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Dietary Reference Intakes (DRI)
The Dietary Reference Intakes (DRI) include two sets of values In addition to the values that serve as goals for nutrient in
that serve as goals for nutrient intake—Recommended Dietary takes (presented in the tables on these two pages), the DRI in
Allowances (RDA) and Adequate Intakes (AI). The RDA reflect clude a set of values called Tolerable Upper Intake Levels (UL).
the average daily amount of a nutrient considered adequate to The UL represent the maximum amount of a nutrient that ap
meet the needs of most healthy people. If there is insufficient pears safe for most healthy people to consume on a regular ba
evidence to determine an RDA, an AI is set. AI are more ten sis. Turn the page for a listing of the UL for selected vitamins
tative than RDA, but both may be used as goals for nutrient and minerals.
intakes. (Chapter 9 provides more details.)
Estimated Energy Requirements (EER), Recommended Dietary Allowances (RDA),

and Adequate Intakes (AI) for Water, Energy, and the Energy Nutrients
ht
ht
g/da cid c
Heig
y)
m 2) BMI
Wei
y)
(g/d te
g/da id
g/da
ay) d
l/da
AI ( lenic A
ay)
RDA ohydra
AI ( leic Ac
AI ( l Fiber
(kg/ rence
cm rence
kg ( rence
y)
y)
y)
y)
y)
AI ( l Fat
(g/k
(kca
(g/d
g/da
g/da
L/da
RDA ein
RDA ein
a
EER b gy
(in)
lb)
AI ( er
Refe
Refe
Refe
C ar b
r
Lino
Lino
Tota
Tota
P ro t
P ro t
Wat
Ene
Age (yr)
Males
0–0.5 — 62 (24) 6 (13) 0.7e 570 60 — 31 4.4 0.5 9.1 1.52
0.5–1 — 71 (28) 9 (20) 0.8f 743 95 — 30 4.6 0.5 11 1.20
1–3g — 86 (34) 12 (27) 1.3 1046 130 19 — 7 0.7 13 1.05
4–8g 15.3 115 (45) 20 (44) 1.7 1742 130 25 — 10 0.9 19 0.95
9–13 17.2 144 (57) 36 (79) 2.4 2279 130 31 — 12 1.2 34 0.95
14–18 20.5 174 (68) 61 (134) 3.3 3152 130 38 — 16 1.6 52 0.85
19–30 22.5 177 (70) 70 (154) 3.7 3067h 130 38 — 17 1.6 56 0.80
31–50 22.5i 177 (70)i 70 (154)i 3.7 3067h 130 38 — 17 1.6 56 0.80
>50 22.5i 177 (70)i 70 (154)i 3.7 3067h 130 30 — 14 1.6 56 0.80
Females
0–0.5 — 62 (24) 6 (13) 0.7e 520 60 — 31 4.4 0.5 9.1 1.52
0.5–1 — 71 (28) 9 (20) 0.8f 676 95 — 30 4.6 0.5 11 1.20
1–3g — 86 (34) 12 (27) 1.3 992 130 19 — 7 0.7 13 1.05
4–8g 15.3 115 (45) 20 (44) 1.7 1642 130 25 — 10 0.9 19 0.95
9–13 17.4 144 (57) 37 (81) 2.1 2071 130 26 — 10 1.0 34 0.95
14–18 20.4 163 (64) 54 (119) 2.3 2368 130 26 — 11 1.1 46 0.85
19–30 21.5 163 (64) 57 (126) 2.7 2403 j 130 25 — 12 1.1 46 0.80
i i i
31–50 21.5 163 (64) 57 (126) 2.7 2403 j 130 25 — 12 1.1 46 0.80
>50 21.5i 163 (64)i 57 (126)i 2.7 2403 j 130 21 — 11 1.1 46 0.80
Pregnancy
1st trimester 3.0 +0 175 28 — 13 1.4 46 0.80
2nd trimester 3.0 +340 175 28 — 13 1.4 71 1.10
3rd trimester 3.0 +452 175 28 — 13 1.4 71 1.10
Lactation
1st 6 months 3.8 +330 210 29 — 13 1.3 71 1.30
2nd 6 months 3.8 +400 210 29 — 13 1.3 71 1.30
h
NOTE: For all nutrients, values for infants are AI. Dashes indicate until age 19. Chapter 8 provides equations and tables to determine For males, subtract 10 kcalories per day for each year of age
that values have not been determined. estimated energy requirements. above 19.
a c i
The water AI includes drinking water, water in beverages, and wa- The linolenic acid referred to in this table and text is the omega-3 Because weight need not change as adults age if activity is main-
ter in foods; in general, drinking water and other beverages contrib- fatty acid known as alpha-linolenic acid. tained, reference weights for adults 19 through 30 years are applied
d
ute about 70 to 80 percent, and foods, the remainder. Conversion The values listed are based on reference body weights. to all adult age groups.
e j
factors: 1 L = 33.8 fluid oz; 1 L = 1.06 qt; 1 cup = 8 fluid oz. Assumed to be from human milk. For females, subtract 7 kcalories per day for each year of age
b f
The Estimated Energy Requirement (EER) represents the average Assumed to be from human milk and complementary foods and above 19.
dietary energy intake that will maintain energy balance in a healthy beverages. This includes approximately 0.6 L (∼21⁄2 cups) as total
person of a given gender, age, weight, height, and physical activity fluid including formula, juices, and drinking water. SOURCE: Adapted from the Dietary Reference Intakes series, National
g
level. The values listed are based on an “active” person at the refer- For energy, the age groups for young children are 1–2 years and Academies Press. Copyright 1997, 1998, 2000, 2001, 2002, 2004,
ence height and weight and at the midpoint ages for each group 3–8 years. 2005, 2011 by the National Academies of Sciences.
A Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203
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Recommended Dietary Allowances (RDA) and Adequate Intakes (AI) for Vitamins
day) acid
/day e
/day a
/day b
/day c
day) d
)
)
)
)
/day
/day
/day
/day
c
/day
(µg 12
day)
AI ( otheni
ay)
ay)
(mg 6
RDA flavin
RDA in A
RDA min D
RDA min B
RDA min B
AI ( min K
RDA min C
RDA min E
(IU/
RDA min
(mg
(mg
(mg
(mg
(mg
µg/d
µg/d
AI ( ine
(µg
(µg
mg/
mg/
RDA in
RDA e
n
m
t
i
Ribo
l
Niac
Thia
Biot
Fola
Vita
Vita
Vita
Vita
Vita
Vita
Vita
Cho
Pan
AI (
Age (yr)
Infants
0–0.5 0.2 0.3 2 5 1.7 0.1 65 0.4 125 40 400 400 (10 µg) 4 2.0
0.5–1 0.3 0.4 4 6 1.8 0.3 80 0.5 150 50 500 400 (10 µg) 5 2.5
Children
1–3 0.5 0.5 6 8 2 0.5 150 0.9 200 15 300 600 (15 µg) 6 30
4–8 0.6 0.6 8 12 3 0.6 200 1.2 250 25 400 600 (15 µg) 7 55
Males
9–13 0.9 0.9 12 20 4 1.0 300 1.8 375 45 600 600 (15 µg) 11 60
14–18 1.2 1.3 16 25 5 1.3 400 2.4 550 75 900 600 (15 µg) 15 75
19–30 1.2 1.3 16 30 5 1.3 400 2.4 550 90 900 600 (15 µg) 15 120
31–50 1.2 1.3 16 30 5 1.3 400 2.4 550 90 900 600 (15 µg) 15 120
51–70 1.2 1.3 16 30 5 1.7 400 2.4 550 90 900 600 (15 µg) 15 120
>70 1.2 1.3 16 30 5 1.7 400 2.4 550 90 900 800 (20 µg) 15 120
Females
9–13 0.9 0.9 12 20 4 1.0 300 1.8 375 45 600 600 (15 µg) 11 60
14–18 1.0 1.0 14 25 5 1.2 400 2.4 400 65 700 600 (15 µg) 15 75
19–30 1.1 1.1 14 30 5 1.3 400 2.4 425 75 700 600 (15 µg) 15 90
31–50 1.1 1.1 14 30 5 1.3 400 2.4 425 75 700 600 (15 µg) 15 90
51–70 1.1 1.1 14 30 5 1.5 400 2.4 425 75 700 600 (15 µg) 15 90
>70 1.1 1.1 14 30 5 1.5 400 2.4 425 75 700 800 (20 µg) 15 90
Pregnancy
≤18 1.4 1.4 18 30 6 1.9 600 2.6 450 80 750 600 (15 µg) 15 75
19–30 1.4 1.4 18 30 6 1.9 600 2.6 450 85 770 600 (15 µg) 15 90
31–50 1.4 1.4 18 30 6 1.9 600 2.6 450 85 770 600 (15 µg) 15 90
Lactation
≤18 1.4 1.6 17 35 7 2.0 500 2.8 550 115 1200 600 (15 µg) 19 75
19–30 1.4 1.6 17 35 7 2.0 500 2.8 550 120 1300 600 (15 µg) 19 90
31–50 1.4 1.6 17 35 7 2.0 500 2.8 550 120 1300 600 (15 µg) 19 90
c
NOTE: For all nutrients, values for infants are AI. Vitamin A recommendations are expressed as retinol activity equivalents (RAE).
a d
Niacin recommendations are expressed as niacin equivalents (NE), except for recommendations for infants Vitamin D recommendations are expressed as cholecalciferol and assume an absence of adequate exposure
younger than 6 months, which are expressed as preformed niacin. to sunlight.
b e
Folate recommendations are expressed as dietary folate equivalents (DFE). Vitamin E recommendations are expressed as α-tocopherol.
Recommended Dietary Allowances (RDA) and Adequate Intakes (AI) for Minerals
)
)
(µg m
)
)
/day
/day
/day
/day
/day
/day
/day
/day
/day
(mg us
day)
day)
day)
day)
day)
RDA esium
AI ( ganese
ay)
RDA bdenu
AI ( ssium
AI ( mium
RDA nium
or
AI ( r ide
AI ( r ide
(mg
RDA ium
µg/d
(mg
(mg
(mg
AI ( um
(µg
(µg
(µg
sph
mg/
mg/
mg/
mg/
mg/
per
n e
n
y
Chro
Pota
Zinc
i
Chlo
Fluo
Sele
Iron
Iodi
Man
Calc
Mag
Pho
Cop
S od
Mol
RDA
RDA
RDA
RDA
RDA
Age (yr)
Infants
0–0.5 120 180 400 200 100 30 0.27 2 110 15 200 0.003 0.01 0.2 2
0.5–1 370 570 700 260 275 75 11 3 130 20 220 0.6 0.5 5.5 3
Children
1–3 1000 1500 3000 700 460 80 7 3 90 20 340 1.2 0.7 11 17
4–8 1200 1900 3800 1000 500 130 10 5 90 30 440 1.5 1.0 15 22
Males
9–13 1500 2300 4500 1300 1250 240 8 8 120 40 700 1.9 2 25 34
14–18 1500 2300 4700 1300 1250 410 11 11 150 55 890 2.2 3 35 43
19–30 1500 2300 4700 1000 700 400 8 11 150 55 900 2.3 4 35 45
31–50 1500 2300 4700 1000 700 420 8 11 150 55 900 2.3 4 35 45
51–70 1300 2000 4700 1000 700 420 8 11 150 55 900 2.3 4 30 45
>70 1200 1800 4700 1200 700 420 8 11 150 55 900 2.3 4 30 45
Females
9–13 1500 2300 4500 1300 1250 240 8 8 120 40 700 1.6 2 21 34
14–18 1500 2300 4700 1300 1250 360 15 9 150 55 890 1.6 3 24 43
19–30 1500 2300 4700 1000 700 310 18 8 150 55 900 1.8 3 25 45
31–50 1500 2300 4700 1000 700 320 18 8 150 55 900 1.8 3 25 45
51–70 1300 2000 4700 1200 700 320 8 8 150 55 900 1.8 3 20 45
>70 1200 1800 4700 1200 700 320 8 8 150 55 900 1.8 3 20 45
Pregnancy
≤18 1500 2300 4700 1300 1250 400 27 12 220 60 1000 2.0 3 29 50
19–30 1500 2300 4700 1000 700 350 27 11 220 60 1000 2.0 3 30 50
31–50 1500 2300 4700 1000 700 360 27 11 220 60 1000 2.0 3 30 50
Lactation
≤18 1500 2300 5100 1300 1250 360 10 13 290 70 1300 2.6 3 44 50
19–30 1500 2300 5100 1000 700 310 9 12 290 70 1300 2.6 3 45 50
31–50 1500 2300 5100 1000 700 320 9 12 290 70 1300 2.6 3 45 50
NOTE: For all nutrients, values for infants are AI.
Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203 B
Tolerable Upper Intake Levels (UL) for Vitamins
/day 6
(µg min A
(IU/ min D
(mg min B
(mg min C
(mg min E
/day a
/day c
/day b
/day a
)
)
)
day)
(mg ine
/day
/day
(mg in
(µg te
l
Niac
Fola
Vita
Vita
Vita
Vita
Vita
Cho
Age (yr)
Infants
0–0.5 — — — — — 600 1000 (25 µg) —
0.5–1 — — — — — 600 1500 (38 µg) —
Children
1–3 10 30 300 1000 400 600 2500 (63 µg) 200
4–8 15 40 400 1000 650 900 3000 (75 µg) 300
9–13 20 60 600 2000 1200 1700 4000 (100 µg) 600
Adolescents
14–18 30 80 800 3000 1800 2800 4000 (100 µg) 800
Adults
19–70 35 100 1000 3500 2000 3000 4000 (100 µg) 1000
>70 35 100 1000 3500 2000 3000 4000 (100 µg) 1000
Pregnancy
≤18 30 80 800 3000 1800 2800 4000 (100 µg) 800
19–50 35 100 1000 3500 2000 3000 4000 (100 µg) 1000
Lactation
≤18 30 80 800 3000 1800 2800 4000 (100 µg) 800
19–50 35 100 1000 3500 2000 3000 4000 (100 µg) 1000
a c
The UL for niacin and folate apply to synthetic forms obtained from The UL for vitamin E applies to any form of supplemental
supplements, fortified foods, or a combination of the two. α-tocopherol, fortified foods, or a combination of the two.
b
The UL for vitamin A applies to the preformed vitamin only.
Tolerable Upper Intake Levels (UL) for Minerals
/day num
(mg phorus
(mg nesium
(mg ganese
(mg dium
/day d
(µg nium
(mg r ide
(mg r ide
e
)
)
(mg ium
)
(mg um
/day
/day
/day
/day
/day
/day
/day
/day
/day
/day
/day
ybd
(µg er
/day
/day
/day
(µg ne
(mg el
(mg n
p
a
s
Boro
Nick
i
Chlo
Fluo
Sele
Iron
Zinc
Iodi
Man
Calc
Mag
Pho
Van
Cop
S od
Mol
(mg
(mg
(µg
Age (yr)
Infants
0–0.5 — — 1000 — — 40 4 — 45 — — 0.7 — — — —
0.5–1 — — 1500 — — 40 5 — 60 — — 0.9 — — — —
Children
1–3 1500 2300 2500 3000 65 40 7 200 90 1000 2 1.3 300 3 0.2 —
4–8 1900 2900 2500 3000 110 40 12 300 150 3000 3 2.2 600 6 0.3 —
9–13 2200 3400 3000 4000 350 40 23 600 280 5000 6 10 1100 11 0.6 —
Adolescents
14–18 2300 3600 3000 4000 350 45 34 900 400 8000 9 10 1700 17 1.0 —
Adults
19–50 2300 3600 2500 4000 350 45 40 1100 400 10,000 11 10 2000 20 1.0 1.8
51–70 2300 3600 2000 4000 350 45 40 1100 400 10,000 11 10 2000 20 1.0 1.8
>70 2300 3600 2000 3000 350 45 40 1100 400 10,000 11 10 2000 20 1.0 1.8
Pregnancy
≤18 2300 3600 3000 3500 350 45 34 900 400 8000 9 10 1700 17 1.0 —
19–50 2300 3600 2500 3500 350 45 40 1100 400 10,000 11 10 2000 20 1.0 —
Lactation
≤18 2300 3600 3000 4000 350 45 34 900 400 8000 9 10 1700 17 1.0 —
19–50 2300 3600 2500 4000 350 45 40 1100 400 10,000 11 10 2000 20 1.0 —
d
The UL for magnesium applies to synthetic forms obtained from supplements or drugs only. SOURCE: Adapted with permission from the Dietary Reference Intakes series, National Academies Press.
NOTE: An Upper Limit was not established for vitamins and minerals not listed and for those age groups Copyright 1997, 1998, 2000, 2001, 2002, 2005, 2011 by the National Academies of Sciences.
listed with a dash (—) because of a lack of data, not because these nutrients are safe to consume at any
level of intake. All nutrients can have adverse effects when intakes are excessive.
C Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203
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ADVANCED NUTRITION
AND HUMAN METABOLISM
EIGHTH EDITION
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
This is an electronic version of the print textbook. Due to electronic rights restrictions,
some third party content may be suppressed. Editorial review has deemed that any suppressed
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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
may be reproduced or distributed in any form or by any means, except as
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Printed in the United States of America

