Professor Paul Engel is professor of biochemistry at University College Dublin. He has had extensive experience teaching biochemistry to undergraduate and postgraduate biochemistry, biology, medicine, engineering and nursing students, in his current post at UCD and at the universities of Sheffield, Oxford and Hong Kong. He is on the Editorial Board of Biochemical Journal, and Essays in Biochemistry, and is the author of the highly successful textbook: Enzyme Kinetics (Outline Series) published by Chapman and Hall.

"For students embarking on careers in the health sciences, it is an essential academic companion and, retailing at approximately £28, it is excellent value for money". (The Biochemist, 1 June 2011) "It's not every day that one picks up a textbook that can claim to occupy a unique niche, given the multitude of scientific textbooks that are vying for a medical readership. However, with the recent publication of Pain-Free Biochemistry: An Essential Guide for the Health Sciences, which is specifically aimed at students of medicine and nursing, one could be left wondering just why nobody thought of this sooner." (Irish Medical News, September 08, 2010)

• Preface xSection 1 Foundations 1Topic 1 Why biochemistry? 3Topic 2 Remarkableness of life 9Chemistry I The basic structure of substances: atoms, molecules, elements and compounds 13Chemistry II Atomic structure, valency and bonding 16Chemistry III Protons, acids, bases, concentration and the pH scale 25Topic 3 Shape, molecular recognition and proteins: an example 32Topic 4 Proteins: molecular necklaces 38Topic 5 Chemical transformations in the living organism: metabolism 43Topic 6 Reactions, catalysts and enzymes 46Topic 7 Specificity, saturation and active sites 49Topic 8 Structure of metabolism: anabolism and catabolism 54Chemistry IV Equilibrium 56Topic 9 Catabolism: degradation vs energy metabolism 60Chemistry V Oxidation and reduction 63Topic 10 Oxidation and reduction in metabolism 67Section 2 Catabolism 71Chemistry VI Aldehydes, ketones and sugars 73Topic 11 Carbohydrates: sugars and polysaccharides in metabolism 81Topic 12 Glucose inside the body 85Topic 13 Breakdown of sugar: glycolysis 88Topic 14 Aerobic oxidation of pyruvate: Krebs cycle 94Topic 15 Respiratory chain, oxidative phosphorylation and overall ATP yields 98Topic 16 Mobilising the carbohydrate store: glycogenolysis 103Chemistry VII Alcohols, esters, glycerol, fatty acids and triglycerides 106Chemistry VIII Hydrophobic, hydrophilic and amphiphilic 109Topic 17 Phospholipids and membranes 111Chemistry IX Saturated and unsaturated 114Topic 18 Fats as an energy source 119Topic 19 Fats: digestion, transport, storage and mobilisation 122Topic 20 Fats: oxidation of fatty acids 126Topic 21 Ketone bodies in health and disease 133Topic 22 Dietary fat: essential fatty acids 137Topic 23 Protein and amino acid breakdown 140Topic 24 Shedding excess amino groups: urea cycle 147Section 3 Anabolism and Control 151Topic 25 Is anabolism just catabolism backwards? 153Topic 26 Making new glucose: gluconeogenesis 156Topic 27 Fatty acid biosynthesis 161Topic 28 Providing reducing power: NADPH and the pentose phosphate pathway 164Chemistry X Isotopes 168Topic 29 Red cells and white cells: defence against reactive oxygen and reactive oxygen as defence! 172Topic 30 The need for metabolic control 176Topic 31 Relationship of fats and carbohydrates: use by different tissues 179Section 4 Genes and Protein Synthesis 181Topic 32 The idea of genes 183Topic 33 The chemistry of genes: DNA and the double helix 186Topic 34 The genetic code and mRNA 190Topic 35 Protein synthesis, ribosomes and tRNA 193Topic 36 Genetic differences and disease 197Topic 37 Genetic variability: drug metabolism and disease susceptibility 203Topic 38 Mutation, radiation and ageing 205Topic 39 Switching genes on and off: development, tissue specificity, adaptation and tolerance 207Topic 40 DNA and protein synthesis as targets: chemotherapy, antibiotics, etc. 210Section 5 Physiological Systems and Clinical Issues 217Topic 41 Hormones and second messengers 219Topic 42 Switching enzymes on and off: coarse and fine control 222Topic 43 Insulin, glucagon and adrenaline 224Topic 44 Diabetes 229Topic 45 Steroid hormones and receptors: fertility control, pregnancy testing, etc. 233Topic 46 Pituitary hormones and feedback loops 238Topic 47 Thyroid hormones 240Topic 48 Adrenal cortex 244Topic 49 Prostaglandins and inflammation: aspirin 246Topic 50 Membrane transport 250Topic 51 Nerve and muscle 255Topic 52 pH homeostasis 258Topic 53 Diagnostic markers: biochemical tests 261Topic 54 Blood, bleeding and clotting 265Section 6 Appendices 271Appendix 1 pH and neutrality 273Appendix 2 Crystallography 274Appendix 3 Protein forces, secondary structure and folding 277Appendix 4 Equilibrium constant 281Appendix 5 Phosphorus, phosphoric acid and phosphate esters 283Appendix 6 Coenzymes, cofactors and prosthetic groups 285Appendix 7 Coenzyme A 287Appendix 8 Krebs cycle and evidence for a catalytic reaction sequence 289Appendix 9 Knoop’s experiment pointing to β-oxidation of fatty acids 291Appendix 10 Isoenzymes 293Appendix 11 Genetic code 295Appendix 12 Different kinds of mutation 297Appendix 13 Restriction enzymes 299Appendix 14 Enzyme inhibition 301Appendix 15 Electrophoresis to separate proteins 304Appendix 16 Chromatography and mass spectrometry toseparate and identify metabolites 308Glossary 313MCQ answers 322Index 323

