Biology: Class 11 — Chapter 12: Respiration in Plants

Introduction to Cellular Respiration

All living organisms require energy to perform life activities such as absorption, transport, movement, reproduction, and breathing. This energy is obtained through the oxidation of macromolecules, collectively termed as "food".

12.1 Do Plants Breathe?

Plants require $O_2$ for respiration and release $CO_2$, but they do not possess specialized respiratory organs like animals. Instead, they rely on stomata and lenticels for gaseous exchange.

Reasons Plants Can Function Without Respiratory Organs

  1. Independence of Parts: Each plant part takes care of its own gas-exchange needs. There is very little transport of gases from one part to another.
  2. Low Gas Exchange Demands: Roots, stems, and leaves respire at rates far lower than animals. Large volumes of gas exchange occur only during photosynthesis, where $O_2$ is released internally within the photosynthesising cells.
  3. Short Diffusion Distance: Most living cells in a plant are located close to the surface. In thick woody stems, living cells are organized in thin layers beneath the bark (with lenticels), while the interior cells are dead and provide only mechanical support.

The complete combustion of glucose yields $CO_2$, $H_2O$, and energy (mostly as heat):

$$C_6H_{12}O_6 + 6O_2 \rightarrow 6CO_2 + 6H_2O + \text{Energy}$$
12.2 Glycolysis

The term glycolysis originates from the Greek words glycos (sugar) and lysis (splitting). It is also known as the EMP pathway, named after Gustav Embden, Otto Meyerhof, and J. Parnas.

Key Steps of Glycolysis

  1. Phosphorylation of Glucose: Glucose is phosphorylated to Glucose-6-phosphate by the enzyme hexokinase, consuming 1 ATP.
  2. Isomerisation: Glucose-6-phosphate isomerises to Fructose-6-phosphate.
  3. Second Phosphorylation: Fructose-6-phosphate is phosphorylated to Fructose-1,6-bisphosphate, consuming another ATP.
  4. Splitting: Fructose-1,6-bisphosphate is split into two 3-carbon molecules: Triose phosphate (PGAL - 3-phosphoglyceraldehyde) and Dihydroxyacetone phosphate (which are interconvertible).
  5. Oxidation and Phosphorylation: PGAL is oxidised and phosphorylated to 1,3-bisphosphoglycerate (BPGA). During this step, $NAD^+$ is reduced to $NADH + H^+$.
  6. ATP Generation (Substrate-level): BPGA is converted to 3-phosphoglyceric acid (PGA), yielding 1 ATP.
  7. Mutase and Dehydration: PGA is converted to 2-phosphoglycerate, and then to Phosphoenolpyruvate (PEP) with the elimination of water.
  8. Pyruvate Formation: PEP is converted to Pyruvic acid, yielding another ATP.

Net Yield of Glycolysis: For one molecule of glucose, 4 ATP are produced and 2 ATP are consumed, resulting in a net gain of 2 ATP and 2 $NADH + H^+$.

12.3 Fermentation

Fermentation is the incomplete oxidation of glucose under anaerobic conditions. It occurs in many prokaryotes, unicellular eukaryotes (like yeast), and animal muscle cells under oxygen-depleted conditions.

Types of Fermentation

  • Alcoholic Fermentation: Pyruvic acid is converted to $CO_2$ and ethanol. Catalysed by pyruvic acid decarboxylase and alcohol dehydrogenase. Occurs in yeast.
  • Lactic Acid Fermentation: Pyruvic acid is reduced to lactic acid by lactate dehydrogenase. Occurs in some bacteria and in animal muscle cells during vigorous exercise when oxygen is inadequate.

In both processes, the reducing agent is $NADH + H^+$, which is reoxidised to $NAD^+$. No extra ATP is produced beyond what was generated during glycolysis.

Characteristics and Hazards

  • Less than 7% of the energy in glucose is released, and not all of it is trapped as ATP.
  • The products (acid or alcohol) are hazardous. Yeasts poison themselves to death when the concentration of alcohol reaches about 13%.
12.4 Aerobic Respiration

Aerobic respiration is the process that leads to the complete oxidation of organic substances in the presence of oxygen, releasing $CO_2$, water, and a large amount of energy. It occurs within the mitochondria of eukaryotes.

Crucial Events in Aerobic Respiration

  1. Complete oxidation of pyruvate by the stepwise removal of all hydrogen atoms, leaving three molecules of $CO_2$ (takes place in the mitochondrial matrix).
  2. Passing on of electrons removed as part of hydrogen atoms to molecular $O_2$ with simultaneous synthesis of ATP (located on the inner mitochondrial membrane).