Print Number: 01 Print Year: 2020
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
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
Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203 vii
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
Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203 ix
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
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
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
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
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
xiv co n te n ts
11.3 Magnesium 485 Respiratory Regulation 519

Sources 485 Renal Regulation 520
Digestion, Absorption, and Transport 486 Summary 521
Regulation and Homeostasis 487 PERSPECTIVE Macrominerals and Hypertension 522
Interactions with Other Nutrients 489
Excretion 489 CHAPTER 13 Essential Trace and Ultratrace
Recommended Dietary Allowance 489
Deficiency 489 Minerals 525
Toxicity 491 13.1 Iron 525
Assessment of Nutriture 491 Sources 526
Summary 492 Digestion, Absorption, Transport,
PERSPECTIVE Osteoporosis and Diet 493 and Storage 528
Turnover 540
CHAPTER 12 Water and Electrolytes 499 Interactions with Other Nutrients 541
Excretion 542
12.1 Water Functions 499
12.3 Body Water Content and Distribution 500
Deficiency 542
12.3 Water Losses, Sources, and Absorption 501 Toxicity 544
12.4 Recommended Water Intake 501 Assessment of Nutriture 544
12.5 Water (Fluid) and Sodium Balance 502 13.2 Zinc 546
Osmotic Pressure 502 Sources 546
Hydrostatic (Fluid/Capillary) Pressure 503 Digestion, Absorption, Transport,
Colloidal Osmotic Pressure 504 and Storage 547
Extracellular Fluid Volume and Osmolarity and Functions and Mechanisms of Action 551
Hormonal Controls 504 Interactions with Other Nutrients 554
THE KIDNEYS: A BRIEF REVIEW 505 Excretion 555
12.6 Sodium 508 Recommended Dietary Allowance 555
Sources 508 Deficiency 555
ELECTROLYTES: CALCULATING MILLIEQUIVALENTS (MEQ) 509 Toxicity 556
Absorption and Transport 510 Assessment of Nutriture 556
Functions and Interactions with Other Nutrients 511 13.3 Copper 557
Excretion 511 Sources 557
Recommendations, Deficiency, Toxicity, and Assessment Digestion, Absorption, Transport, and Storage 557
of Nutriture 511 Functions and Mechanisms of Action 560
12.7 Potassium 512 Interactions with Other Nutrients 562
Sources 512 Excretion 563
Absorption, Secretion, and Transport 512 Recommended Dietary Allowance 564
Functions and Interactions with Other Nutrients 513 Deficiency 564
Excretion 513 Toxicity 565
Recommendations, Deficiency, Toxicity, and Assessment Assessment of Nutriture 565
of Nutriture 513 13.4 Selenium 566
12.8 Chloride 514 Sources 566
Sources 514 THE SHIFTING SANDS OF SELENIUM 567
Absorption, Secretion, and Transport 514 Digestion, Absorption, Transport, and Storage 568
Functions 515 Metabolism 568
Excretion 515 Functions and Mechanisms of Action 570
Recommendations, Deficiency, Toxicity, and Assessment Interactions with Other Nutrients 572
of Nutriture 516 Excretion 573
12.9 Acid–Base Balance: Control of Hydrogen Ion Recommended Dietary Allowance 573
Concentration 516 Deficiency 573
Chemical Buffer Systems 517 Toxicity 574
PRINCIPLES OF BUFFERS 517
co n te n ts xv
13.5 Chromium 575 Assessment of Nutriture 590

