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Chapter 13: How Cells Obtain Energy

Energy Sources

  • Energy is primarily derived from:
    • Carbohydrates
    • Fats
    • Proteins

Molecules that Can Store Chemical Energy

  • Common energy storage molecules include:
    • ATP (Adenosine Triphosphate)
    • NADH (Nicotinamide Adenine Dinucleotide)
    • NADPH (Nicotinamide Adenine Dinucleotide Phosphate)
    • FADH₂ (Flavin Adenine Dinucleotide)

Energy Currency of Cells

ATP Overview
  • ATP is known as the energy “currency” of cells.
    • Discovered by Karl Lohmann in 1929.
    • Identified as the main energy molecule by Fritz Albert Lipmann in 1941.
  • Structure of ATP:
    • Ribose (a 5-carbon sugar)
    • Adenine
    • Three phosphate groups
Energy Storage in ATP
  • ATP stores energy in the bonds between phosphates:
    • Reason for high energy: Phosphates are negatively charged, which leads to:
    • Repulsion between phosphates.
    • Increased energy required to keep the phosphates bound to each other.
    • Increased energy is released when the bond between two phosphates is broken.

ATP Hydrolysis

  • Reaction:
    • When the bond between phosphates is broken:
    • ATP + H2O ightarrow ADP + Pi + ext{energy released}
    • This process is called hydrolysis.
    • ADP (Adenosine Diphosphate) and P_i (inorganic phosphate) are the products of ATP hydrolysis, which is a reversible reaction.
Gibbs Free Energy Change
  • Gibbs Free Energy Change for hydrolysis of ATP:
    • riangleG=−30.5extkJ/molriangle G = -30.5 ext{ kJ/mol} (-7.3 kcal/mol)
    • When coupled to a reaction, as in:
    • A + ATP + H2O ightleftharpoons B + ADP + Pi + H^+
    • This yields riangleG=−13.8extkJ/molriangle G = -13.8 ext{ kJ/mol}.
    • ATP acts to make reactions progress forward rather than revert, leading cells to maintain high levels of ATP.

ATP Structure

  • ATP contains two energy-rich phosphoanhydride bonds:
    • Structure depicted shows phosphate groups and their arrangement with ribose and adenine.

ATP Synthesis and Hydrolysis

  • Catabolic Reactions:
    • Energy from the oxidation of nutrients is stored in high-energy bonds, contributing to ATP synthesis.
  • Anabolic Reactions:
    • High-energy bonds are broken, releasing energy.

High Energy Electron Carriers

  • Some reactions utilize high-energy electron carriers instead of ATP:
    • NAD+: Nicotinamide adenine dinucleotide
    • NADP+: Nicotinamide adenine dinucleotide phosphate
    • FAD: Flavin adenine dinucleotide
  • These carriers are involved in oxidation-reduction reactions and transfer 2 electrons.

Oxidation-Reduction Reactions

Definitions
  • Oxidation: Loss of electrons; does not always include the addition of oxygen.
  • Reduction: Gain of electrons; often involves the addition of hydrogen.
  • Electrons in organic redox reactions are typically transferred in the form of a hydride ion (a proton and two electrons), linking to hydrogenation and dehydrogenation reactions.
Reaction Dynamics
  • Oxidation and reduction occur coupled, as illustrated in the transitions of ADP and similar molecules during these reactions.

Respiation

Aerobic Respiration
  • Overall reaction for glucose oxidation:
    • C6H{12}O6 + 6O2
      ightarrow 6CO2 + 6H2O + E
    • The change in free energy: ∆Gext −686extkcal/mol∆G ext{ ~} -686 ext{kcal/mol} for glucose, with energy released needing to be produced in small steps.

