explain the involvement of chloroplasts, peroxisomes, and mitochondria, detailing the reactions occurring in each organelle. The significance should be discussed in terms of its evolutionary context, its drawbacks, and potential adaptations plants have developed to mitigate its effects. A clear, step-by-step explanation with relevant biochemical details is crucial.

Model Answer

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Introduction

Photorespiration is a metabolic pathway that occurs in plants alongside photosynthesis, particularly under conditions of high oxygen and low carbon dioxide. Often described as a wasteful process, it involves the oxygenation of RuBP by the enzyme RuBisCO, instead of its carboxylation. This initiates a complex series of reactions spanning three different organelles – chloroplasts, peroxisomes, and mitochondria – demonstrating a remarkable example of biochemical compartmentation. Understanding this compartmentalization is key to appreciating the intricacies of plant metabolism and its evolutionary history.

Compartmentation of Biochemical Reactions in Photorespiration

Photorespiration isn't a single reaction but a cycle of reactions distributed across three organelles. This compartmentalization is crucial for efficient functioning and preventing interference with other metabolic processes.

1. Chloroplasts: The Initiation Phase

  • The process begins in the chloroplast when RuBisCO oxygenates RuBP (Ribulose-1,5-bisphosphate) instead of carboxylating it.
  • This yields one molecule of 3-PGA (3-phosphoglycerate) – which enters the Calvin cycle – and one molecule of 2-phosphoglycolate.
  • 2-phosphoglycolate is toxic and cannot be directly used by the plant. It is rapidly converted to glycolate.
  • Glycolate is then transported to the peroxisome.

2. Peroxisomes: The Major Processing Center

  • In the peroxisome, glycolate is oxidized to glyoxylate by glycolate oxidase, producing hydrogen peroxide (H2O2).
  • Hydrogen peroxide is then broken down into water and oxygen by catalase.
  • Glyoxylate is transaminated to glycine.
  • Two molecules of glycine are transported to the mitochondria.

3. Mitochondria: The Release of CO2 and Nitrogen

  • Within the mitochondrial matrix, two molecules of glycine undergo a complex reaction catalyzed by the glycine decarboxylase complex (GDC).
  • This reaction releases one molecule of CO2, one molecule of ammonia (NH3), and one molecule of serine.
  • The released ammonia is re-assimilated into glutamate via the glutamine synthetase/glutamate synthase (GS/GOGAT) pathway, requiring ATP and reducing power.
  • Serine is transported back to the peroxisome.

4. Return to Peroxisome and Chloroplast

  • In the peroxisome, serine is converted to hydroxypyruvate.
  • Hydroxypyruvate is reduced to glycerate.
  • Glycerate is transported back to the chloroplast.
  • In the chloroplast, glycerate is phosphorylated to 3-PGA, re-entering the Calvin cycle.

This cyclical process, involving the transport of metabolites between organelles, highlights the importance of compartmentation in managing the potentially harmful byproducts of photorespiration and ultimately recovering some of the carbon lost.

Significance of Photorespiration

While often considered a wasteful process, photorespiration has several important implications:

  • Evolutionary Relic: It is believed that photorespiration evolved when atmospheric CO2 levels were much higher and O2 levels were lower. RuBisCO, in that environment, was more efficient at carboxylation. As O2 levels rose, the oxygenase activity of RuBisCO became significant.
  • Photoprotection: Photorespiration can act as a protective mechanism against photoinhibition, preventing damage to the photosynthetic apparatus under high light intensity. By consuming excess light energy, it dissipates energy that could otherwise lead to the formation of reactive oxygen species (ROS).
  • Nitrogen Metabolism: The ammonia released during glycine decarboxylation is reassimilated, contributing to the plant’s nitrogen metabolism.
  • Metabolic Intermediates: Photorespiration generates metabolic intermediates that can be used in other biosynthetic pathways.

However, the energetic cost of photorespiration is substantial. It consumes ATP and reducing power, and releases CO2, reducing the net efficiency of photosynthesis. C4 and CAM plants have evolved mechanisms to minimize photorespiration by concentrating CO2 around RuBisCO, thereby favoring carboxylation over oxygenation.

Conclusion

In conclusion, photorespiration is a complex metabolic pathway intricately compartmentalized across chloroplasts, peroxisomes, and mitochondria. While energetically costly, it likely played a crucial role in the evolution of photosynthesis and continues to offer some photoprotective benefits. The development of C<sub>4</sub> and CAM pathways demonstrates the evolutionary pressure to overcome the limitations of photorespiration, highlighting the ongoing adaptation of plants to changing environmental conditions. Further research into optimizing photosynthetic efficiency and minimizing photorespiration remains a key area in plant biology.

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, the enzyme responsible for carbon fixation in photosynthesis, but also catalyzes the oxygenation of RuBP, initiating photorespiration.
Photoinhibition
Damage to the photosynthetic apparatus caused by excessive light energy, leading to reduced photosynthetic efficiency. Photorespiration can help mitigate photoinhibition.

Key Statistics

Photorespiration can reduce photosynthetic efficiency by up to 25-50% in C3 plants.

Source: Taiz & Zeiger, Plant Physiology and Development (2010)

Globally, approximately 3-16% of carbon fixed by photosynthesis is lost due to photorespiration.

Source: Sage, R. F. (2004). The evolution of C4 photosynthesis. New Phytologist, 164(2), 317–338.

Examples

C4 Plants

Maize and sugarcane are C4 plants. They spatially separate initial CO<sub>2</sub> fixation from the Calvin cycle, concentrating CO<sub>2</sub> around RuBisCO in bundle sheath cells, thus minimizing photorespiration.

Frequently Asked Questions

Why is photorespiration considered wasteful?

Photorespiration is considered wasteful because it consumes ATP and NADPH, releases CO<sub>2</sub>, and does not produce sugars directly. It effectively reverses some of the work done by photosynthesis.