Sources 575 PERSPECTIVE Nutrient–Drug Interactions 591
Digestion, Absorption, Transport,
and Storage 575
Functions and Mechanisms of Action 576 CHAPTER 14 Nonessential Trace and Ultratrace
Excretion 577 Minerals 595
Adequate Intake 577
14.1 Fluoride 595
Deficiency 577
Sources 595
Toxicity 577
Absorption, Transport, Tissue Uptake, Storage,
and Excretion 597
13.6 Iodine 578
Functions and Deficiency 597
Sources 578
Recommended Intake, Toxicity, and Assessment
Digestion, Absorption, Transport, and Storage 579
of Nutriture 598
14.2 Boron 598
Sources 598
Excretion 582
Absorption, Transport, Tissue Uptake, Storage,
and Excretion 599
Deficiency 582
Toxicity 583
of Nutriture 600
13.7 Manganese 584
14.3 Silicon 600
Sources 584
Sources 600
Digestion, Absorption, Transport,
Absorption, Transport, Storage, and Excretion 601
and Storage 584
of Nutriture 601
Excretion 586
Adequate Intake 586 14.4 Vanadium 602
Deficiency 586 Sources 602
Toxicity 586 Absorption, Transport, Storage, and Excretion 602
Assessment of Nutriture 586 Functions and Deficiency 602
13.8 Molybdenum 587
of Nutriture 603
Sources 587
Digestion, Absorption, Transport, 14.5 Cobalt 603
and Storage 587 Summary 604
Functions and Mechanisms of Action 587 PERSPECTIVE No, Silver Is Not Another Essential Ultratrace
Interactions with Other Nutrients 589 Mineral: Tips to Identifying Bogus Claims and Selecting Dietary
Excretion 590 Supplements 605
Deficiency 590 Glossary 609
Toxicity 590 Index 615
PREFACE
S Chapter 2 The Digestive System:

ince the first edition was published in 1990, much
has changed in the science of nutrition. But the pur-
pose of the text—to provide thorough coverage of
Mechanism for Nourishing the Body
normal metabolism for upper-division undergraduate and ●● Expanded information on the structural features of the
graduate students majoring in nutrition or other health- small intestine
related fields—remains the same. We continue to strive for ●● Added new information on probiotics and intestinal
a level of detail and scope of material that satisfy the needs
conditions
of both instructors and students. With each succeeding
edition, we have responded to suggestions from instruc-
tors, content reviewers, and students that have improved Chapter 3 Carbohydrates
the text by enhancing the clarity of the material and by
ensuring accuracy. In addition, we have included the latest ●● Reorganized the chapter sections to improve flow and
and most pertinent nutrition science available to provide readability
future nutrition professionals with the fundamental infor- ●● Revised sections on stereoisomers, ring structures, and
mation vital to their careers and to provide the basis for derivatives of monosaccharides
assimilating new scientific discoveries. ●● Added new information related to dextrins and dex-
Just as the body of information on nutrition science trose equivalents
has increased, so has the team of authors working on this
●● Added new information on SGLTs and GLUTs
text. Dr. James Groff and Dr. Sara Hunt coauthored the
first edition. In subsequent editions, Dr. Sareen Grop- ●● Expanded sections on blood–tissue barriers and the
per became a coauthor as Dr. Hunt entered retirement. electron transport chain
In the fourth edition, Dr. Jack L. Smith joined the author ●● Reorganized sections related to carbohydrate absorp-
team now led by Dr. Gropper. In the seventh and eighth tion and transport; added new discussion on membrane
editions, Dr. Tim Carr has provided additional expertise transport
and coauthorship on several chapters following Dr. Smith’s ●● Revised section on metabolic regulation; added new
retirement. information on enzyme translocation
●● Updated and modified several figures and figure
legends
NEW TO THIS EDITION ●● Added new Box feature on syrups
All chapters of the eighth edition have been updated and ●● Added new Box feature on uncoupling oxidative
feature new or enhanced tables and illustrations. The orga- phosphorylation
nization of the content among the chapters has remained ●● Updated the end-of-chapter Perspective
similar to the previous editions.
Chapter 4 Fiber
Chapter 1 The Cell: A Microcosm of Life
●● Added new information on another form of resistant
●● Expanded content in several sections including, for starch
example, the nucleus where additional information is ●● Provided new information on the properties of fiber
presented on genes and chromosomes important for laxation
●● Added additional information on mechanism of apoptosis ●● Added information on a new mechanism by which phy-
●● Created new Perspective on Nutritional Genomics tochemicals may regulate mRNA translation
Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203 xvii
xviii P reface
Chapter 5 Lipids ●● Added new Box feature of how to measure what people
eat
●● Added new section on odd-chain and branched-chain
fatty acids
●● Expanded discussion related to conjugated linoleic acid Chapter 9 Water-Soluble Vitamins
●● Revised information related to trans fatty acids, ●● Updated daily values
mono- and diacylglycerols, and the biological roles of ●● Added photos showing the physical manifestations of
phospholipids several vitamin deficiencies
●● Updated information on fatty acid transport into ●● Added information on another coenzyme role of thia-
enterocytes min tied to fatty acid oxidation
●● Added new section on the lipid hypothesis ●● Expanded the figures showing pantothenic acid
●● Expanded discussion on β-oxidation, including metabolism
new sections related to oxidation of odd-chain and ●● Added new information on the functions of panto-
branched-chain fatty acids thenic acid linking it to folate metabolism
●● Updated and modified several figures and figure ●● Expanded the information on the non-coenzyme roles
legends of biotin
●● Added new Box feature on the gallbladder ●● Added a new figure showing folate metabolism within
●● Added new Box feature on coconut oil the cytosol, nucleus, and mitochondria
●● Expanded the discussion of intracellular chaperones
involved in vitamin B12 transport
Chapter 6 Protein
●● Added information on the Dietary Reference Intakes,
●● Added a new figure showing intestinal amino acid including chronic disease risk reduction
transport ●● Developed a new Perspective addressing types of
●● Expanded the discussion addressing the mechanisms of research study designs
protein degradation
●● Expanded the discussion on the need for protein with
aging Chapter 10 Fat-Soluble Vitamins
●● Expanded the section addressing plant proteins ●● Updated daily values
●● Provided more details on the mechanisms of absorption
Chapter 7 Integration and of vitamins A, E, and K
Regulation of Metabolism and ●● Added a new figure and information on the functions
and metabolism of vitamin E
the Impact of Exercise ●● Added new figures showing some manifestations of
●● Revised Table 7.1 deficiencies of vitamins A and D
●● Added new section on adiponectin; updated Table 7.3 ●● Added a new figure showing phylloquinone metabolism
to include adiponectin ●● Created a new table providing the phylloquinone and
●● Revised figures and figure legends menaquinone contents of foods
●● Added new Box feature on the metabolic similarity ●● Expanded information on the carotenoid content of foods
between type 1 diabetes and starvation
Chapter 11 Major Minerals

