RESPIRATION IN PLANTS

Introduction

  • Life Processes and Energy: All living organisms require energy for essential life processes.
  • Energy Sources: Energy is obtained through the oxidation of macromolecules known as “food.”

Photosynthesis and Food Production

  • Green Plants and Cyanobacteria: Capable of photosynthesis, converting light energy into chemical energy stored in carbohydrates (glucose, sucrose, starch).
  • Food Translocation: Not all cells in green plants photosynthesize; food must be translocated to non-green parts.

Heterotrophic Organisms

  • Animal and Fungal Nutrition: Animals (herbivores, carnivores) obtain food directly or indirectly from plants. Fungi are saprophytes, relying on dead and decaying matter.
  • Ultimate Source: Regardless of the organism’s nutritional strategy, the ultimate source of food for respiration is photosynthesis.

Cellular Respiration Process

  • Cellular Locations: Photosynthesis occurs in chloroplasts, while cellular respiration takes place in the cytoplasm and mitochondria (eukaryotes).
  • C-C Bond Breakdown: Respiration involves the breakdown of C-C bonds through oxidation, releasing energy.
  • Respiratory Substrates: Carbohydrates are commonly oxidized, but proteins, fats, and organic acids may also serve as respiratory substrates in specific conditions.

ATP Synthesis and Utilization

  • Series of Reactions: Energy release in respiration occurs in a series of controlled, enzymatic reactions.
  • ATP Synthesis: Energy released is trapped as chemical energy in the form of ATP.
  • Energy Currency: ATP serves as the energy currency of the cell, utilized in various energy-requiring processes.
  • Carbon Skeleton Utilization: Carbon skeletons produced during respiration serve as precursors for the biosynthesis of other molecules in the cell.
  • Photosynthesis Adaptations: During photosynthesis, O2 release within cells addresses O2 availability.

Gaseous Exchange Mechanisms

  • Cell Proximity to Surface: Living cells in plants are close to the surface, facilitating efficient gas exchange.
  • Parenchyma Cells: Loose packing of parenchyma cells in leaves, stems, and roots creates interconnected air spaces.

Energy Production and Utilization

  • Complete Glucose Combustion: Glucose combustion produces CO2, H2O, and energy (mostly as heat).
  • Energy Utilization Strategy: Cells catabolize glucose in multiple small steps, coupling energy release to ATP synthesis.
  • Story of Respiration: Respiration involves oxygen utilization, releasing CO2, water, and energy.

Anaerobic Adaptations

  • Oxygen Availability: Some cells live in environments with varying oxygen availability.
  • Adaptation to Anaerobic Conditions: Early cells likely existed in oxygen-lacking atmospheres; present-day organisms adapted to anaerobic conditions.

Glycolysis: Anaerobic Glucose Breakdown

  • Enzymatic Machinery: Living organisms retain enzymatic machinery for partial glucose oxidation without oxygen.
  • Glycolysis: Breakdown of glucose to pyruvic acid in the absence of oxygen.

Glycolysis: Breaking Down Glucose for Energy

Introduction

  • Origin of the Term: “Glycolysis” from Greek words glycos (sugar) and lysis (splitting).
  • Pioneers: Gustav Embden, Otto Meyerhof, and J. Parnas developed the glycolysis scheme.
  • Anaerobic Process: Predominant in anaerobic organisms; occurs in the cytoplasm.

Glycolytic Pathway in Plants

  • Glucose Source: Derived from sucrose (photosynthesis end product) or storage carbohydrates.
  • Sucrose Conversion: Invertase converts sucrose into glucose and fructose.
  • Initial Steps: Glucose and fructose phosphorylation to form glucose-6-phosphate.

Ten Steps of Glycolysis

  1. Hexokinase Activity: Glucose to glucose-6-phosphate.
  2. Isomerization: Glucose-6-phosphate to fructose-6-phosphate.
  3. Common Metabolism: Subsequent metabolism steps for glucose and fructose are the same.
  4. ATP Utilization: Twice during glucose conversion and fructose-6-phosphate conversion.
  5. Fructose 1,6-Bisphosphate Formation: ATP consumption.
  6. Bisphosphate Split: Fructose 1,6-bisphosphate to dihydroxyacetone phosphate and 3-phosphoglyceraldehyde.

