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 ADVANCED NUTRITION
AND HUMAN METABOLISM
EIGHTH EDITION

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 ADVANCED NUTRITION
AND HUMAN METABOLISM
EIGHTH EDITION

Sareen S. Gropper
FLORIDA ATLANTIC UNIVERSITY
AUBURN UNIVERSITY (PROFESSOR EMERITUS)

Jack L. Smith
UNIVERSITY OF DELAWARE

Timothy P. Carr
UNIVERSITY OF NEBRASKA-LINCOLN

Australia • Brazil • Canada • Mexico • Singapore • United Kingdom • United States

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 Advanced Nutrition and Human Metabolism, © 2022, 2018, 2013 Cengage Learning, Inc.
Eighth Edition
Sareen S. Gropper, Jack L. Smith, and
ALL RIGHTS RESERVED. No part of this work covered by the copyright herein
Timothy P. Carr
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 To my children Michelle and Michael and their spouses, and to my husband,
Daniel, for their ongoing encouragement, support, faith, and love and to the
students who continue to impress and inspire me.
Sareen Gropper

To my wife, Carol, for her continued support, constant inspiration, and

assistance in the preparation of this book.
Jack Smith

To my wife, Marion, and my children, Erin and Rebecca, for their love,
humor, and support. And to the many students who have made my career so
worthwhile.
Tim Carr

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BRIEF CONTENTS

Preface xvii

SECTION I Cells and Their Nourishment

1 The Cell: A Microcosm of Life 1
2 The Digestive System: Mechanism for Nourishing The Body 29

SECTION II Macronutrients and Their Metabolism

3 Carbohydrates 63
4 Fiber 113
5 Lipids 131
6 Protein 187
7 Integration and Regulation of Metabolism and the
Impact of Exercise 261
8 Energy Expenditure, Body Composition, and Healthy Weight 293

SECTION III The Regulatory Nutrients

9 Water-Soluble Vitamins 321
10 Fat-Soluble Vitamins 401
11 Major Minerals 463
12 Water and Electrolytes 499
13 Essential Trace and Ultratrace Minerals 525
14 Nonessential Trace and Ultratrace Minerals 595

Glossary 609
Index 615

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CONTENTS

Preface xvii Regulatory Peptides 57

Summary 59
PERSPECTIVE The Nutritional Impact of Roux-En-Y Gastric Bypass,
SECTION I
A Surgical Approach for the Treatment of Obesity 60
CELLS AND THEIR NOURISHMENT

CHAPTER 1 The Cell: A Microcosm of Life 1 SECTION II

1.1 Components of Cells 1 MACRONUTRIENTS AND THEIR
Plasma Membrane 1 METABOLISM
Cytosol and Cytoskeleton 4
Mitochondrion 5 CHAPTER 3 Carbohydrates 63
Nucleus 6
3.1 Simple Carbohydrates 63
Endoplasmic Reticulum and Golgi Apparatus 10
Monosaccharides 63
Lysosomes and Peroxisomes 11
Disaccharides 66
1.2 Selected Cellular Proteins 11
SYRUPS – LIQUID SUGAR 67
Receptors 11
Catalytic Proteins (Enzymes) 13 3.2 Complex Carbohydrates 68
Oligosaccharides 68
1.3 Apoptosis 17
Polysaccharides 69
1.4 Biological Energy 18
3.3 Digestion 69
Energy Release and Consumption in Chemical
Digestion of Polysaccharides 70
Reactions 18
Digestion of Disaccharides 70
Units and Expressions of Energy 19
The Role of High-Energy Phosphate in Energy 3.4 Absorption and Transport 72
Storage 22 Membrane Transport 72
Coupled Reactions in the Transfer of Energy 23 Intestinal Absorption of Glucose and Galactose 75
Reduction Potentials 24 Intestinal Absorption of Fructose 75
Hepatic Metabolism of Dietary Monosaccharides 76
Summary 25
3.5 Maintenance of Blood Glucose Concentration 76
PERSPECTIVE Nutritional Genomics 26
Role of Insulin 76
Blood–Tissue Barriers 78
CHAPTER 2 The Digestive System: Mechanism for Glycemic Response to Carbohydrates 78
Nourishing The Body 29 3.6 Integrated Metabolism in Tissues 80
Glycogenesis 80
2.1 The Structures of the Digestive Tract and the Glycogenolysis 83
Digestive and Absorptive Processes 29 Glycolysis 85
The Oral Cavity 33 The Tricarboxylic Acid Cycle 88
The Esophagus 34 Formation of ATP 92
The Stomach 36 The Pentose Phosphate Pathway (Hexose
The Small Intestine 41 Monophosphate Shunt) 98
The Accessory Organs 45 UNCOUPLING ELECTRON TRANSPORT AND ATP SYNTHESIS 98
The Absorptive Process 50 Gluconeogenesis 100
The Colon (Large Intestine) 52
3.7 Regulation of Metabolism 103
2.2 Coordination and Regulation of the Allosteric Enzyme Modulation 103
Digestive Process 56 Covalent Regulation 104
Neural Regulation 56 Directional Shifts in Reversible Reactions 104