Chapter 8 Energy Expenditure, Body
●● Expanded discussion of calcium functions
Composition, and Healthy Weight ●● Expanded the discussion providing an overview of
●● Added new information on the origin of body mass bone
index ●● Expanded discussions of calcium, phosphate, and mag-
●● Revised section related to adiponectin regulation nesium homeostasis
●● Added new and updated information in the end-of-chapter ●● Added a new table showing factors regulating serum
Perspective on eating disorders phosphate
●● Revised figures and figure legends ●● Improved figure depicting phosphate absorption
P reface xix
●● Added information on the use of topical magnesium and helps generate reader interest. The Perspectives pro-
oils vide applications or expansion of the information in the
●● Updated daily values chapter text.
Because this book focuses on normal human nutrition
and physiological function, it is an effective resource for
Chapter 12 Water and Electrolytes students majoring in either nutrition sciences or dietetics
and for other health care professionals enrolled in a gradu-
●● Expand list of dietary sources for sodium and potassium ate nutrition course. Intended for a course in advanced
●● Added a table with food sources of potassium nutrition, the text presumes a sound background in the
●● Updated recommendations and daily values biological sciences. At the same time, however, it provides
●● Added information on the chronic disease risk reduc- a review of the basic sciences, particularly biochemistry
tion recommendation for sodium and physiology, which are important to understanding
the material. This text applies biochemistry to nutrient
●● Updated the Perspective on macrominerals and hyper- use from consumption through digestion, absorption,
tension to reflect the latest dietary recommendations distribution, and cellular metabolism. Health practitio-
ners may find that the book is a useful resource to refresh
Chapter 13 Essential Trace their memories with regard to metabolic and physiological
interrelationships and to obtain a concise update on cur-
and Ultratrace Minerals rent concepts related to human nutrition.
●● Revised tables showing food sources of trace minerals We continue to present nutrition as the science that
integrates life processes from the molecular to the cellu-
●● Expanded section on iron as a pro-oxidant
lar level and on through the multisystem operation of the
●● Added new information describing zinc/cancer whole organism. Our primary goal is to give a comprehen-
association sive picture of cell reactions at the tissue, organ, and system
●● Added new information on mercury/selenium levels. Subject matter has been selected for its relevance to
interaction meeting this goal.
●● Updated the Daily Value for each mineral
●● Added photos showing deficiency symptoms of zinc,
copper, selenium, and iodine ORGANIZATION
●● Revised figures and figure legends Each of the 14 chapters begins with a topic outline, fol-
●● Added new Box feature on selenium in the lowed by a brief introduction to the chapter’s subject mat-
environment ter. These features are followed in order by the chapter text,
a brief summary that ties together the ideas presented in
the chapter, a reference list, and a Perspective with its own
Chapter 14 Nonessential Trace reference list.
and Ultratrace Minerals The text is divided into three sections. Section I (Chap-
ters 1 and 2) focuses on cell structure, gastrointestinal
●● Expanded the sections on the sources of fluoride and
tract anatomy, and function with respect to digestion and
supplemental forms of boron
absorption.
●● Expanded the discussions addressing fluoride’s and Section II (Chapters 3–8) discusses metabolism of the
boron’s mechanisms of action macronutrients. This section reviews primary metabolic
●● Updated information on recommendations for intake pathways for carbohydrates, lipids, and proteins, empha-
of boron sizing those reactions particularly relevant to issues of
●● Updated information on toxicity-related concerns with health. Since most of the body’s energy production is
vanadium associated with glycolysis or the tricarboxylic acid cycle
by way of the electron transport chain and oxidative phos-
●● Expanded the Perspective to include information to
phorylation, the carbohydrates chapter (Chapter 3) cov-
consider when buying supplements
ers these aspects of energy transformation. We include a
separate chapter (Chapter 4) on fiber. The metabolism of
alcohol, which contributes to the caloric intake of many
PRESENTATION people, is discussed within the lipids chapter (Chapter 5).
Alcohol’s chemical structure more closely resembles that
The presentation of the text is designed to make the book of carbohydrates, but its metabolism is more similar to
easy for the reader to use. The added color(s) draws atten- that of lipids. Chapter 7 discusses the interrelationships
tion to important elements in the text, tables, and figures among the metabolic pathways that are common to the
xx P reface
macronutrients. This chapter also includes a discussion of the product manager, Courtney Heilman; our art direc-
the regulation of the metabolic pathways and a description tor, Lizz Anderson; our marketing manager, Shannon
of the metabolic dynamics of the fed-fast cycle, along with Hawkins; our content manager, Samantha Rundle; and
a presentation of the effects of physical exertion on the our permissions analysts, Ann Hoffman. We extend spe-
body’s metabolic pathways. Chapter 8 focuses on energy cial thanks to our production team and our copy editor,
expenditure, energy balance, and healthy weight and also Laura Specht Patchkofsky.
includes a brief discussion of measuring body composition We appreciate the writing contribution of Karsten
and the health implications of altered body weight. Koehler, PhD, for the Perspective “The Role of Dietary
Section III (Chapters 9–14) concerns those nutrients Supplements in Sports Nutrition.”
considered regulatory in nature: the water- and fat-soluble We owe special thanks to the reviewers whose thought-
vitamins and the minerals, including the major minerals, ful comments, criticisms, and suggestions were indispens-
trace minerals, and ultratrace minerals. These chapters able in shaping this text.
cover nutrient features such as digestion, absorption,
transport, function, metabolism, excretion, deficiency,
toxicity, and assessment of nutriture, as well as the latest Eighth Edition Reviewers
Recommended Dietary Allowances or Adequate Intakes Michael Crosier, Framingham State University
for each nutrient. Information about the major minerals Janet Colson, Middle Tennessee State University
has been split into two chapters: Chapter 11 addresses La-Tonya J. Dixon, Alabama A&M University
calcium, phosphorus, and magnesium, and Chapter 12 Erika Ireland, California State University, Fresno
discusses sodium, potassium, and chloride. Chapter Jennifer Farrell, Florida State University
12 integrates coverage of the maintenance of the body’s Long Wang, California State University, Long Beach
homeostatic environment—including discussions of body Norma L. Dawkins, Tuskegee University
fluids, electrolyte balance, and pH maintenance—with the
presentation of the electrolytes.
Seventh Edition Reviewers
Michael E. Bizeau, Metropolitan State University
SUPPLEMENTARY MATERIAL of Denver
Janet Colson, Middle Tennessee State University
MindTap for Gropper's Advanced Nutrition and Human Michael Crosier, Framingham State University
Metabolism, 8th Edition, is a digital learning solution J. Andrew Doyle, Georgia State University
that empowers learners to go beyond memorization— Elizabeth A. Kirk, Bastyr University
enabling a deeper understanding of concepts and topics. Kevin L. Schalinske, Iowa State University
MindTap provides engaging content and activities that Long Wang, California State University, Long Beach
help build student confidence. Accelerate progress with
MindTap. Visit cengage.com/login to learn more. Addi-
tional instructor resources for this product are available Sixth Edition Reviewers
online. Instructor assets include an Instructor’s Manual, Jodee L. Dorsey, Florida State University
Educator’s Guide, PowerPoint® slides, and a test bank Jennifer Hemphill, Florida State University
powered by Cognero®. Sign up or sign in at www.cengage Elizabeth A. Kirk, Bastyr University and University
.com to search for and access this product and its online of Washington
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Scott K. Reaves, California Polytechnic State University,
San Luis Obispo
ACKNOWLEDGMENTS Karla P. Shelnutt, University of Florida
Although this textbook represents countless hours of

work by the authors, it is also the work of many other
Fifth Edition Reviewers
hardworking individuals. We cannot possibly list every- Richard C. Baybutt, Kansas State University
one who has helped, but we would like to call attention Patricia B. Brevard, James Madison University
to a few individuals who have played particularly impor- Marie A. Caudill, California Polytechnic State University,
tant roles. We thank our undergraduate and graduate Pomona
nutrition students for their ongoing feedback. We thank Prithiva Chanmugam, Louisiana State University
P reface xxi
Michele M. Doucette, Georgia State University William R. Proulx, State University of New York, Oneonta
Michael A. Dunn, University of Hawaii at Mānoa Scott K. Reaves, California Polytechnic State University,
Steve Hertzler, Ohio State University San Luis Obispo
Steven Nizielski, Grand Valley State University Donato F. Romagnolo, University of Arizona, Tucson
Kimberli Pike, Ball State University James H. Swain, Case Western Reserve University
THE CELL:
1
A MICROCOSM OF LIFE
LEARNING OBJECTIVES
1.1 Identify cellular components and their functions.
1.2 Describe the roles of cell receptors and enzymes.
1.3 Explain the mechanisms by which enzymatic reactions are regulated.
1.4 Discuss the need for and pathways involved in apoptosis.
1.5 Describe how energy is released and utilized in chemical reactions.
C
ELLS ARE THE VERY ESSENCE OF LIFE. Cells may be defined as the basic
living, structural, and functional units of the human body. They vary
greatly in size, chemical composition, and function, but each one is a
remarkable miniaturization of human life. Cells move, grow, ingest “food,”
excrete wastes, react to their environment, and reproduce. This chapter
provides a brief review of the basics of a cell, including cellular components,
biological energy, and an overview of a cell’s natural life span.
Cells of multicellular organisms are called eukaryotic cells (from the Greek
eu meaning “true” and karyon meaning “nucleus”). Eukaryotic cells evolved
from simpler, more primitive cells called prokaryotic cells (from the Greek
meaning “before nucleus”). One distinguishing feature between the two cell
types is that eukaryotic cells possess a defined nucleus, whereas prokaryotic
cells do not. Also, eukaryotic cells are larger and much more complex structur-
ally and functionally than their ancestors. Because this text addresses human
metabolism and nutrition, all descriptions of cellular structure and function in
this and subsequent chapters pertain to eukaryotic cells.
While specialization among cells is necessary for life, cells, in general, have
certain basic similarities. All human cells have a plasma membrane and a
nucleus (or have had a nucleus), and most contain an endoplasmic reticulum,
Golgi apparatus, and mitochondria. For convenience of discussion, a “typical
cell” is presented (Figure 1.1) to enable the identification of the various organ-
elles and their functions, which characterize cellular life. Our discussion begins
with the plasma membrane, which forms the outer boundary of the cell, and
then moves inward to examine the organelles found within the cell.
1.1 COMPONENTS OF CELLS

Plasma Membrane
The plasma membrane is a sheetlike structure that encapsulates and surrounds
the cell, allowing it to exist as a distinct unit. The plasma membrane, like other
membranes within the cell, has distinct structural characteristics and functions.
Copyright 2022 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-203 1
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offered him entertainment, which he scrupulously refused. ‘Then I
went to my lord and said, I was sent by God to cast the devil out of
his son, by faith in Christ. He seemed to be like that lord who had the
charge of the gate in Samaria. Then I said to him: “My lord, do you
not believe me?” Then he bade me go and 1720.
speak to many ministers that was near by
him; but I said I was not sent to them. Then he went to them himself,
and spoke to them what I said; but they would not hear of it; so I
went to three witches and a warlock, to examine them, in sundry
places. Two of them denied, and two of them confessed. I have no
time to relate here all that I said to them, and what they said; but I
asked them, “When they took on in that service?” The wife said:
“Many years;” and the man said: “It was ten years to him.” Then I
asked the wife: “What was her reason for taking on with the devil?”
And she said: “He promised her riches, and she believed him.” Then
she called him many a cheat and liar in my hearing. Then I went to
the man, because he was a great professor, and could talk of religion
with any of the parish, as they that was his neighbours said, and he
was at Bothwell Bridge fighting against the king; and because of that,
I desired to ask questions at him; but my lord’s officer said: “His lord
would not allow me.” I said I would not be hindered neither by my
lord nor by the devil, before many there present. Then I asked:
“What iniquity he found in God, that he left his service?” He got up
and said: “Oh, sir, are ye a minister?” So ye see the devil knows me to
be a minister better than the magistrates. He said: “He found no fault
in God; but his wife beguiled him;” and he said: “Wo be to the
woman his wife!” and blamed her only, as Adam did his wife, and the
woman blamed the devil; so ye see it is from the beginning. This is a
caution to us all never to hearken to our wives except they have
Scripture on their side. Then I asked at him: “Did he expect heaven?”
“Yes,” said he. Then I asked at him if he could command the devil to
come and speak to me? He said: “No.” Then I said again: “Call for
him, that I may speak with him.” He said again: “It was not in his
power.” Then my lord sent more servants, that hindered me to ask
any more questions, otherwise I might have seen the devil, and I
would have spoken about his son.’[539]
On this fast-day, a sermon was preached in Calder kirk by the Rev.
Mr John Wilkie, minister of Uphall, the alleged sorcerers being all
present. Lord Torphichen subsequently caused the discourse to be
printed. His boy in time recovered, and going to sea, rose by merit to
the command of an East Indiaman, but perished in a storm. It brings
us strangely near to this wild-looking affair, 1720.
that the present tenth Lord Torphichen
(1860) is only nephew to the witch-boy of Calder.
The exportation of some corn from Feb. 5.