Oxidative Decarboxylation (Link Reaction)

Pyruvate enters the mitochondrial matrix and undergoes oxidative decarboxylation catalysed by pyruvate dehydrogenase, requiring several coenzymes including $NAD^+$ and Coenzyme A (CoA):

$$\text{Pyruvic acid} + \text{CoA} + \text{NAD}^+ \xrightarrow[\text{Pyruvate dehydrogenase}]{\text{Mg}^{2+}} \text{Acetyl CoA} + \text{CO}_2 + \text{NADH} + \text{H}^+$$

12.4.1 Tricarboxylic Acid Cycle (Krebs' Cycle)

The Tricarboxylic Acid (TCA) Cycle, or Krebs' Cycle, begins with the condensation of acetyl CoA with oxaloacetic acid (OAA) and water to yield citric acid, catalysed by citrate synthase.

  • Citrate is isomerised to isocitrate.
  • Two successive decarboxylations follow, leading to the formation of $\alpha$-ketoglutaric acid and then succinyl-CoA.
  • Succinyl-CoA is converted to succinic acid, coupled with the synthesis of GTP from GDP (substrate-level phosphorylation). GTP is subsequently converted to ATP.
  • Succinic acid is oxidised to malic acid (reducing $FAD^+$ to $FADH_2$), which is then oxidised back to OAA (reducing $NAD^+$ to $NADH + H^+$).

Summary Equation for TCA Cycle (per Pyruvic Acid):

$$\text{Pyruvic acid} + 4\text{NAD}^+ + \text{FAD}^+ + 2\text{H}_2\text{O} + \text{ADP} + \text{Pi} \rightarrow 3\text{CO}_2 + 4\text{NADH} + 4\text{H}^+ + \text{FADH}_2 + \text{ATP}$$

12.4.2 Electron Transport System (ETS) and Oxidative Phosphorylation

The Electron Transport System (ETS) is a metabolic pathway present in the inner mitochondrial membrane through which electrons pass from one carrier to another to reduce oxygen to water.

  • Complex I: NADH dehydrogenase (oxidises NADH produced in the matrix).
  • Complex II: Succinate dehydrogenase (receives reducing equivalents via $FADH_2$).
  • Complex III: Cytochrome $bc_1$ complex.
  • Complex IV: Cytochrome $c$ oxidase (contains cytochromes $a$, $a_3$, and two copper centres).
  • Cytochrome c: A small, mobile protein attached to the outer surface of the inner membrane that transfers electrons between Complex III and IV.
  • Complex V: ATP synthase, which utilizes the proton gradient to synthesise ATP.

Oxidative Phosphorylation: Unlike photophosphorylation which uses light energy, respiration uses energy from oxidation-reduction reactions to establish a proton gradient.

  • Oxidation of 1 molecule of NADH yields 3 ATP.
  • Oxidation of 1 molecule of $FADH_2$ yields 2 ATP.
  • Oxygen acts as the final hydrogen acceptor, removing protons from the system to form $H_2O$.

Mechanism of ATP Synthesis: ATP synthase consists of two components:

  • $F_1$ Headpiece: A peripheral membrane protein complex containing the catalytic site for ATP synthesis from ADP and inorganic phosphate.
  • $F_0$ Part: An integral membrane protein complex forming the channel through which protons cross the inner membrane down the electrochemical gradient. For each ATP produced, $4H^+$ pass through $F_0$ from the intermembrane space to the matrix.
12.5 The Respiratory Balance Sheet

Calculations of net ATP gain are based on several theoretical assumptions:

Though these assumptions are not strictly valid in living systems (as pathways run simultaneously and intermediates are dynamically withdrawn/added), they help estimate that there is a net gain of 38 ATP molecules during the aerobic respiration of one glucose molecule.

12.6 Amphibolic Pathway

Glucose is the favoured substrate for respiration, but fats and proteins can also be broken down:

Because the respiratory pathway is involved in both breakdown (catabolism) and synthesis (anabolism) of these substrates (e.g., withdrawing acetyl CoA to synthesise fatty acids), it is more accurately described as an amphibolic pathway rather than a purely catabolic one.

12.7 Respiratory Quotient (RQ)

The Respiratory Quotient (RQ) or respiratory ratio is the ratio of the volume of $CO_2$ evolved to the volume of $O_2$ consumed during respiration:

$$\text{RQ} = \frac{\text{Volume of } CO_2 \text{ evolved}}{\text{Volume of } O_2 \text{ consumed}}$$

The RQ value depends on the type of respiratory substrate used:

In living organisms, respiratory substrates are rarely pure; mixtures of proteins, fats, and carbohydrates are typically used.