Professor Paul Engel is professor of biochemistry at University College Dublin. He has had extensive experience teaching biochemistry to undergraduate and postgraduate biochemistry, biology, medicine, engineering and nursing students, in his current post at UCD and at the universities of Sheffield, Oxford and Hong Kong. He is on the Editorial Board of Biochemical Journal, and Essays in Biochemistry, and is the author of the highly successful textbook: Enzyme Kinetics (Outline Series) published by Chapman and Hall.

"For students embarking on careers in the health sciences, it is an essential academic companion and, retailing at approximately £28, it is excellent value for money". (The Biochemist, 1 June 2011) "It's not every day that one picks up a textbook that can claim to occupy a unique niche, given the multitude of scientific textbooks that are vying for a medical readership. However, with the recent publication of Pain-Free Biochemistry: An Essential Guide for the Health Sciences, which is specifically aimed at students of medicine and nursing, one could be left wondering just why nobody thought of this sooner." (Irish Medical News, September 08, 2010)

• Preface xSection 1 Foundations 1Topic 1 Why biochemistry? 3Topic 2 Remarkableness of life 9Chemistry I The basic structure of substances: atoms, molecules, elements and compounds 13Chemistry II Atomic structure, valency and bonding 16Chemistry III Protons, acids, bases, concentration and the pH scale 25Topic 3 Shape, molecular recognition and proteins: an example 32Topic 4 Proteins: molecular necklaces 38Topic 5 Chemical transformations in the living organism: metabolism 43Topic 6 Reactions, catalysts and enzymes 46Topic 7 Specificity, saturation and active sites 49Topic 8 Structure of metabolism: anabolism and catabolism 54Chemistry IV Equilibrium 56Topic 9 Catabolism: degradation vs energy metabolism 60Chemistry V Oxidation and reduction 63Topic 10 Oxidation and reduction in metabolism 67Section 2 Catabolism 71Chemistry VI Aldehydes, ketones and sugars 73Topic 11 Carbohydrates: sugars and polysaccharides in metabolism 81Topic 12 Glucose inside the body 85Topic 13 Breakdown of sugar: glycolysis 88Topic 14 Aerobic oxidation of pyruvate: Krebs cycle 94Topic 15 Respiratory chain, oxidative phosphorylation and overall ATP yields 98Topic 16 Mobilising the carbohydrate store: glycogenolysis 103Chemistry VII Alcohols, esters, glycerol, fatty acids and triglycerides 106Chemistry VIII Hydrophobic, hydrophilic and amphiphilic 109Topic 17 Phospholipids and membranes 111Chemistry IX Saturated and unsaturated 114Topic 18 Fats as an energy source 119Topic 19 Fats: digestion, transport, storage and mobilisation 122Topic 20 Fats: oxidation of fatty acids 126Topic 21 Ketone bodies in health and disease 133Topic 22 Dietary fat: essential fatty acids 137Topic 23 Protein and amino acid breakdown 140Topic 24 Shedding excess amino groups: urea cycle 147Section 3 Anabolism and Control 151Topic 25 Is anabolism just catabolism backwards? 153Topic 26 Making new glucose: gluconeogenesis 156Topic 27 Fatty acid biosynthesis 161Topic 28 Providing reducing power: NADPH and the pentose phosphate pathway 164Chemistry X Isotopes 168Topic 29 Red cells and white cells: defence against reactive oxygen and reactive oxygen as defence! 