Dundee being connected unfavourably in
the minds of the populace with a rise of the markets, a tumult took
place, with a view to keeping the grain within the country. The mob
not only took possession of two vessels loaded with bear lying in the
harbour, the property of Mr George Dempster, merchant, but
attacked and gutted the house, shops, cellars, and lofts of that
gentleman, carrying off everything of value they contained, including
twelve silver spoons, a silver salver, and two silver boxes, one of
them containing a gold chain and twelve gold rings, some hair ones,
and others set with diamonds. Dempster advertised that whoever
shall discover to him ‘the havers of his goods,’ should have ‘a
sufficient reward and the owner’s kindness, and no questions
asked.’[540]
A similar affair took place at Dundee nine years later. The country-
people in and about the town then ‘carried their resentment so high
against the merchants for transporting of victual, that they furiously
mobbed them, carried it out of lofts, and cut the sacks of those that
were bringing it to the barks.’[541]
Died, Alexander Rose, who had been Mar. 20.

appointed Bishop of Edinburgh just before
the Revolution. He was the last survivor of the unfortunate
episcopate of Scotland, and also outlived all the English bishops who
forfeited their sees at the Revolution. Though strenuous during all
these thirty-one years as a nonjuror—for which in 1716 he was
deprived of a pension assigned him by Queen Anne—he is testified to
by his presbyter Robert Keith as ‘a sweet-natured man.’[542] His
aspect, latterly, was venerable, and the gentleness of his life secured
him the respect of laymen of all parties. Descended of the old House
of Kilravock, he had married a daughter of Sir Patrick Threipland of
Fingask, a family which maintained fidelity to the House of Stuart
with a persistency beyond all parallel, never once swerving in
affection from the days of the Commonwealth down to recent times.
‘Mr John Rose, son of the Bishop of Edinburgh,’ is in the list of rebels
who pled guilty at Carlisle in December 1716. The good bishop,
having come to his sister’s house in the Canongate, to see a brother
who was there lying sick, had a sudden fainting-fit, and calmly
breathed his last. He was buried in the 1720.
romantic churchyard of Restalrig, which
has ever since been a favourite resting-place of the members of the
Scottish Episcopal communion.
There was a jubilation in Edinburgh on Apr. 28.

what appears to us an extraordinary
occasion. The standing dryness between the king and Prince of Wales
had come to a temporary end. The latter had gone formally to the
palace, and been received by his father ‘with great marks of
tenderness’ [the king was sixty, and the prince thirty-seven]. At a
court held on the occasion, ‘the officers and servants on both sides,
from the highest to the lowest, caressed one another with mutual
civilities,’ and there were great acclamations from the crowd outside.
The agreeable news having been received in the northern metropolis,
the magistrates set the bells a-ringing, and held an entertainment for
all persons of note then in town, at which loyal toasts were drunk,
with feux de joie from the City Guard. Demonstrations of a like
nature took place at Glasgow—the music-bells rung—the stairs of the
town-house covered with carpets—toast-drinking—and discharges of
firearms from the Earl of Stair’s regiment. Nor was there a similar
expression of joy wanting even in the Cavalier city of Aberdeen—
where, however, such expressions were certainly more desirable.[543]
One is startled at finding in the May 2.

Edinburgh Evening Courant of this date
the following advertisement: ‘Taken up a stolen negro: whoever owns
him, and gives sufficient marks of his being theirs, before the end of
two weeks from the date hereof, may have him again upon payment
of expenses laid out upon him; otherwise the present possessor must
dispose of him at his pleasure.’
Yet true it is that colonial negro slaves who had accompanied their
masters to the British shores, were, till fifty-five years after this
period, regarded as chattels. One named Joseph Knight came with
his master, John Wedderburn, Esq., to Glasgow in 1771, and
remained with him as his bound slave for two years. A love-affair
then set the man upon the idea of attempting to recover his liberty,
which a recent decision by Lord Mansfield in England seemed to
make by no means hopeless. With the help of friends, he carried his
claim through a succession of courts, till a 1720.
decision of the Court of Session in 1775
finally established that, however he might be a slave while in the
West Indies, he, being now in Scotland, was a free man.
Horse-racing had for many years been considerably in vogue in

Scotland. There were advertised in the course of this year—a race at
Cupar in Fife; one at Gala-rig, near Selkirk, for a piece of plate given
by the burgh, of £12 value; a race at Hamilton Moor for £10; a race
on Lanark Moor for a plate of £12, given by the burgh; a race on the
sands of Leith for a gold cup of about a hundred guineas value, and
another, for a plate of £50 value, given by the city of Edinburgh;
finally, another race at Leith for a silver punch-bowl and ladle, of £25
value, given by the captains of the Trained Bands of Edinburgh—the
bowl bearing an inscription which smacks wonderfully like the
produce of the brain of Allan Ramsay:
‘Charge me with Nantz and limpid spring,
Let sour and sweet be mixt;
Bend round a health syne to the king,
To Edinburgh captains next,
Wha formed me in sae blithe a shape,
And gave me lasting honours;
Take up my ladle, fill and lap,
And say: “Fair fa’ the donors.”’
The genius of Scott has lent an Oct. 17.

extraordinary interest to a murder
perpetrated at this date. Nicol Mushet appears to have been a young
man of some fortune, being described as ‘of Boghall,’ and he had
studied for the profession of a surgeon; but for some time he had led
an irregular and dissipated life in Edinburgh, where he had for one of
his chief friends a noted profligate named Campbell of Burnbank,
ordnance store-keeper in the Castle. The unhappy young man was
drawn into a marriage with a woman named Hall, for whom he soon
discovered that he had neither affection nor respect; and he then
became so eager to be free from the connection, as to listen to a
project by Burnbank for obtaining a divorce by dishonourable
means. An obligation passed between the parties in November 1719,
whereby a claim of Burnbank for an old debt of nine hundred merks
(about £50) was to be discharged by Mushet, as soon as Burnbank
should be able to furnish evidence calculated to criminate the
woman. Burnbank then deliberately hired a wretch like himself, one
Macgregor, a teacher of languages, to enter into a plot for placing
Mrs Mushet in criminative circumstances; 1720.
and some progress was made in this plan,
which, however, ultimately misgave. It was then suggested by
Burnbank that they should go a step further, and remove the woman
by poison. One James Mushet and his wife—a couple in poor
circumstances—readily undertook to administer it. Several doses
were actually given, but the stomach of the victim always rejected
them. Then the project for debauching her was revived, and Mushet
undertook to effect it; but it was not carried out. Dosing with poison
was resumed, without effect; other plans of murder were considered.
James Mushet undertook to knock his sister-in-law on the head for
twenty guineas, and got one or two in hand by anticipation, part of
which he employed in burying a child of his own. These diabolically
wicked projects occupied Mushet, his brother, his brother’s wife, and
Burnbank, in the Christian city of Edinburgh, during a course of
many months, without any one, to appearance, ever feeling the
slightest compunction towards the poor woman, though it is
admitted she loved her husband, and no real fault on her side has
ever been insinuated.
At length, the infatuated Nicol himself borrowed a knife one day,
hardly knowing what he wanted it for, and, taking his wife with him
that night, as on a walk to Duddingston, he embraced the
opportunity of killing her at a solitary place in the King’s Park. He
went immediately after to his brother’s, to tell him what he had done,
but in a state of mind which made all afterwards seem a blank to
him. Next morning, the poor victim was found lying on the ground,
with her throat cut to the bone, and many other wounds, which she
had probably received in struggling with her brutal murderer.
Mushet was seized and examined, when he readily related the
whole circumstances of the murder and those which had led to it. He
was adjudged to be hanged in the Grassmarket on the ensuing 6th of
January. His associate Burnbank was declared infamous, and
sentenced to banishment. The common people, thrilled with horror
by the details of the murder, marked their feelings in the old national
mode by raising a cairn on the spot where it took place; and Mushet’s
Cairn has ever since been a recognised locality.[544]
There was published this year in Edinburgh a small treatise at the