NADH + H+ Formation

  • Redox Reaction: 3-phosphoglyceraldehyde to 1, 3-bisphosphoglycerate.
  • Hydrogen Transfer: NADH + H+ formation.

ATP Synthesis

  • Energy Yielding Steps: Conversion of 1, 3-bisphosphoglycerate to 3-phosphoglyceric acid.
  • ATP Formation: During the conversion of PEP to pyruvic acid.

Overall ATP Calculation

  • Direct ATP Synthesis: Calculate the total ATP molecules directly synthesized from one glucose molecule.

Pyruvic Acid as a Product

  • Metabolic Fate: Determined by cellular needs.
  • Possibilities: Lactic acid fermentation, alcoholic fermentation, or aerobic respiration.

Fermentation: Anaerobic Energy Production

Overview

  • Definition: Incomplete oxidation of glucose under anaerobic conditions.
  • Byproducts: CO2 and ethanol (yeast), or lactic acid (bacteria and muscles).
  • Enzymes: Pyruvic acid decarboxylase and alcohol dehydrogenase (for alcohol fermentation).

Yeast Fermentation

  1. Pyruvic Acid Conversion: Catalyzed by pyruvic acid decarboxylase.
  2. Ethanol Formation: Through alcohol dehydrogenase.
  3. Energy Release: Limited energy release (less than 7% of glucose energy).
  4. ATP Synthesis: Not highly efficient; some ATP formed.

Lactic Acid Fermentation

  • Bacterial Process: Some bacteria convert pyruvic acid to lactic acid.
  • Animal Cells: Muscles during anaerobic conditions convert pyruvic acid to lactic acid.
  • NADH+H+ Reoxidation: NAD+ regeneration via reduction of pyruvic acid.

Energy Yield and Hazards

  • Limited ATP Production: Not all energy is trapped as high-energy ATP bonds.
  • Hazardous Byproducts: Production of acid (lactic acid) or alcohol.
  • Yeast Poisoning: Yeasts are affected when alcohol concentration reaches about 13%.

Net ATP Synthesis

  • Calculation: Deduct ATP used during glycolysis from ATP synthesized during fermentation.

Maximum Alcohol Concentration

  • Yeast Limitation: Yeasts poison themselves at high alcohol concentrations.
  • Natural Fermentation: Maximum alcohol content limited by yeast tolerance.

Complete Oxidation in Aerobic Respiration

  • Mitochondrial Steps: Complete oxidation occurs within mitochondria.
  • Oxygen Dependency: Requires oxygen (O2).
  • Products: CO2, water, and substantial energy release for ATP synthesis.
  • Prevalence: Common in higher organisms.

Aerobic Respiration: Mitochondrial Processes

Overview

  • Location: Mitochondria (matrix and inner membrane).
  • Pyruvate Transport: From cytoplasm to mitochondria.
  • Crucial Events:
    1. Oxidation of Pyruvate:
      • Removal of all hydrogen atoms.
      • Production of three molecules of CO2.
      • Occurs in the matrix of mitochondria.
    2. Electron Transfer to O2:
      • Electrons passed to molecular O2.
      • Simultaneous ATP synthesis.
      • Located on the inner membrane of mitochondria.

Pyruvate Oxidation

  1. Pyruvate Formation: Glycolytic catabolism in cytosol produces pyruvate.
  2. Transport to Mitochondria: Pyruvate enters mitochondrial matrix.
  3. Oxidative Decarboxylation: Catalyzed by pyruvic dehydrogenase.
  4. Coenzymes: Involves NAD+ and Coenzyme A.
  5. Reaction:

NADH Production

  • NADH Synthesis: Two molecules of NADH from the metabolism of two pyruvic acid molecules (from one glucose during glycolysis).

Tricarboxylic Acid Cycle (TCA Cycle)

Overview

  • Start: Condensation of acetyl group with oxaloacetic acid (OAA) and water.
  • Enzyme: Catalyzed by citrate synthase.
  • Product: Citric acid and release of CoA.
  • Isomerization: Citrate is isomerized to isocitrate.
  • Decarboxylation Steps:
    1. Formation of α-ketoglutaric acid.
    2. Formation of succinyl-CoA.
  • Oxidation Steps:
    • Succinyl-CoA to succinic acid.
    • GTP synthesis in substrate-level phosphorylation.
    • GTP converted to GDP, producing ATP.
  • Reduction Steps:
    • Three points where NAD+ is reduced to NADH + H+.
    • One point where FAD+ is reduced to FADH2.
  • Continued Cycle: Requires replenishment of oxaloacetic acid and regeneration of NAD+ and FAD+.