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x co n te n ts

Enzyme Translocation 104 Phospholipid Digestion 148

Genetic Regulation 105 Cholesterol Ester Digestion 148
Metabolic Control of Glycolysis and 5.4 Absorption 148
Gluconeogenesis 105 Fatty Acid, Monoacylglycerol, and Lysophospholipid
Summary 106 Absorption 148
PERSPECTIVE What Carbohydrates Do Americans Eat? 109 Cholesterol Absorption 149
Lipid Release into Circulation 150
5.5 Transport and Storage 151
CHAPTER 4 Fiber 113 Lipoprotein Structure 151
4.1 Definitions 113 Lipoprotein Metabolism 153
4.2 Fiber and Plants 114 5.6 Lipids, Lipoproteins, and Cardiovascular
4.3 Chemistry and Characteristics of Fiber 114 Disease Risk 159
Cellulose 114 The Lipid Hypothesis 160
Hemicellulose 117 Lipoprotein(a) 160
Pectins 117 Apolipoprotein E 160
Lignin 117 Dietary Cholesterol 161
Gums 117 Saturated and Unsaturated Fatty Acids 161
β-Glucans 118 COCONUT OIL: HERO OR VILLAIN? 162
Fructans 118 Trans Fatty Acids 162
Galactans 118 5.7 Integrated Metabolism in Tissues 163
Resistant Starch 118 Catabolism of Triacylglycerols and Fatty Acids 163
Mucilages (Psyllium) 119 Formation of Ketone Bodies 167
Polydextrose and Polyols 119 Synthesis of Fatty Acids 169
Chitin and Chitosan 119 Synthesis of Triacylglycerols and Phospholipids 174
4.4 Selected Properties of Fiber and Their Synthesis, Catabolism, and Whole-Body Balance
Physiological Impact 120 of Cholesterol 174
Solubility in Water 120 5.8 Regulation of Lipid Metabolism 176
Viscosity and Gel Formation 121 Fatty Acids 176
Fermentability 121 Cholesterol 176
4.5 Health Benefits of Fiber 122 5.9 Brown Fat Thermogenesis 177
Cardiovascular Disease 122 5.10 Ethyl Alcohol: Metabolism and
Diabetes Mellitus 123 Biochemical Impact 178
Appetite and/or Satiety and Weight Control 123 The Alcohol Dehydrogenase Pathway 179
Gastrointestinal Disorders 123 The Microsomal Ethanol Oxidizing System 179
4.6 Food Labels and Health Claims 124 The Catalase System 179
4.7 Recommended Fiber Intake 125 Alcoholism: Biochemical and Metabolic
Summary 126 Alterations 180
PERSPECTIVE The Flavonoids: Roles in Health and Disease
Alcohol in Moderation: The Brighter Side 181
Prevention 127 Summary 181
PERSPECTIVE The Role of Lipoproteins and Inflammation in
Atherosclerosis 184
CHAPTER 5 Lipids 131
5.1 Structure and Biological Importance 132
Fatty Acids 132 CHAPTER 6 Protein 187
Triacylglycerols (Triglycerides) 135
6.1 Amino Acid Classification 187
Phospholipids 137
Structure 188
Sphingolipids 139
Net Electrical Charge 188
Sterols 140
Polarity 188
5.2 Dietary Sources 142 Essentiality 190
Recommended Intakes 145
6.2 Sources of Amino Acids 191
5.3 Digestion 145
6.3 Digestion 191
Triacylglycerol Digestion 145
Stomach 191
THE GALLBLADDER 146
Small Intestine 193