price of a shilling, under the title of Rules of 1720.
Good Deportment and Good Breeding. The
author was Adam Petrie, who is understood as having commenced
life as domestic tutor in the family of Sinclair of Stevenston, and to
have ended it in the situation of a parish schoolmaster in East
Lothian. He dedicated his treatise to the magistrates of Edinburgh,
acknowledging them to be ‘so thoroughly acquainted with all the
steps of civility and good breeding, that it is impossible for the least
misrepresentation of them to escape your notice.’
Adam sets out with the thesis, that ‘a courteous way gilds a denial,
sweetens the sharpness of truth ... sets off the defects of reason,
varnishes slights, paints deformities ... in a word, disguises
everything that is unsavoury.’ Everything, however, required to have
some reference to religion in that age, and Adam takes care to
remind us that civility has a divine basis, in the injunctions, ‘Be
courteous to all men,’ and ‘Give honour to whom honour is due.’
As to ordinary demeanour, Mr Petrie was of opinion that ‘a
gentleman ought not to run or walk too fast in the streets, lest he be
suspected to be going a message.’ ‘When you walk with a superior,
give him the right hand; but if it be near a wall, let him be next to it.’
The latter rule, he tells us, was not yet followed in Scotland, though
established in England and Ireland. ‘When you give or receive
anything from a superior, be sure to pull off your glove, and make a
show of kissing your hand, with a low bow after you have done.’ In
this and some other instances, it strikes us that a too ceremonious
manner is counselled; but such was the tendency of the time. There
was, however, no want of rude persons. ‘I have,’ says Adam, ‘seen
some noblemen treat gentlemen that have not been their
dependents, and men of ancienter families than they could pretend
to, like their dependents, and carry to the ambassadors of Jesus
Christ as if they had been their footmen.’
Mr Petrie deemed it proper not to come amongst women abruptly,
‘without giving them time to appear to advantage: they do not love to
be surprised.’ He also thought it was well ‘to carry somewhat
reserved from the fair sex.’ One should not enter the house or
chamber of a great person with a great-coat and boots, or without
gloves—though ‘it is usual in many courts that they deliver up their
gloves with their sword before they enter the court, because some
have carried in poison on their gloves, and have conveyed the same
to the sovereign that way.’ Women, on their 1720.
part, are equally advised against
approaching superiors of their own sex with their gown tucked up.
‘Nor,’ says he, ‘is it civil to wear a mask anywhere in company of
superiors, unless they be travelling together on a journey.’ In that
case, ‘when a superior makes his honours to her, she is to pull off her
mask, and return him his salute, if it be not tied on.’
There is a good deal about the management of the handkerchief,
with one general recommendation to ‘beware of offering it to any,
except they desire it.’ We also are presented with a rule which one
could wish to see more universally observed than it is, against
making any kind of gesticulations or noises in company.
There were customs of salutation then, which it is now difficult to
imagine as having ever been practised. ‘In France,’ says Adam, ‘they
salute ladies on the cheek; but in Britain and Ireland they salute
them on the lips.’ Our Scottish Chesterfield seems to have felt that
the custom should be abated somewhat; or perhaps it was going out.
‘If,’ says he, ‘a lady of quality advance to you, and tender her cheek,
you are only to pretend to salute her by putting your head to her
hoods: when she advances, give her a low bow, and when you retreat,
give her another.’ He adds: ‘It is undecent to salute ladies but in
civility.’
Formulæ of address and for the superscription of letters are fully
explained; but Adam could not allow ‘the Right Reverend Father in
God the Lord Bishop of London’ to pass as an example of Episcopal
style, without remarking that many have not ‘clearness’ to use such
titles. Adam is everywhere inclined to an infusion of piety. He
denounces ‘an irreligious tippling’ of coffee, tea, and chocolate,
which he observed to be continually going on in coffee-houses,
‘because not one in a hundred asks a blessing to it.’ He is very much
disposed, too, to launch out into commonplace morals. Rather
unexpectedly in a lover of the politenesses, he sets his face wholly
against cards and dice, stage-plays, and promiscuous dancing,
adducing a great number of learned references in support of his
views.
The editor of a very scarce reprint of this curious volume,[545]
remarks that, from the manifest sincerity of the author’s delineations
of good breeding, and the graphic character of many of his scenes, it
may fairly be presumed that they were painted from nature. We are
told by the same writer, that ‘Helen Countess of Haddington, who
died in 1768, at the advanced age of ninety-one, and to whom Petrie
was well known, used to describe his own 1720.
deportment and breeding as in strict
accordance with his rules.’
James Dougal, writing the news of 1721. Jan. 4.

Edinburgh to his friend Wodrow at
Eastwood, has a sad catalogue to detail. ‘There was four pirates
hanged at Leith this day ... very hardened. They were a melancholy
sight, and there is three to be hanged next Wednesday. Nicol Mushet
is to be hanged on Friday ... for murdering his wife: he appears to be
more concerned than he was before. Ane woman brought from Leith
is to die the first Wednesday of February for putting down
[destroying] a child. Another man is laid up in prison, that is thought
to have murdered his wife. The things falling out now are very
humbling.’
He goes on to tell that several persons ‘in trouble of mind’ are
frequently prayed for in Edinburgh churches. ‘But they do not name
them but after such a manner—A man there is in such trouble (or a
woman), and desires the congregation to praise God with them for
signal deliverance that the Lord hath given them from great troubles
that they have been in.’
The end of the letter is terrible: ‘There is some of the Lord’s people
that lives here, that are feared for melancholy days, iniquity doth so
abound, and profanity; and if there were not a goodly remnant in this
town, it would sink.’[546]
A sperm whale, ‘the richest that has ever Jan. 30.

been seen in this country,’ was advertised in
the Courant as having come ashore in the Firth of Forth near
Culross, and to be sold by public roup.
At the end of June 1730, three wounded whales ran ashore at
Kilrenny in Fife, on the property of Mr Bethune of Balfour. The
produce, consisting of a hundred and forty-six barrels of speck, or
blubber, and twenty-three barrels of spermaceti speck, was
afterwards advertised for sale.
With regard to several of the forfeited Oct.