Substrate-Level Phosphorylation

  • Process: Conversion of succinyl-CoA to succinic acid.
  • Result: Synthesis of one molecule of GTP.
  • Coupled Reaction: GTP converted to GDP, simultaneously synthesizing ATP from ADP.

NADH and FADH2 Production

  • NADH Production: Three points in the cycle.
  • FADH2 Production: One point in the cycle.

Cycle Significance

  • Purpose: Continued oxidation of acetyl CoA.
  • Requirements:
    1. Replenishment of oxaloacetic acid.
    2. Regeneration of NAD+ and FAD+ from NADH and FADH2.
  • Products So Far: CO2, eight NADH + H+, two FADH2, and two ATP.

Electron Transport System (ETS) and Oxidative Phosphorylation

Overview

  • Objective: Release and utilize the energy stored in NADH+H+ and FADH2.
  • Location: Inner mitochondrial membrane.
  • Process: Oxidation of NADH and FADH2 through the electron transport system (ETS).
  • Result: Formation of H2O, coupled with ATP synthesis.

Electron Transport System (ETS) Components

  1. NADH Dehydrogenase (Complex I):
    • Oxidizes electrons from NADH produced in the mitochondrial matrix.
    • Transfers electrons to ubiquinone.
  2. Ubiquinone:
    • Receives electrons from NADH dehydrogenase (complex I) and FADH2 (complex II).
    • Transfers electrons to cytochrome c.
  3. Cytochrome c and Cytochrome bc1 Complex (Complex III):
    • Cytochrome c acts as a mobile carrier.
    • Receives electrons from reduced ubiquinone.
    • Transfers electrons to cytochrome c oxidase complex (complex IV).
  4. Cytochrome c Oxidase Complex (Complex IV):
    • Contains cytochromes a and a3, and two copper centers.
    • Accepts electrons from cytochrome c.
    • Transfers electrons to oxygen, forming H2O.

ATP Synthesis

  • Coupled with ETS: ATP synthase (complex V) couples with the electron transport chain (ETS).
  • Components:
    • F1 Headpiece: Peripheral membrane protein complex with ATP synthesis site.
    • F0 Integral Membrane Protein: Forms a channel for proton movement.
  • Proton Movement: Protons move through F0 from the intermembrane space to the matrix down the electrochemical proton gradient.
  • Proton Passage Coupling: Coupled to the catalytic site of the F1 component for ATP production.
  • ATP Synthesis: For each ATP produced, 4H+ passes through F0.

Role of Oxygen

  • Terminal Stage: Oxygen acts as the final hydrogen acceptor.
  • Importance: Drives the entire process by removing hydrogen from the system.
  • Role: Oxygen is the final electron acceptor, forming water.

Oxidative Phosphorylation

  • Definition: The process is termed oxidative phosphorylation due to the utilization of energy from oxidation-reduction for ATP synthesis.
  • Comparison to Photophosphorylation: Unlike photophosphorylation, where light energy is utilized, oxidative phosphorylation uses the energy from oxidation-reduction.

ATP Synthase (Complex V)

  • Components:
    • F1 Headpiece: Peripheral membrane protein complex.
    • F0 Integral Membrane Protein: Forms the proton channel.
  • Energy Source: Energy released during the electron transport system.
  • Function: Synthesizes ATP from ADP and inorganic phosphate.
  • Hypothesis: Mechanism explained by the chemiosmotic hypothesis.

ATP Production

  • NADH Oxidation: Produces 3 molecules of ATP.
  • FADH2 Oxidation: Produces 2 molecules of ATP.

Importance of Oxygen in Respiration

  • Limited Role: Oxygen’s role is primarily in the terminal stage of respiration.
  • Vital Presence: Vital for driving the entire process by acting as the final hydrogen acceptor.

