UPSC MainsBOTANY-PAPER-II202515 Marks
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Q21.

What is biological nitrogen fixation? Explain the mechanism of symbiotic nitrogen fixation with examples.

How to Approach

The answer should begin by defining biological nitrogen fixation and highlighting its significance. The core of the answer will then delve into the detailed mechanism of symbiotic nitrogen fixation, focusing on the Rhizobium-legume association as a prime example. Key stages like nodule formation, the role of the nitrogenase enzyme, and the crucial function of leghemoglobin must be explained comprehensively. Examples of both leguminous and non-leguminous symbiotic fixers should be included.

Model Answer

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Introduction

Nitrogen, though abundant in the atmosphere as inert N<sub>2</sub> gas, is often a limiting nutrient for plant growth as most organisms cannot directly utilize it. Biological Nitrogen Fixation (BNF) is a critical biological process by which certain prokaryotic microorganisms convert atmospheric nitrogen into usable forms, primarily ammonia (NH<sub>3</sub>), which plants can then assimilate for synthesizing vital biomolecules like proteins and nucleic acids. This natural process is indispensable for maintaining ecosystem productivity and soil fertility, offering a sustainable alternative to synthetic nitrogen fertilizers whose excessive use contributes to environmental pollution and greenhouse gas emissions. Among the various types of BNF, symbiotic nitrogen fixation stands out due to its profound agricultural and ecological importance.

What is Biological Nitrogen Fixation?

Biological Nitrogen Fixation (BNF) is the biochemical process mediated by specialized microorganisms (diazotrophs) that transform atmospheric dinitrogen (N2) gas into ammonia (NH3). This conversion is catalyzed by the enzyme complex nitrogenase. The overall reaction is:

N2 + 8H+ + 8e- + 16 ATP → 2NH3 + H2 + 16 ADP + 16 Pi

This process is energetically intensive, requiring a significant amount of ATP, which is supplied by the host plant in symbiotic associations. BNF is a cornerstone of the global nitrogen cycle, making nitrogen available to the biosphere and supporting primary productivity.

Mechanism of Symbiotic Nitrogen Fixation

Symbiotic nitrogen fixation involves a mutualistic relationship between nitrogen-fixing microorganisms and plants, where both partners benefit. The most well-studied and agriculturally significant example is the association between leguminous plants (e.g., peas, soybeans, clover) and bacteria of the genus Rhizobium (collectively known as rhizobia).

Stages of Symbiotic Nitrogen Fixation (Rhizobium-Legume Symbiosis):

  1. Signaling and Attraction:
    • Leguminous roots release chemical signals called flavonoids into the rhizosphere.
    • These flavonoids attract specific Rhizobium bacteria present in the soil.
    • In response, Rhizobium produces Nod factors (Nodulation factors), which are lipo-oligosaccharides, triggering changes in the plant root.
  2. Root Hair Infection and Curling:
    • Nod factors induce curling of the root hair cells of the host plant.
    • The bacteria then attach to the curled root hair and invade it.
    • This leads to the formation of an infection thread, a tubular structure that guides the bacteria through the root hair cell and into the cortical cells of the root.
  3. Nodule Formation:
    • As the infection thread reaches the cortical cells, the bacteria are released into these cells, but remain enclosed within a plant-derived membrane, forming symbiosomes.
    • The infected cortical cells undergo rapid division and enlargement, leading to the formation of specialized structures called root nodules.
    • Within the mature nodule, the rhizobia differentiate into pleomorphic, nitrogen-fixing forms called bacteroids.
  4. Nitrogen Fixation and Ammonia Assimilation:
    • Inside the bacteroids, the nitrogenase enzyme complex converts atmospheric N2 into ammonia (NH3).
    • The nitrogenase enzyme is extremely sensitive to oxygen and is irreversibly inactivated in its presence. To protect nitrogenase, the plant synthesizes a protein called leghemoglobin (leghaemoglobin).
    • Leghemoglobin, similar to hemoglobin in animal blood, binds oxygen, creating a micro-anaerobic environment essential for nitrogenase activity, while still allowing enough oxygen for the bacteroids' respiration (which provides the ATP for fixation).
    • The fixed ammonia (NH3) is rapidly protonated to ammonium (NH4+) and then assimilated by the plant into amino acids (e.g., glutamine, aspartate) and other nitrogenous compounds, which are then transported to other parts of the plant. The plant supplies carbohydrates from photosynthesis to the bacteroids as an energy source.

