CHM219-L7_74db0d70f1c8ea5d52a41de6c43b60e9

Page 1: Introduction

  • Course Title: Biochemistry I (CHM219)

  • Instructor: Assist. Prof. Dr. Esra Aydemir

Page 2: Topic Introduction

  • Focus on Enzymes II: The Biological Catalysts

Page 3: Outline of Key Topics

  • Kinetics of Enzymatic Catalysis

  • Enzyme Inhibition

  • Cofactors, Vitamins, and Essential Metals

  • Diversity of Enzymatic Function

  • Nonprotein Biocatalysts: Catalytic Nucleic Acids

  • Regulation of Enzyme Activity: Allosteric Enzymes

  • Covalent Modifications: Used to Regulate Enzyme Activity

Page 4: Conformational Selection vs. Induced Fit

  • Induced Fit Model: Enzyme is homogenous when unbound; changes upon substrate binding.

  • Conformational Selection Model: Enzyme exists in multiple conformations; substrate binds only to the specific conformation.

  • Le Chatelier’s Principle: Binding of substrate alters the equilibrium towards the ES complex.

Page 5: Kinetics of Enzymatic Catalysis

  • Expression for a reaction with a single substrate and product.

  • Assumption: k1, k-1, k3 >> k2 simplifies rate equations.

  • kcat: Apparent rate constant for substrate to product conversion.

Page 6: Reaction Rates and Steady-State

  • Initial conditions assumed to neglect reverse reaction between E and P.

  • Rate: First-order reaction dependent on [ES] and kcat.

  • Velocity expressed as: v = kcat[ES]

  • Difficulty in measuring [ES] during experiments.

Page 7: Steady-State Assumption

  • Desirable to express rate (v) in terms of [S] and total enzyme concentration.

  • Assumes equilibrium between E, S, and ES.

  • Steady-state conditions persist until substrate is almost consumed.

Page 8: Michaelis–Menten Equation

  • Rate equation relates initial reaction rate (v0), max rate (Vmax), and substrate concentration [S].

  • Michaelis Constant (KM): Measure of substrate-binding affinity.

Page 9: Reaction Velocity vs. Substrate Concentration

  • Graph of reaction velocity and [S] according to Michaelis–Menten model.

  • At [S] = KM, reaction rate is half of Vmax.

  • Vmax approached asymptotically.

Page 10: Steady-State Implications

  • Steady-state implies constant ES concentration.

  • KM indicates substrate concentration for half Vmax.

  • Turnover Number (kcat): Measures catalytic rate.

Page 11: Michaelis-Menten Parameters Table

Enzyme

Reaction Catalyzed

m (mol/L)

kcat (s-1)

Efficiency (kcat/KM)

Chymotrypsin

Ac-Phe-Ala to Ac-Phe

1.5 x 10-2

0.14

9.3

Pepsin

Phe-Gly to Phe

3 x 10-4

0.5

1.7 x 10³

Tyrosyl-tRNA synthetase

Tyrosine + tRNA

9 x 10-4

7.6

8.4 x 10³

Ribonuclease

Cytidine 2', 3' to

7.9 x 10-3

7.9 x 10²

1.0 x 10⁵

Carbonic anhydrase

Water

2.6 x 10-2

1.5 x 10⁷

Page 12: Preferences of Chymotrypsin

Amino Acid

Side Chain

kcat/km

Glycine

-H

1.3 x 10-1

Norvaline

-CH2CH2CH3

3.6 x 10²

Norleucine

-CH2CH2CH2CH3

3.0 x 10³

Phenylalanine

CH2

1.0 x 10⁵

Page 13: Analysis of Initial Rates

(a) Initial rates determined from varying substrate concentrations. (b) Plotted data shows adherence to Michaelis–Menten kinetics.

Page 14: Lineweaver–Burk Plot

  • Double reciprocal plot of 1/v vs. 1/[S] for determining Vmax and KM.

  • Prone to errors due to the nature of reciprocal measurements.

Page 15: Eadie–Hofstee Plot

  • Graph of v vs. V/[S], yielding Vmax at (V/[S]) = 0 and KM from slope.

  • Offers a graphical method for kinetic analysis.

Page 16: Kinetics Analysis Using Plots

  • Lineweaver–Burk and Eadie–Hofstee plots help determine KM and kcat.

  • Mutation effects analyzed through changes in KM and kcat values.

Page 17: Multisubstrate Reactions

  • Classes based on substrate binding order:

    • Random

    • Ordered

    • Ping-pong

Page 18: Random Substrate Binding

  • Either substrate can bind first, promoting the binding of the other substrate.

Page 19: Ordered Substrate Binding

  • One substrate must bind before the second can bind significantly.

Page 20: The Ping-Pong Mechanism

  • Cycle where one substrate binds, one product is released; then a second substrate and product.

Page 21: Rate Constants by Chymotrypsin

  • Table of rate constants for hydrolysis of N-acyl amino acid esters by chymotrypsin.

Page 22: Enzyme Inhibition Overview

  • Inhibition can be reversible or irreversible.

  • Competitive Inhibition: Inhibitor competes with substrate for the active site; increases apparent KM.

Page 23: Equilibrium and Substrate Interaction

  • The equilibrium state involves bound and free enzyme interactions with substrate and inhibitor.