172Topic 30 The need for metabolic control 176Topic 31 Relationship of fats and carbohydrates: use by different tissues 179Section 4 Genes and Protein Synthesis 181Topic 32 The idea of genes 183Topic 33 The chemistry of genes: DNA and the double helix 186Topic 34 The genetic code and mRNA 190Topic 35 Protein synthesis, ribosomes and tRNA 193Topic 36 Genetic differences and disease 197Topic 37 Genetic variability: drug metabolism and disease susceptibility 203Topic 38 Mutation, radiation and ageing 205Topic 39 Switching genes on and off: development, tissue specificity, adaptation and tolerance 207Topic 40 DNA and protein synthesis as targets: chemotherapy, antibiotics, etc. 210Section 5 Physiological Systems and Clinical Issues 217Topic 41 Hormones and second messengers 219Topic 42 Switching enzymes on and off: coarse and fine control 222Topic 43 Insulin, glucagon and adrenaline 224Topic 44 Diabetes 229Topic 45 Steroid hormones and receptors: fertility control, pregnancy testing, etc. 233Topic 46 Pituitary hormones and feedback loops 238Topic 47 Thyroid hormones 240Topic 48 Adrenal cortex 244Topic 49 Prostaglandins and inflammation: aspirin 246Topic 50 Membrane transport 250Topic 51 Nerve and muscle 255Topic 52 pH homeostasis 258Topic 53 Diagnostic markers: biochemical tests 261Topic 54 Blood, bleeding and clotting 265Section 6 Appendices 271Appendix 1 pH and neutrality 273Appendix 2 Crystallography 274Appendix 3 Protein forces, secondary structure and folding 277Appendix 4 Equilibrium constant 281Appendix 5 Phosphorus, phosphoric acid and phosphate esters 283Appendix 6 Coenzymes, cofactors and prosthetic groups 285Appendix 7 Coenzyme A 287Appendix 8 Krebs cycle and evidence for a catalytic reaction sequence 289Appendix 9 Knoop’s experiment pointing to β-oxidation of fatty acids 291Appendix 10 Isoenzymes 293Appendix 11 Genetic code 295Appendix 12 Different kinds of mutation 297Appendix 13 Restriction enzymes 299Appendix 14 Enzyme inhibition 301Appendix 15 Electrophoresis to separate proteins 304Appendix 16 Chromatography and mass spectrometry toseparate and identify metabolites 308Glossary 313MCQ answers 322Index 323

Professor Paul Engel is professor of biochemistry at University College Dublin. He has had extensive experience teaching biochemistry to undergraduate and postgraduate biochemistry, biology, medicine, engineering and nursing students, in his current post at UCD and at the universities of Sheffield, Oxford and Hong Kong. He is on the Editorial Board of Biochemical Journal, and Essays in Biochemistry, and is the author of the highly successful textbook: Enzyme Kinetics (Outline Series) published by Chapman and Hall.

"For students embarking on careers in the health sciences, it is an essential academic companion and, retailing at approximately £28, it is excellent value for money". (The Biochemist, 1 June 2011) "It's not every day that one picks up a textbook that can claim to occupy a unique niche, given the multitude of scientific textbooks that are vying for a medical readership. However, with the recent publication of Pain-Free Biochemistry: An Essential Guide for the Health Sciences, which is specifically aimed at students of medicine and nursing, one could be left wondering just why nobody thought of this sooner." (Irish Medical News, September 08, 2010)