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 co n te n ts xi

6.4 Absorption 193 6.11 Catabolism of Tissue/Cell Proteins and

Intestinal Cell Absorption 194 Protein Turnover 243
Extraintestinal Cell Absorption 197 Autophagy-Lysosome Systems 243
6.5 Amino Acid Catabolism 197 Ubiquitin Proteasomal Pathway 244
Transamination of Amino Acids 198 Calpains 245
Deamination of Amino Acids 199 6.12 Changes in Body Mass with Age 246
Disposal of Ammonia 200 Loss of Muscle Mass and Disease 246
Carbon Skeleton/α-Keto Acid Uses 201 6.13 Protein Quality and Protein and Amino
Hepatic Catabolism and Uses of Aromatic Acid Needs 248
Amino Acids 202 Evaluation of Protein Quality 248
Hepatic Catabolism and Uses of Sulfur-Containing Protein Information on Food Labels 251
Amino Acids 205 Assessing Protein and Amino Acid Needs 251
Hepatic Catabolism and Uses of Branched-Chain Recommended Protein and Amino Acid Intakes 252
Amino Acids 209 Protein Deficiency/Malnutrition 254
Hepatic Catabolism and Uses of Basic Summary 255
Amino Acids 209 PERSPECTIVE Stress and Inflammation: Impact on Protein 257
SOME ROLES OF NITRIC OXIDE 211
Hepatic Catabolism and Uses of Other Selected
Amino Acids 212 CHAPTER 7 Integration and Regulation of
6.6 Protein Synthesis 214 Metabolism and the Impact of Exercise 261
Slow versus Fast Proteins 214
7.1 Energy Homeostasis in the Cell 262
Plant versus Animal Proteins 214
Regulatory Enzymes 262
Hormonal Effects 214
mTOR, Intracellular Signaling, and Amino Acids 215 7.2 Integration of Carbohydrate, Lipid, and
Protein Intake, Distribution, and Quantity at Meals 216 Protein Metabolism 266
Interconversion of Fuel Molecules 266
6.7 Protein Structure and Organization 216
Energy Distribution among Tissues 267
6.8 Functional Roles of Proteins 219
7.3 The Fed-Fast Cycle 271
Catalysts 219
The Fed State 271
Messengers 219
The Postabsorptive State 273
Structural Elements 219
The Fasting State 274
Buffers 220
The Starvation State 274
Fluid Balancers 220
Immunoprotectors 220 7.4 Hormonal Regulation of Metabolism 278
Transporters 221 Insulin 278
Acute-Phase Responders 222 HOW IS TYPE 1 DIABETES SIMILAR TO STARVATION? 279