estates which lay in inaccessible situations
in the Highlands, the Commissioners had been up to this time
entirely baffled, having never been able even to get surveys of them
effected. In this predicament in a special manner lay the immense
territory of the Earl of Seaforth, extending from Brahan Castle in
Easter Ross across the island to Kintail, and 1721.
including the large though unfertile island
of Lewis. The districts of Lochalsh and Kintail, on the west coast, the
scene of the Spanish invasion of 1719, were peculiarly difficult of
access, there being no approach from the south, east, or north,
except by narrow and difficult paths, while the western access was
only assailable to a naval force. To appearance, this tract of ground,
the seat of many comparatively opulent ‘tacksmen’ and cattle-
farmers, was as much beyond the control of the six Commissioners
assembled at their office in Edinburgh, as if it had been amongst the
mountains of Tibet or upon the shores of Madagascar.
During several years after the insurrection, the rents of this district
were collected, without the slightest difficulty, for the benefit of the
exiled earl, and regularly transmitted to him. At one time, a
considerable sum was sent to him in Spain, and the descendants of
the man who carried it continued for generations to bear ‘the
Spanyard’ as an addition to their name.[547] The chief agent in the
business was Donald Murchison, descendant of a line of faithful
adherents of the ‘high chief of Kintail’—the first of whom, named
Murcho, had come from Ireland with Colin the son of Kenneth, the
founder of the clan Mackenzie in the thirteenth century. The later
generations of the family had been intrusted in succession with the
keeping of Ellan Donan Castle, a stronghold dear to the modern
artist as a picturesque ruin, but formerly of serious importance as
commanding a central point from which radiate Loch Alsh and Loch
Duich, in the midst of the best part of the Mackenzie country. Donald
was a man worthy of a more prominent place in his country’s annals
than he has yet attained; he acted under a sense of right which,
though unfortunately defiant of acts of parliament, was still a very
pure sense of right; and in the remarkable actions which he
performed, he looked solely to the good of those towards whom he
had a feeling of duty. A more disinterested hero—and he was one—
never lived.
When Lord Seaforth brought his clan to fight for King James in
1715, Donald Murchison and a senior brother, John, went as field-
officers of the regiment—Donald as lieutenant-colonel, and John as
major. Sir Roderick I. Murchison, the distinguished geologist, great-
grandson of John, possesses a large ivory and silver ‘mill,’ which
once contained the commission sent from France to Donald, as
colonel, bearing the inscription: ‘James 1721.
Rex: Forward and Spare not.’ John fell at
Sheriffmuir, in the prime of life; Donald, returning with the remains
of the clan, was intrusted by the banished earl with the management
of estates no longer legally his, but still virtually so, through the
effect of Highland feelings in connection with very peculiar local
circumstances. And for this task Donald was in various respects well
qualified, for, strange to say, the son of the castellan of Ellan Donan
—the Sheriffmuir colonel—had been ‘bred a writer’ in Edinburgh,
and was as expert at the business of a factor or estate-agent as in
wielding the claymore.[548]
In bold and avowed insubordination to the government of George
the First, the Mackenzie tenants continued for ten years to pay their
rents to Donald Murchison, on account of their forfeited and exiled
lord, setting at nought all fear of ever being compelled to repeat the
payment to the commissioners.
In 1720, these gentlemen made a movement for asserting their
claims upon the property. In William Ross of Easterfearn, and
Robert Ross, a bailie of Tain, they found two men bold enough to
undertake the duty of stewardship in their behalf over the Seaforth
property, and also the estates of Grant of Glenmorriston and
Chisholm of Strathglass. Little, however, was done that year beyond
sending out notices to the tenants, and preparing for strenuous
measures to be entered upon next year. The stir they made only
produced excitement, not dismay. Some of the duine-wassels from
about Loch Carron, coming down with their cattle to the south-
country fairs, were heard to declare that the two factors would never
get anything but leaden coin from the Seaforth tenantry. Donald was
going over the whole country, shewing a letter he had got from the
earl, encouraging his people to stand out; at the same time telling
them that the old countess was about to come north with a factory
for the estate, when she would allow as paid any rents which they
might now hand to him. The very first use to be made of this money
was, indeed, to bring both the old and the young countesses home
immediately to Brahan Castle, where they would live as they used to
do. Part of the funds thus acquired, he used in keeping on foot a
party of about sixty armed Highlanders, whom, in virtue of his
commission as colonel, he proposed to employ in resisting any
troops of George the First which might be sent to Kintail. Nor did he
wait to be attacked, but, in June 1720, 1721.
hearing of a party of excisemen passing
near Dingwall with a large quantity of aqua-vitæ, he fell upon them,
and rescued their prize. The collector of the district reported this
transaction to the Board of Excise; but no notice was taken of it.
In February 1721, the two factors sent officers of their own into the
western districts, to assure the tenants of good usage, if they would
make a peaceable submission; but the men were seized, robbed of
their papers, money, and arms, and quietly remanded over the Firth
of Attadale, though only after giving solemn assurance that they
would never attempt to renew their mission. Resenting this
procedure, the two factors caused a constable to take a military party
from Bernera barracks into Lochalsh, and, if possible, capture those
who had been guilty. They made a stealthy night-march, and took
two men; but the alarm was given, the two men escaped, and began
to fire down upon their captors from a hillside; then they set fire to
the bothy as a signal, and such a coronach went over all Kintail and
Lochalsh, as made the soldiers glad to beat a quick retreat.
After some further proceedings, all of them ineffectual, the two
factors were enabled, on the 13th of September, to set forth from
Inverness with a party of thirty soldiers and some armed servants of
their own, with the design of enforcing submission to their legal
claims. Let it be remembered there were then no roads in the
Highlands, nothing but a few horse-tracks along the principal lines in
the country, where not the slightest effort had ever been made to
smooth away the natural difficulties of the ground. In two days, the
factors had got to Invermorriston; but here they were stopped for
three days, waiting for their heavy baggage, which was storm-stayed
in Castle Urquhart, and there nearly taken in a night-attack by a
partisan warrior bearing the name of Evan Roy Macgillivray. The
tenantry of Glenmorriston at first fled with their bestial; but
afterwards a number of them came in and made at least the
appearance of submission. The party then moved on towards
Strathglass, while Evan Roy respectfully followed, to pick up any
man or piece of baggage that might be left behind. At Erchless Castle,
and at Invercannich, seats of the Chisholm, they held courts, and
received the submission of a number of the tenants, whom, however,
they subsequently found to be ‘very deceitful.’
There were now forty or fifty miles of the wildest Highland country
before them, where they had reason to believe they should meet
groups of murderous Camerons and 1721.
Glengarry Macdonalds, and also encounter
the redoubted Donald Murchison, with his guard of Mackenzies,
unless their military force should be of an amount to render all such
opposition hopeless. An appointment having been made that they
should receive an addition of fifty soldiers from Bernera, with whom
to pass through the most difficult part of their journey, it seemed
likely that they would appear too strong for resistance; and, indeed,
intelligence was already coming to them, that ‘the people of Kintail,
being a judicious opulent people, would not expose themselves to the
punishments of law,’ and that the Camerons were absolutely
determined to give no further provocation to the government. Thus
assured, they set out in cheerful mood along the valley of Strathglass,
and, soon after passing a place called Knockfin, were reinforced by
Lieutenant Brymer, with the expected fifty men from Bernera. There
must have now been about a hundred well armed men in the invasive
body. They spent the next day (Sunday) together in rest, to gather
strength for the ensuing day’s march of about thirty arduous miles,
by which they hoped to reach Kintail.
At four in the morning of Monday the 2d October, the party set
forward, the Bernera men first, and the factors in the rear. They were
as yet far from the height of the country, and from its more difficult
passes; but they soon found that all the flattering tales of non-
resistance were groundless, and that the Kintail men had come a
good way out from their country in order to defend it. The truth was,
that Donald Murchison had assembled not only his stated band of
Mackenzies, but a levy of the Lewis men under Seaforth’s cousin,
Mackenzie of Kildun; also an auxiliary corps of Camerons, Glengarry
and Glenmorriston men, and some of those very Strathglass men
who had been making appearances of submission. Altogether, he
had, if the factors were rightly informed, three hundred and fifty men
with long Spanish firelocks, under his command, and all posted in
the way most likely to give them an advantage over the invading
force.
The rear-guard, with the factors, had scarcely gone a mile, when
they received a platoon of seven shots from a rising ground near
them to the right, with, however, only the effect of piercing a soldier’s
hat. The Bernera company, as we are informed, left the party at eight
o’clock, as they were passing Lochanachlee, and from this time is
heard of no more: how it made its way out of the country does not
appear. The remainder still advancing, Easterfearn, as he rode a little
before his men, had eight shots levelled at 1721.
him from a rude breastwork near by, and
was wounded in two places, but was able to appear as if he had not
been touched. Then calling out some Highlanders in his service, he
desired them to go before the soldiers, and do their best, according to
their own mode of warfare, to clear the ground of such lurking
parties, so that the troops might advance in safety. They performed
this service pretty effectually, skirmishing as they went on, and the
main body advanced safely about six miles. They were here arrived at
a place called Aa-na-Mullich (Ford of the Mull People), where the
waters, descending from the Cralich and the lofty mountains of
Kintail, issue eastwards through a narrow gorge into Loch Affaric. It
was a place remarkably well adapted for the purposes of a resisting
party. A rocky boss, called Tor-an-Beatich, then densely covered with
birch, closes up the glen as with a gate. The black mountain stream,
‘spear-deep,’ sweeps round it. A narrow path wound up the rock,
admitting only of passengers in single file. Here lay Donald with the
best of his people, while inferior adherents were ready to make
demonstrations at a little distance. As the invasive party approached,
they received a platoon from a wood on the left, but nevertheless
went on. When, however, they were all engaged in toiling up the
pass, forty men concealed in the heather close by fired with deadly
effect, inflicting a mortal wound on Walter Ross, Easterfearn’s son,
while Bailie Ross’s son was also hurt by a bullet which swept across
his breast. The bailie called to his son to retire, and the order was
obeyed; but the two wounded youths and Bailie Ross’s servant were
taken prisoners, and carried up the hill, where they were quickly
divested of clothes, arms, money, and papers. Young Easterfearn
died next morning. The troops faced the ambuscade manfully, and
are said to have given their fire thrice, and to have beat the
Highlanders from the bushes near by; but, observing at this juncture
several parties of the enemy on the neighbouring heights, and being
informed of a party of sixty in their rear, Easterfearn deemed it best
to temporise.
He sent forward a messenger to ask who they were that opposed
the king’s troops, and what they wanted. The answer was that, in the
first place, they required to have Ross of Easterfearn delivered up to
them. This was pointedly refused; but it was at length arranged that
Easterfearn should go forward, and converse with the leader of the
opposing party. The meeting took place at Bal-aa-na-Mullich (the
Town of the Mull Men’s Ford), and Easterfearn found himself
confronted with Donald Murchison. It 1721.
ended with Easterfearn giving up his
papers, and covenanting, under a penalty of five hundred pounds,
not to officiate in his factory any more; after which he gladly
departed homewards with his associates, under favour of a guard of
Donald’s men, to conduct them safely past the sixty men lurking in
the rear. It was alleged afterwards that the commander was much
blamed by his own people for letting the factors off with their lives
and baggage, particularly by the Camerons, who had been five days
at their post with hardly anything to eat; and Murchison only
pacified them by sending them a good supply of meat and drink. He
had in reality given a very effectual check to the two gentlemen-
factors, to one of whom he imparted in conversation that any scheme
of a government stewardship in Kintail was hopeless, for he and
sixteen others had sworn that, if any person calling himself a factor
came there, they would take his life, whether at kirk or at market,
and deem it a meritorious action, though they should be cut to pieces
for it next minute.[549]
A bloody grave for young Easterfearn in Beauly Cathedral
concluded this abortive attempt to take the Seaforth estates within
the scope of a law sanctioned by statesmen, but against which the
natural feelings of nearly a whole people revolted.[550]
A newspaper advertisement informed the 1721. Dec.
world that ‘There is a certain gentleman
living at Glasgow, who has put forth a problem to the learned—
proposing, if no man answer it, to do it himself in a few weeks—viz.,
Whether or not it is possible so to dispose a ship, either great or
small, that, although she, or it, be rent in the bottom, and filled full of
water, or however tossed with tempest, she, or it, shall never sink
below the water; and also that the same may be reduced to
practice.’[551]
An election of a member of parliament for 1722. Apr. 27.

a Highland county was apt to bring forth
somewhat strenuous sentiments, and the scene sometimes partook a
good deal of the nature of a local civil war.
A representative of Ross-shire being to be chosen, there came, the
night before, to Fortrose, the greatest man of the north, the Earl of
Sutherland, heading a large body of armed and mounted retainers,
who made a procession round the streets, while an English sloop-of-
war, in friendly alliance with him, came up to the town and fired its
guns. Hundreds of Highlanders, his lordship’s retainers, at the same
time lounged about. The reason of all this was, that the opposition
interest was in a decided majority, and a defeat to the Whig
candidate seemed impending. When the election came on, there
were thirty-one barons present, of whom eighteen gave their votes
for General Charles Ross of Balnagowan, the remainder being for
Captain Alexander Urquhart of Newhall. Hereupon, Lord
Sutherland’s relative and friend, Sir William Gordon of Invergordon,
sheriff of the county, retired with the minority, and went through the
form of electing their own man, notwithstanding a protest from the
other candidate. ‘Immediately after this separation, Colin Graham of
Drynie, one of the deputy-lieutenants of the county, came into the
court-house, with his sword in his hand, accompanied by Robert
Gordon of Haughs and Major John Mackintosh, with some of the
armed Highlanders whom they had posted at the door, with drawn
swords and cocked firelocks, and did require the majority (who
remained to finish the election), in the name of the Earl of
Sutherland, to remove out of the house, otherwise they must expect
worse treatment. Major Mackintosh said they would be dragged out
by the heels. Upon which the barons protested against those violent
proceedings, declaring their resolution to 1722.
remain in the court-house till the election
was finished, though at the hazard of their lives; which they
accordingly did.’[552]
The Catholics had of late been getting up Apr. 29.