• Preface xSection 1 Foundations 1Topic 1 Why biochemistry? 3Topic 2 Remarkableness of life 9Chemistry I The basic structure of substances: atoms, molecules, elements and compounds 13Chemistry II Atomic structure, valency and bonding 16Chemistry III Protons, acids, bases, concentration and the pH scale 25Topic 3 Shape, molecular recognition and proteins: an example 32Topic 4 Proteins: molecular necklaces 38Topic 5 Chemical transformations in the living organism: metabolism 43Topic 6 Reactions, catalysts and enzymes 46Topic 7 Specificity, saturation and active sites 49Topic 8 Structure of metabolism: anabolism and catabolism 54Chemistry IV Equilibrium 56Topic 9 Catabolism: degradation vs energy metabolism 60Chemistry V Oxidation and reduction 63Topic 10 Oxidation and reduction in metabolism 67Section 2 Catabolism 71Chemistry VI Aldehydes, ketones and sugars 73Topic 11 Carbohydrates: sugars and polysaccharides in metabolism 81Topic 12 Glucose inside the body 85Topic 13 Breakdown of sugar: glycolysis 88Topic 14 Aerobic oxidation of pyruvate: Krebs cycle 94Topic 15 Respiratory chain, oxidative phosphorylation and overall ATP yields 98Topic 16 Mobilising the carbohydrate store: glycogenolysis 103Chemistry VII Alcohols, esters, glycerol, fatty acids and triglycerides 106Chemistry VIII Hydrophobic, hydrophilic and amphiphilic 109Topic 17 Phospholipids and membranes 111Chemistry IX Saturated and unsaturated 114Topic 18 Fats as an energy source 119Topic 19 Fats: digestion, transport, storage and mobilisation 122Topic 20 Fats: oxidation of fatty acids 126Topic 21 Ketone bodies in health and disease 133Topic 22 Dietary fat: essential fatty acids 137Topic 23 Protein and amino acid breakdown 140Topic 24 Shedding excess amino groups: urea cycle 147Section 3 Anabolism and Control 151Topic 25 Is anabolism just catabolism backwards? 153Topic 26 Making new glucose: gluconeogenesis 156Topic 27 Fatty acid biosynthesis 161Topic 28 Providing reducing power: NADPH and the pentose phosphate pathway 164Chemistry X Isotopes 168Topic 29 Red cells and white cells: defence against reactive oxygen and reactive oxygen as defence! 172Topic 30 The need for metabolic control 176Topic 31 Relationship of fats and carbohydrates: use by different tissues 179Section 4 Genes and Protein Synthesis 181Topic 32 The idea of genes 183Topic 33 The chemistry of genes: DNA and the double helix 186Topic 34 The genetic code and mRNA 190Topic 35 Protein synthesis, ribosomes and tRNA 193Topic 36 Genetic differences and disease 197Topic 37 Genetic variability: drug metabolism and disease susceptibility 203Topic 38 Mutation, radiation and ageing 205Topic 39 Switching genes on and off: development, tissue specificity, adaptation and tolerance 207Topic 40 DNA and protein synthesis as targets: chemotherapy, antibiotics, etc. 210Section 5 Physiological Systems and Clinical Issues 217Topic 41 Hormones and second messengers 219Topic 42 Switching enzymes on and off: coarse and fine control 222Topic 43 Insulin, glucagon and adrenaline 224Topic 44 Diabetes 229Topic 45 Steroid hormones and receptors: fertility control, pregnancy testing, etc. 233Topic 46 Pituitary hormones and feedback loops 238Topic 47 Thyroid hormones 240Topic 48 Adrenal cortex 244Topic 49 Prostaglandins and inflammation: aspirin 246Topic 50 Membrane transport 250Topic 51 Nerve and muscle 255Topic 52 pH homeostasis 258Topic 53 Diagnostic markers: biochemical tests 261Topic 54 Blood, bleeding and clotting 265Section 6 Appendices 271Appendix 1 pH and neutrality 273Appendix 2 Crystallography 274Appendix 3 Protein forces, secondary structure and folding 277Appendix 4 Equilibrium constant 281Appendix 5 Phosphorus, phosphoric acid and phosphate esters 283Appendix 6 Coenzymes, cofactors and prosthetic groups 285Appendix 7 Coenzyme A 287Appendix 8 Krebs cycle and evidence for a catalytic reaction sequence 289Appendix 9 Knoop’s experiment pointing to β-oxidation of fatty acids 291Appendix 10 Isoenzymes 293Appendix 11 Genetic code 295Appendix 12 Different kinds of mutation 297Appendix 13 Restriction enzymes 299Appendix 14 Enzyme inhibition 301Appendix 15 Electrophoresis to separate proteins 304Appendix 16 Chromatography and mass spectrometry toseparate and identify metabolites 308Glossary 313MCQ answers 322Index 323