Other Roles 222 Glucagon 280

6.9 Functional Roles of Nitrogen-Containing
Epinephrine 280
Nonprotein Compounds 223
Cortisol 280
Glutathione 223 Growth Hormone 280
Carnitine 223 Adiponectin 281
Creatine 225 7.5 Exercise and Nutrition 281
Carnosine 226 Muscle Function 281
Choline 226 Energy Sources in Resting Muscle 282
Purine and Pyrimidine Bases 227 Muscle ATP Production during Exercise 282
6.10 Interorgan “Flow” of Amino Acids and
Fuel Sources during Exercise 284
Organ-Specific Metabolism 232 Summary 287
Intestinal Cell Amino Acid Metabolism 232 PERSPECTIVE The Role of Dietary Supplements in Sports Nutrition
Amino Acids in the Plasma 233 by Karsten Koehler, PhD 289
Glutamine and the Muscle, Intestine, Liver,
and Kidneys 234
CHAPTER 8 Energy Expenditure, Body
Alanine and the Liver and Muscle 235
Skeletal Muscle Use of Amino Acids 235 Composition, and Healthy Weight 293
Amino Acid Metabolism in the Kidneys 239 8.1 Measuring Energy Expenditure 293
Brain and Accessory Tissues and Amino Acids 241 Direct Calorimetry 294

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xii co n te n ts

Indirect Calorimetry 294 Deficiency: Beriberi 344

Doubly Labeled Water 296 Toxicity 346
HOW TO MEASURE WHAT PEOPLE EAT 297 Assessment of Nutriture 346
8.2 Components of Energy Expenditure 298 9.3 Riboflavin (Vitamin B2) 346
Basal and Resting Metabolic Rate 298 Sources 346
Energy Expenditure of Physical Activity 299 Digestion and Absorption 348
Thermic Effect of Food 300 Transport, Tissue Uptake, and Storage 348
Thermoregulation 301 Functions and Mechanisms of Action 349
8.3 Body Weight: What Should We Weigh? 301 Metabolism and Excretion 351
Ideal Body Weight Formulas 301 Recommended Dietary Allowance 351
Body Mass Index 302 Deficiency: Ariboflavinosis 351
8.4 Measuring Body Composition 303 Toxicity 352
Field Methods 304 Assessment of Nutriture 352
Laboratory Methods 306 9.4 Niacin (Vitamin B3) 352
8.5 Regulation of Energy Balance and Sources 353
Body Weight 307 Digestion and Absorption 354
Hormonal Influences 308 Transport, Tissue Uptake, and Storage 354
Intestinal Microbiota 310 Functions and Mechanisms of Action 355
Environmental Chemicals 310 Metabolism and Excretion 356
Lifestyle Influences 311 Recommended Dietary Allowance 357
Deficiency: Pellagra 357
8.6 Health Implications of Altered Body Weight 311
Toxicity 358
Metabolic Syndrome 311
Assessment of Nutriture 358
Insulin Resistance 312
Weight-Loss Methods 313 9.5 Pantothenic Acid 358
Sources 358
Summary 313
Digestion and Absorption 360
PERSPECTIVE Eating Disorders 315
Transport, Tissue Uptake, and Storage 360
Functions and Mechanisms of Action 360
SECTION III Metabolism and Excretion 363
Adequate Intake 363
THE REGULATORY NUTRIENTS Deficiency: Burning Foot Syndrome 363
Toxicity 363
CHAPTER 9 Water-Soluble Vitamins 321 Assessment of Nutriture 363
DIETARY REFERENCE INTAKES (DRIs) 325 9.6 Biotin (Vitamin B7) 364
DAILY VALUES AND PERCENTAGE DAILY VALUES 326 Sources 364
9.1 Vitamin C (Ascorbic Acid) 326 Digestion, Absorption, Transport, Tissue Uptake,
Sources 327 and Storage 364
Digestion and Absorption 328 Functions and Mechanisms of Action 365
Transport, Tissue Uptake, and Storage 329 Metabolism and Excretion 368
Functions and Mechanisms of Action 329 Adequate Intake 369
Interactions with Other Nutrients 335 Deficiency 369
Metabolism and Excretion 335 Toxicity 369
Recommended Dietary Allowance 335 Assessment of Nutriture 370
Deficiency: Scurvy 336 9.7 Folate (Vitamin B9) 370
Toxicity 337 Sources 370
Assessment of Nutriture 337 Digestion and Absorption 372
9.2 Thiamin (Vitamin B1) 338 Transport, Tissue Uptake, and Storage 372
Sources 338 Functions and Mechanisms of Action 373
Digestion and Absorption 339 Interactions with Other Nutrients 379
Transport, Tissue Uptake, and Storage 339 Association with Diseases 379
Functions and Mechanisms of Action 340 Metabolism and Excretion 380
Metabolism and Excretion 344 Recommended Dietary Allowance 381
Recommended Dietary Allowance 344 Deficiency: Megaloblastic Macrocytic Anemia 381