their heads in the north, especially in
districts over which the Gordon family held sway; and the open
practice of the Romish rites before large congregations in the
Banffshire valleys, was become a standing subject of complaint and
alarm in the church-courts. When at length the government obtained
scent of the Jacobite plot in which Bishop Atterbury was concerned,
it sympathised with these groans of the laden spirits in Scotland, and
permitted some decided measures of repression to be taken.
Accordingly, this day, being Sunday, as the Duchess-Dowager of
Gordon—Elizabeth Howard, daughter of the Duke of Norfolk—was
having mass performed at her house in the Canongate, Edinburgh, in
the presence of about fifty professors like herself of the Catholic
religion, Bailie Hawthorn, a magistrate of the Canongate, broke open
the doors, and seized the whole party. The ladies were bailed, and
allowed to depart; but the priest, Mr John Wallace, was marched to
prison. We are informed by Wodrow that Wallace had been ordained
a Protestant minister thirty-five years before.[553] The Lord Advocate
would not at first listen to any proposal for his liberation, though
several persons of distinction came to plead for it; but at length bail
was taken for him to the extent of a thousand merks Scots. Being
indicted under the statute of 1700, he failed to stand his trial, and
was outlawed.[554]
Before the upbreak of this plot, considerable numbers of
gentlemen under attainder daily presented themselves on the streets
of Edinburgh, emboldened of course by the mildness of the
government; but, one or two of them having been seized and put up
in the Castle, it came to pass, 15th May, that not one was any longer
to be seen. Mr Wodrow, who records these circumstances, expresses
the feeling of the hour. ‘It’s certain we are in a most divided and
defenceless state; divisions on the one hand, rancour and malice on
the other, and a wretched indolence among too many. But the Lord
liveth!’[555]
The Canongate, which had so often, in the Aug. 3.

sixteenth century, 1722.
been reddened
with the best blood in Scotland, was still occasionally the scene of
wild transactions, though arising amongst a different class of persons
and from different causes. A local journalist chronicles a dreadful
tragedy as occurring on its pavé at this date.
‘In the afternoon, Captain Chiesley and Lieutenant Moodie, both of
Cholmly’s Regiment, which lies encamped at Bruntsfield Links,
having quarrelled some time before in the camp, meeting on the
street of the Canongate, the captain, as we are told, asked Mr Moodie
whether he had in a certain company called him a coward? And he
owning he had, the captain beat him first with his fist, and then with
a cane; whereupon Mr Moodie drew his sword, and, shortening it,
run the captain into the great artery. The captain, having his sword
drawn at the same time, pushed at Mr Moodie, who was rushing on
him with his sword shortened, and thus run him into the lower belly,
of which in a few minutes he died, without speaking one word,
having had no more strength or life left him than to cross the street,
and reach the foot of the stair of his lodgings, where he dropped
down dead. The captain lived only to step into a house near by, and
to pray shortly that God might have mercy on his soul, without
speaking a word more. ’Tis said Mr Moodie’s lady was looking over
the window all the while this bloody tragedy was acting.’[556]
A duel which happened about the same time between Captains
Marriot and Scroggs proved fatal to both.
‘Four of those poor deluded people called Aug. 7.

Quakers, two men and two women, came
about noon to the Cross [of Edinburgh], when one of the women,
who by her accent seemed to be of Yorkshire, after several violent
agitations, said, that she was appointed by God to preach repentance
to this sinful city; that a voice of mortality, as she called it, had
sounded in her ears, and that desolation and all kinds of miseries
would befall the inhabitants if they did not repent. After she had
spoke about a quarter of an hour, a party of the city-guard carried
her and the other three prisoners to the main guard.’
Some years after, one Thomas Erskine, a brewer, made himself
conspicuous as a Quaker preacher in Edinburgh. One Saturday,
January 17, 1736, he ‘made a religious peregrination through this
city. He made his first station at the Bow-head [reputed as the head-
quarters of the saints in Edinburgh], where 1722.
he pronounced woes and judgments on the
inhabitants of the Good Town, if they did not speedily repent. Thence
he walked to the Cross, where he recapitulated what he had
evangelised by the way, and concluded with desiring his auditory to
remember well what he had told them. However, he gave them forty
days to think on’t.’
One day, in the ensuing July, Erskine sent a notice to the quiet
little country town of Musselburgh, to the effect that the Spirit had
appointed him to hold forth to them in the marketplace at five in the
afternoon; and, accordingly, at the appointed hour, he mounted the
Cross, and discoursed to a large audience.[557]
A second attempt was now made to Aug. 29.

obtain possession of the forfeited Seaforth
estates for the government. It was calculated that what the two
factors and their attendants, with a small military force, had failed to
accomplish in the preceding October, when they were beat back with
a fatal loss at Aa-na-Mullich, might now be effected by means of a
good military party alone, if they should make their approach
through a less critical passage. A hundred and sixty of Colonel Kirk’s
regiment left Inverness under Captain M‘Neil, who had at one time
been commander of the Highland Watch. They proceeded by
Dingwall, Strath Garve, and Loch Carron, a route to the north of that
adopted by the factors, and an easier, though a longer way. Donald
Murchison, nothing daunted, got together his followers, and
advanced to the top of Maam Attadale, a high pass from Loch Carron
to the head of Loch Long, separating Lochalsh from Kintail. Here a
gallant relative named Kenneth Murchison, and a few others,
volunteered to go forward and plant themselves in ambush in the
defiles of the Choille Van [White Wood], while the bulk of the party
should remain where they were. It would appear that this ambush
party consisted of thirteen men, all peculiarly well armed.
On approaching this dangerous place, the captain went forward
with a sergeant and eighteen men to clear the wood, while the main
body came on slowly in the rear. At a place called Altanbadn, in the
Choille Van, he encountered Kenneth and his associates, whose fire
wounded himself severely, killed one of his grenadiers, and wounded
several others of the party. He persisted in advancing, and attacked
the handful of natives with sufficient 1722.
resolution. They slowly withdrew, as unable
to resist; but the captain now obtained intelligence that a large body
of Mackenzies was posted in the mountain-pass of Attadale. It
seemed as if there was a design to draw him into a fatal ambuscade.
His own wounded condition probably warned him that a better
opportunity might occur afterwards. He turned his forces about, and
made the best of his way back to Inverness. Kenneth Murchison
quickly rejoined Colonel Donald on Maam Attadale, with the
cheering intelligence that one salvo of thirteen guns had repelled the
hundred and sixty sidier roy.[558] After this, we hear of no renewed
attempt to comprise the Seaforth property.
Strange as it may seem, Donald Murchison, two years after thus a
second time resisting the government troops, came down to
Edinburgh with eight hundred pounds of the earl’s rents, that he
might get the money sent abroad for his lordship’s use. He remained
a fortnight in the city unmolested. He would on this occasion appear
in the garb of a Lowland gentleman; he would mingle with old
acquaintances, ‘doers’ and writers; and appear at the Cross amongst
the crowd of gentlemen who assembled there every day at noon.
Scores would know all about his doings at Aa-na-Mullich and the
Choille Van; but thousands might have known, without the chance of
one of them betraying him to government.
General Wade, writing a report to the king in 1725, states that the
Seaforth tenants, formerly reputed the richest of any in the
Highlands, are now become poor, by neglecting their business, and
applying themselves to the use of arms. ‘The rents,’ he says, ‘continue
to be collected by one Donald Murchison, a servant of the late earl’s,
who annually remits or carries the same to his master into France.
The tenants, when in a condition, are said to have sent him free gifts
in proportion to their several circumstances, but are now a year and
a half in arrear of rent. The receipts he gives to the tenants are as
deputy-factor to the Commissioners of the Forfeited Estates, which
pretended power he extorted from the factor (appointed by the said
commissioners to collect those rents for the use of the public), whom
he attacked with above four hundred armed men, as he was going to
enter upon the said estate, having with him a party of thirty of your
majesty’s troops. The last year this Murchison marched in a public
manner to Edinburgh, to remit eight hundred pounds to France for
his master’s use, and remained fourteen days there unmolested. I
cannot omit observing to your majesty, that 1722.
this national tenderness the subjects of
North Britain have one for the other, is a great encouragement for

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Advanced Nutrition and Human

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SECTION I Cells and Their Nourishment

SECTION II Macronutrients and Their Metabolism

SECTION III The Regulatory Nutrients

Preface xvii Regulatory Peptides 57

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

Enzyme Translocation 104 Phospholipid Digestion 148

6.4 Absorption 193 6.11 Catabolism of Tissue/Cell Proteins and

Other Roles 222 Glucagon 280

Indirect Calorimetry 294 Deficiency: Beriberi 344

Toxicity 382 10.3 Vitamin E 435

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

11.3 Magnesium 485 Respiratory Regulation 519

13.5 Chromium 575 Assessment of Nutriture 590

S Chapter 2 The Digestive System:

Chapter 11 Major Minerals

Although this textbook represents countless hours of

1.1 COMPONENTS OF CELLS

The exportation of some corn from Feb. 5.

Died, Alexander Rose, who had been Mar. 20.

There was a jubilation in Edinburgh on Apr. 28.

One is startled at finding in the May 2.

Horse-racing had for many years been considerably in vogue in

The genius of Scott has lent an Oct. 17.

There was published this year in Edinburgh a small treatise at the

James Dougal, writing the news of 1721. Jan. 4.

A sperm whale, ‘the richest that has ever Jan. 30.

With regard to several of the forfeited Oct.

An election of a member of parliament for 1722. Apr. 27.

The Catholics had of late been getting up Apr. 29.

The Canongate, which had so often, in the Aug. 3.

‘Four of those poor deluded people called Aug. 7.

A second attempt was now made to Aug. 29.

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