Page 24: Competitive Inhibition Effects

  • Analysis of how competitive inhibitors affect reaction velocity at varying substrate concentrations.

Page 25: Lineweaver–Burk Plots of Inhibition

  • Competitive inhibitor lines intersect at 1/v demonstrating Vmax remains constant.

Page 26: KM and KI Determination

  • KI can be determined by varying concentrations of inhibitor, yielding KM when [I] = 0.

Page 27: Substrate and Inhibitor Examples

  • Example of UpA substrate competing with UpcA inhibitor for ribonuclease.

Page 28: Uncompetitive Inhibition Characteristics

  • Inhibitor does not compete for the active site, reducing Vmax and KM.

Page 29: Uncompetitive Inhibition Kinetics

  • Model displaying the interaction of E, S, and uncompetitive inhibition effects.

Page 30: Uncompetitive Inhibition Effects

(a) Effect of uncompetitive inhibitor on reaction velocity. (b) Parallel lines in Lineweaver–Burk plots indicate uncompetitive inhibition.

Page 31: Mixed Inhibition Model

  • Inhibitor binds to both free enzyme and ES complex, affecting substrate binding and catalytic events.

Page 32: Mixed Inhibition Mechanism

  • Kinetic model depicting the interaction of the enzyme with substrate and inhibitor.

Page 33: Lineweaver–Burk Plot for Mixed Inhibition

  • Vmax decreased; KM increased indicating mixed inhibition characteristics.

Page 34: Irreversible Inhibitors Overview

  • Irreversible inhibitors often bind covalently to enzymes; example: DFP.

Page 35: Irreversible Enzyme Inhibitors Table

Name

Formula

Source

Mode of Action

Cyanide

CN-

Bitter almonds

Reacts with enzyme metal ions

Diisopropyl F

Fluorophosphate

Synthetic

Inhibits serine-containing enzymes

Sarin

F

Synthetic

Similar to DFP; a nerve gas

Physostigmine

Calabar beans

Similar to DFP

Parathion

Synthetic

Inhibitor of acetylcholinesterase

Penicillin

...

Penicillium fungus

Inhibitor of bacterial cell wall synthesis

Page 36: Cofactors, Vitamins, and Essential Metals

  • Table of important enzyme cofactors and their associated vitamins.

Page 37: Nicotinamide Nucleotides

  • NAD+ as a key cofactor derived from niacin, crucial in redox reactions.

Page 38: NAD+ as Reducing Agent

  • NAD+ acts as an oxidizing agent in biological reactions, reversible electron transfer.

Page 39: Typical Reactions with NAD+

  • Role of NAD+ in converting alcohols to aldehydes or ketones via dehydrogenase.

Page 40: Trace Elements as Cofactors

  • Importance of trace metals like Fe, Cu, Zn in enzymatic functions.

Page 41: Diversity of Enzymatic Function

  • Some enzymatic reactions depend on metal ions for catalytic activity.

Page 42: Enzyme Classification by IUBMB

  • Six major enzyme classes defined by reaction types:

    1. Oxidoreductases

    2. Transferases

    3. Hydrolases

    4. Lyases

    5. Isomerases

    6. Ligases

Page 43: Examples of Major Enzyme Classes

Class

Example

Reaction Catalyzed

Oxidoreductases

Alcohol dehydrogenase

CH3CH2OH to Acetaldehyde

Transferases

Hexokinase

D-Glucose to D-Glucose-6-P

Hydrolases

Carboxypeptidase A

Peptide bond hydrolysis

Ligases

Pyruvate carboxylase

Carboxylation reaction

Page 44: Nonprotein Biocatalysts

  • Ribozymes: RNA molecules capable of catalyzing specific reactions.

Page 45: DNAzymes and Rate Enhancements

  • Examples of reactions catalyzed by DNAzymes, with associated rate enhancements.

Page 46: Allosteric Enzymes

  • Regulation of enzyme activity crucial for metabolic pathways; feedback control.

Page 47: Cooperative Substrate Binding

  • Comparison of kinetic curves for noncooperative vs. allosteric enzymes.

Page 48: Extreme Homoallostery

  • Enzyme activity sharply increases at specific substrate concentration levels.

Page 49: Characteristics of Allosteric Enzymes

  • Display sigmoidal v vs. [S] curves; can be affected by various inhibitors and activators.

Page 50: Control Points in Pyrimidine Synthesis

  • Essential regulation points in the synthetic pathway, involving aspartate carbamoyltransferase.

Page 51: Regulation by ATP and CTP

  • ATP as an activator and CTP as an inhibitor of ATCase activity.

Page 52: Quaternary Structure of ATCase

  • Structure details and transitions involved between T and R states.

Page 53: Covalent Modifications Overview

  • Four types of covalent modifications that influence enzyme activities.

Page 54: Kinases and Phosphatases

  • Mechanisms by which kinases and phosphatases regulate enzyme activity via reversible modification.

Page 55: Proteolytic Cleavage for Activation

  • Certain enzymes activated irreversibly through cleaving zymogens.

Page 56: Activation of Chymotrypsinogen

  • Cleavage process yielding active chymotrypsin from its zymogen form.

Page 57: Blood Clotting Process

  • Cascade of proteolytic activations leading to fibrin formation.

Page 58: Formation of a Blood Clot

  • Detailed explanation of blood clot formation with fibrin monomer interaction.