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 co n te n ts xiii

Toxicity 382 10.3 Vitamin E 435

Assessment of Nutriture 382 Sources 435
9.8 Vitamin B12 (Cobalamin) 383 Digestion and Absorption 437
Sources 384 Transport, Tissue Uptake, and Storage 437
Digestion and Absorption 384 Functions and Mechanisms of Action 438
Transport, Tissue Uptake, and Storage 386 Interactions with Other Nutrients 441
Functions and Mechanisms of Action 386 Metabolism and Excretion 441
Metabolism and Excretion 387 Recommended Dietary Allowance 442
Recommended Dietary Allowance 387 INTERNATIONAL UNITS – VITAMIN E 442
Deficiency: Megaloblastic Macrocytic Anemia Deficiency 442
and Neuropathy 388 Toxicity 443
Toxicity 389 Assessment of Nutriture 443
Assessment of Nutriture 389 10.4 Vitamin K 443
9.9 Vitamin B6 390 Sources 443
Sources 391 Absorption 444
Digestion and Absorption 391 Transport, Tissue Uptake, and Storage 445
Transport, Tissue Uptake, and Storage 391 Functions and Mechanisms of Action 445
Functions and Mechanisms of Action 392 Interactions with Other Nutrients 449
Metabolism and Excretion 395 Metabolism and Excretion 449
Recommended Dietary Allowance 395 Adequate Intake 449
Deficiency 395 Deficiency 449
Toxicity 396 Toxicity 450
Assessment of Nutriture 396 Assessment of Nutriture 450
Summary 397 Summary 451
PERSPECTIVE Types of Human Research Studies and PERSPECTIVE Antioxidant Nutrients, Reactive Species, and
Their Limitations 398 Disease 452

CHAPTER 10 Fat-Soluble Vitamins 401 CHAPTER 11 Major Minerals 463

10.1 Vitamin A and Carotenoids 402 11.1 Calcium 464
Sources 403 Sources 464
Digestion and Absorption 405 Digestion, Absorption, and Transport 465
Transport, Tissue Uptake, and Storage 408 Regulation and Homeostasis 468
Functions and Mechanisms of Action 411 Functions and Mechanisms of Action 470
Interactions with Other Nutrients 419 AN OVERVIEW OF BONE 471
Metabolism and Excretion 419 Interactions with Other Nutrients 474
Recommended Dietary Allowance 420 Excretion 475
INTERNATIONAL UNITS – VITAMIN A 420 Recommended Dietary Allowance 476
Deficiency 420 Deficiency 476
Toxicity 421 Toxicity 477
Assessment of Nutriture 422 Assessment of Nutriture 477
10.2 Vitamin D 423 11.2 Phosphorus 478
Sources 423 Sources 478
Absorption 425 Digestion, Absorption,
Transport, Tissue Uptake, and Storage 425 and Transport 479
Functions and Mechanisms of Action 427 Regulation and Homeostasis 480
Interactions with Other Nutrients 432 Functions and Mechanisms of Action 481
Metabolism and Excretion 432 Excretion 483
Recommended Dietary Allowance 432 Recommended Dietary Allowance 483
Deficiency 432 Deficiency 484
Toxicity 434 Toxicity 484
Assessment of Nutriture 434 Assessment of Nutriture 485

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