Glycolysis

Overview
  • Glycolysis is the process that converts glucose (C₆) into two pyruvate (C₃):
    • A 10-step biochemical pathway.
    • Takes place in the cytoplasm of both prokaryotic and eukaryotic cells.
  • Results in:
    • Formation of 2 pyruvate
    • Net production of 2 ATP molecules by substrate-level phosphorylation
    • Production of 2 NADH molecules through reduction of NAD+
  • Net Reaction:
    • ext{Glucose} + 2NAD^+ + 2Pi + 2ADP ightarrow 2 ext{pyruvate} + 2ATP + 2NADH + 2H2O
Importance of Glucose
  • Glucose is a fundamental substrate in energy metabolism:
    • Primary energy source for most cells.
    • Acquired from dietary carbohydrates (e.g., starch, sucrose, lactose) and stored glucose (glycogen in animals and starch in plants).
Overview of Glycolysis Steps
  • The glycolytic process can be divided into two phases:
    1. Energy Investment Phase:
    • Consumes 2 ATP.
    1. Energy Generation Phase:
    • Produces 4 ATP, resulting in a net gain of 2 ATP.
    • Involves production of 2 NADH from glyceraldehyde 3-phosphate.
  • Overall pathway from Glucose (C₆) to Pyruvate (C₃) illustrated through multiple steps:
    • Glucose → Glucose 6-phosphate → Fructose 6-phosphate → Fructose 1,6-bisphosphate (and so on through the breakdown to pyruvate).
Energy Expenditure and Gain in Glycolysis
  • Summary of ATP and NADH synthesis in glycolysis and overall energy gain outlined in multiple steps within the pathway leading to pyruvate production.

Fate of Pyruvate

Conditions Affecting Fate
  • The fate of pyruvate depends on the availability of oxygen:
    • Aerobic conditions:
    • Pyruvate is completely oxidized to CO2 and H2O.
    • Anaerobic conditions:
    • Fermentation occurs, including lactic acid and alcohol fermentation options.

Pyruvate Oxidation

Overview
  • In the presence of oxygen, pyruvate undergoes oxidation:
    • Occurs in the mitochondrial matrix of eukaryotes while happening in the cytosol for prokaryotes.
  • Process is mediated by pyruvate dehydrogenase, which is a multienzyme complex consisting of over 60 different polypeptide chains, yielding products such as carbon dioxide and acetyl-CoA (2 carbons attached to coenzyme A).
Reaction Catalysis and Transport
  • The transport of pyruvate into mitochondria is facilitated via active transport due to its charge, while the products resulting from pyruvate oxidation include 1 CO2, 1 NADH, and 1 acetyl-CoA.

Krebs Cycle / TCA Cycle

Overview
  • The Krebs Cycle is responsible for the complete oxidation of acetyl-CoA:
    • Takes place in the mitochondrial matrix; consists of 8 total steps.
    • Overall reaction per turn:
    • Acetyl-CoA + 3NAD^+ + FAD + GDP + Pi ightarrow 2CO2 + 3NADH + FADH_2 + GTP + CoA
Reaction Summary
  • Each turn of the Krebs cycle reduces:
    • 3 NAD+ to NADH,
    • 1 FAD to FADH2,
    • Produces 1 ATP/GTP,
    • Regenerates oxaloacetate and releases 2 CO2.
  • Upon reviewing what has been produced from glycolysis and pyruvate oxidation:
    • 2 CO2, 4 NADH, 2 ATP (produced via substrate-level phosphorylation).
Free-Energy Changes vs. Glucose
  • Changes relative to glucose quantified and diagrammatic representation along with total energy accounting depicted, summarizing yields (2 NADH, 2 ATP, etc.) from glycolysis to Krebs cycle.

Important Steps of Glycolysis

  • Reactions Consuming ATP:
    • Hexokinase, Phosphofructokinase
  • Reactions Synthesizing ATP:
    • Phosphoglycerate Kinase, Pyruvate Kinase
  • Reactions Synthesizing NADH:
    • Glyceraldehyde 3-Phosphate Dehydrogenase
  • Final Products of Glycolysis:
    • 2 ATP, 2 NADH, and 2 Pyruvate

Important Steps of the Krebs Cycle

  • Reactions Synthesizing NADH:
    • Isocitrate Dehydrogenase, α-Ketoglutarate Dehydrogenase, Malate Dehydrogenase
  • Reactions Synthesizing FADH2:
    • Succinate Dehydrogenase
  • Reactions Synthesizing GTP:
    • Succinyl CoA Synthetase
  • Reactions Releasing CO2:
    • Isocitrate Dehydrogenase, α-Ketoglutarate Dehydrogenase

Overall Yield from Complete Glucose Oxidation

  • Pathway Contributions to energy yield are finite as:
    • Glycolysis: 2 ATP, 2 NADH, 0 FADH2, 0 CO2
    • Pyruvate Dehydrogenation: 0 ATP, 2 NADH, 0 FADH2, 2 CO2
    • Krebs Cycle: 2 ATP, 6 NADH, 2 FADH2, 4 CO2
  • Total Yield from glycolysis to Krebs cycle:
    • ATP: 4
    • NADH: 10
    • FADH2: 2
    • CO2: 6