THE RESPIRATORY BALANCE SHEET

Overview

  • Objective: Calculate the theoretical net gain of ATP for every glucose molecule oxidized.
  • Assumptions:
    1. Sequential, orderly pathway: Glycolysis, TCA cycle, and ETS pathway in succession.
    2. Transfer of NADH to mitochondria for oxidative phosphorylation.
    3. No utilization of intermediates for synthesizing other compounds.
    4. Sole respiration of glucose; no alternative substrates entering the pathway.
  • Limitations: Real-life pathways are simultaneous, substrate entry/withdrawal dynamic, and enzymatic rates are controlled by multiple factors.

Theoretical Net Gain

  • Aerobic Respiration:
    • Assumptions-Based Net Gain: 38 ATP molecules for one glucose molecule.
    • Considerations: Sequential functioning, controlled substrate flow, and no diversion of intermediates.

Comparison with Fermentation

  • Fermentation vs. Aerobic Respiration:
    • Partial vs. Complete Breakdown:
      • Fermentation: Partial breakdown of glucose.
      • Aerobic Respiration: Complete degradation to CO2 and H2O.
    • ATP Production:
      • Fermentation: Net gain of only two ATP molecules per glucose degraded to pyruvic acid.
      • Aerobic Respiration: Generates many more ATP molecules under aerobic conditions.
    • NADH Oxidation:
      • Fermentation: Slow oxidation of NADH to NAD+.
      • Aerobic Respiration: Vigorous oxidation of NADH to NAD+.

Appreciating the Living System

  • Dynamic Pathways:
    • Pathways work simultaneously, not sequentially.
    • Substrates enter/exit pathways dynamically.
  • Utilization of ATP:
    • ATP is used as needed, and enzymatic rates are controlled by multiple factors.
  • Exercise Purpose:
    • Appreciate the efficiency of the living system in energy extraction and storage.

AMPHIBOLIC PATHWAY

Overview

  • Glucose as the Primary Substrate:
    • Glucose is the preferred substrate for respiration.
    • All carbohydrates are usually converted into glucose before entering the respiratory pathway.

Entry Points for Different Substrates

  • Points of Entry:
    • Glucose: Primary substrate entering the respiratory pathway.
    • Fats: Break down into glycerol and fatty acids, with entry after degradation to acetyl CoA.
    • Proteins: Degraded by proteases, and amino acids enter the pathway at various stages (as pyruvate, acetyl CoA, or within Krebs’ cycle).

Catabolism vs. Anabolism

  • Traditional Understanding:
    • Respiration seen as a catabolic process, and the respiratory pathway as catabolic.
    • Catabolism involves the breakdown of substrates.

Amphibolic Nature of the Respiratory Pathway

  • Dual Role in Breakdown and Synthesis:
    • Fatty Acids:
      • Breakdown to acetyl CoA during respiration.
      • Acetyl CoA is withdrawn for fatty acid synthesis.
    • Proteins:
      • Involved in both breakdown and synthesis.
    • Recognition of Amphibolic Nature:
      • The respiratory pathway plays a role in both catabolism and anabolism.
      • Recognized as an amphibolic pathway.

RESPIRATORY QUOTIENT (RQ)

Overview

  • Definition:
    • The respiratory quotient (RQ) or respiratory ratio is the ratio of the volume of CO2 evolved to the volume of O2 consumed during aerobic respiration.

Calculation of RQ

  • Formula:
    • RQ=Volume of CO2 evolved Volume of O2 consumed RQ =Volume of O2​ consumed volume of CO2​ evolved​

Dependence on Respiratory Substrate

  • Carbohydrates:
    • RQ is 1 when carbohydrates are completely oxidized.
    • Example: C6H12O6+6O2→6CO2+6H2O+Energy
    • RQ=6CO2/6O2=1
  • Fats:
    • RQ is less than 1 when fats are used.
    • Example (Tripalmitin): C51H98O6​+145O2​→102CO2​+98H2O+Energy
    • RQ=102CO2​​/145O2≈0.7
  • Proteins:
    • RQ for proteins is about 0.9.

Importance

  • Multiple Substrates:
    • Living organisms often use a combination of respiratory substrates.
    • Pure proteins or fats are rarely used as sole respiratory substrates.