Examples of Symbiotic Nitrogen Fixation:

Symbiotic nitrogen fixation is not limited to legumes. Other plant-microbe associations also exist.

  • Legume-Rhizobium Symbiosis:
    • Plants: Peas, soybeans, alfalfa, clover, lentils, chickpeas, groundnuts, beans.
    • Bacteria: Rhizobium species (e.g., Rhizobium leguminosarum, Bradyrhizobium japonicum), Mesorhizobium, Azorhizobium.
  • Non-leguminous Plant Symbiosis (Actinorhizal Symbiosis):
    • Some non-leguminous plants form root nodules with the actinomycete bacterium Frankia. These plants are often pioneer species in nutrient-poor soils.
    • Plants: Alder trees (Alnus sp.), Casuarina, Sea Buckthorn (Hippophae), Sweet Gale (Myrica gale).
  • Cyanobacteria Symbiosis:
    • Certain cyanobacteria (formerly blue-green algae), such as Anabaena and Nostoc, form symbiotic associations with plants.
    • Examples: Azolla (a water fern) with Anabaena azollae; Cycads with Nostoc in their coralloid roots; some bryophytes like Anthoceros with Nostoc.

The pink or reddish color of healthy root nodules is due to the presence of leghemoglobin, indicating active nitrogen fixation. This process is crucial for sustainable agriculture, reducing the reliance on synthetic fertilizers and improving soil health.

Conclusion

Biological nitrogen fixation, particularly symbiotic nitrogen fixation, is a cornerstone of global food security and ecological balance. By converting atmospheric nitrogen into bioavailable forms, it significantly enhances soil fertility and plant productivity. The intricate mechanism involving host plant signals, microbial responses, nodule formation, and the oxygen-scavenging action of leghemoglobin highlights a remarkable evolutionary collaboration. Promoting and optimizing these natural processes, through practices like crop rotation and biofertilizer use, can substantially reduce the environmental footprint of agriculture, contributing to a more sustainable and resilient food system for 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

Diazotrophs
Microorganisms capable of fixing atmospheric nitrogen into ammonia, including bacteria and archaea. Examples include <em>Rhizobium</em>, <em>Azotobacter</em>, and various cyanobacteria.
Nitrogenase
An enzyme complex found in diazotrophs that catalyzes the conversion of atmospheric dinitrogen (N<sub>2</sub>) to ammonia (NH<sub>3</sub>). It is highly sensitive to oxygen and requires a large amount of energy (ATP) to function.

Key Statistics

India's nitrogen fertilizer consumption was approximately 19.43 million tons in 2021-2022. Urea accounts for about 82% of total nitrogen consumption in the country.

Source: FAI (Fertiliser Association of India), 2020; ResearchGate, 2021-2022 data

Globally, biological nitrogen fixation contributes an estimated 100-200 million metric tons of fixed nitrogen per year, which is comparable to or even exceeds industrial nitrogen fixation.

Source: Various ecological and agricultural studies

Examples

Legume Inoculation

Farmers often inoculate legume seeds (e.g., soybean, groundnut) with specific strains of <em>Rhizobium</em> bacteria before planting. This practice ensures the presence of effective nitrogen-fixing symbionts in the soil, leading to improved nodulation, enhanced nitrogen fixation, and higher crop yields without excessive chemical nitrogen fertilizers.

Rice-Azolla Symbiosis

In many Asian countries, the aquatic fern <em>Azolla</em> is used as a biofertilizer in rice paddies. <em>Azolla</em> harbors nitrogen-fixing cyanobacteria (<em>Anabaena azollae</em>) in its fronds. When <em>Azolla</em> grows in rice fields, it fixes atmospheric nitrogen, which becomes available to the rice plants upon <em>Azolla</em>'s decomposition, significantly reducing the need for synthetic nitrogen fertilizers for rice cultivation.

Frequently Asked Questions

Why is leghemoglobin essential for symbiotic nitrogen fixation?

Leghemoglobin is crucial because the nitrogenase enzyme, which catalyzes nitrogen fixation, is highly sensitive to oxygen and gets inactivated in its presence. Leghemoglobin acts as an oxygen scavenger, maintaining a very low (micro-anaerobic) oxygen concentration within the root nodule, thereby protecting nitrogenase while still allowing sufficient oxygen for the high respiratory demands of the bacteria to produce ATP for the energy-intensive fixation process.

Topics Covered

BiologyBotanyAgricultureEcologyNitrogen fixationSymbiosisMicrobiologyPlant nutrition