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Biochemical, Physiological, and
Molecular Aspects of Human
Nutrition

FOURTH EDITION

Martha H. Stipanuk, PhD

Professor, Division of Nutritional Sciences, Colleges of Human Ecology
and, Agriculture and Life Sciences, Cornell University, Ithaca, New York

Marie A. Caudill, PhD, RD

Professor, Division of Nutritional Sciences, Colleges of Human Ecology
and, Agriculture and Life Sciences, Cornell University, Ithaca, New York
Table of Contents

Cover image

Title page

Copyright

Contributors

Reviewers

Preface

Acknowledgments
UNIT I. Nutrients

Chapter 1. Nutrients: History and Definitions

Discovery of the Nutrients

Setting Criteria for Essentiality

Conditionally Essential Nutrients

Shift From Focus on Prevention of Deficiency Diseases to Providing Nutrients for

Maintenance of Good Health
 Recognition of Health Benefits of Nonnutrient Components of Food or of Higher Intakes of
Nutrient Components of Food

Personalized Nutrition

Rise of the Supplement and Functional Food Industries

Thinking Critically—End of Chapter

Chapter 2. Guidelines for Food and Nutrient Intake

Dietary Reference Intakes

Dietary Advice: Goals, Guidelines, and Food Guides

Food and Supplement Labels and Other Nutrition Information

Thinking Critically—End of Chapter

UNIT II. Structure and Properties of the Macronutrients

Chapter 3. Structure, Nomenclature, and Properties of Carbohydrates

Monosaccharides

Glycosidic Linkages, Glycosides, and Polymerization of Sugar Residues

Common Disaccharides and Oligosaccharides

Polysaccharides

Conjugates of Carbohydrates and Proteins or Lipids

Thinking Critically—End of Chapter

Chapter 4. Structure, Nomenclature, and Properties of Lipids

Fatty Acids

Glycerolipids

Sphingolipids

Other Lipids with Linked Fatty Acids—Esters, Thioesters, and Amides

Isoprenoids and Steroids

 Oxidized Derivatives of Fatty Acids

Amphipathic Lipids at Water–Lipid Interfaces

Lipid Autoxidation of Pufas

Thinking Critically — End of Chapter

Chapter 5. Structure, Nomenclature, and Properties of Proteins and Amino

Acids
The Proteinogenic Amino Acids

Synthesis of Peptides and Proteins

Protein Structure

Posttranslational Modifications of Proteins

Trafficking of Proteins to their Destinations

Regulation of the Amount of Protein and its Functional State

Classification of Proteins

Synthesis of Small Peptides or Isopeptide Linkages Without Involvement of the Genetic

Code or Ribosomes

Important Food Proteins: Nutritional and Functional Aspects

Thinking Critically—End of Chapter

UNIT III. Digestion and Absorption of the Macronutrients

Chapter 6. Overview of Digestion and Absorption

General Structure and Function of the Gastrointestinal Tract

The Upper Gastrointestinal System

The Small Intestine

The Large Intestine

General Mechanisms for Regulation of Gi Tract Function

Thinking Critically—End of Chapter

Chapter 7. Digestion and Absorption of Carbohydrate
Overview of Digestion and Absorption of Carbohydrates

Digestion of Available Carbohydrates to Monosaccharides and Absorption of the

Monosaccharides

Regulation of Postprandial Glycemia

Fate of Nondigestible Oligosaccharides and Nonstarch Polysaccharides in the Large

Intestine

Thinking Critically—Clinical Correlation Box—Glucose And Galactose Malabsorption

Thinking Critically—CLinical Correlation Box—Diabetes Mellitus

Thinking Critically—Nutrition Insight Box—Prebiotics or Probiotics or Both

Thinking Critically—Clinical Correlation Box—Short-Bowel Patients

Thinking Critically—End of Chapter

Chapter 8. Digestion and Absorption of Lipids

Overview of Digestion and Absorption of Lipids

Digestion of Lipids

Absorption of Lipid Digestion Products by Enterocytes

Intracellular Metabolism and Secretion of Absorbed Lipid Digestion Products by

Enterocytes

Portal Transport of Fatty Acids

Dietary Lipid Sensing and Signaling Functions in the Gut And Brain–Gut Axis

Thinking Critically — End of Chapter

Chapter 9. Digestion and Absorption of Protein

Overview of Protein Digestion and Absorption

Digestion of Protein in the Gastrointestinal Tract

Absorption of Free Amino Acids and Small Peptides

 Antigen Sampling and Bioactive Peptides

Fate of Undigested Protein or Unabsorbed Peptides and Amino Acids

Determining Dietary Protein Digestibility

Thinking Critically—Nutrition Insight Box—Trypsin Inhibitors

Thinking Critically—Clinical Correlation Box—Cystinuria

Thinking Critically—Clinical Correlation Box—Celiac Disease: Gluten-Sensitive

Enteropathy

Thinking Critically—End of Chapter

UNIT IV. Metabolism of the Macronutrients

Chapter 10. Central Aspects of Macronutrient Metabolism

Central Role of ATP and NADPH in Coupling Fuel Catabolism to Performance of
Anabolic/Chemical, Electrogenic, and Mechanical Work

Overview of Fuel Catabolism

The Oxidative Decarboxylation of Pyruvate, the Citric Acid Cycle, the Electron Transport
Chain, and Oxidative Phosphorylation

ATP Synthesis: The Mitochondrial Electron Transport Chain and Oxidative

Phosphorylation

Reactions Involved in the Reduction of NADP to NADPH

Counting Energy Costs Using ATP Equivalents

Amphibolic Roles of Central Pathways of Metabolism

Thinking Critically—Nutrition Insight Box—Alcohol: Calories and Concerns

Thinking Critically—End of Chapter

Chapter 11. Overview of the Regulation of Macronutrient Metabolism

Metabolism Includes Both Catabolism and Anabolism

Regulation of Macronutrient Metabolism at the Whole-Body Level

Regulation of Macronutrient Metabolism at the Tissue Level

 Macronutrient Metabolism at the Cellular Level

Regulation of Macronutrient Metabolism at the Cellular Level

Integrative Pathways for Regulation of Macronutrient Metabolism at the Cellular Level

Thinking Critically – End of Chapter

Chapter 12. Metabolism of Carbohydrate

Cellular Uptake and Metabolism of Glucose

Glucose Utilization: Glycolysis

Metabolism of Monosaccharides other than Glucose

Glucose Utilization: Glycogen Synthesis from Glucose

Other Pathways of Glucose Utilization

Glucose Production: Glycogen Degradation

Glucose Production: Gluconeogenesis

Other Pathways of Carbohydrate Metabolism; Synthesis of other Sugars and Sugar

Derivatives, Glucuronides, Lactose, Glycoproteins, and Glycolipids

Thinking Critically—Clinical Correlation Box—Hereditary Fructose Intolerance and

Essential Fructosuria

Thinking Critically—Clinical Correlation Box—Glycogen Synthase Deficiency

Thinking Critically—Clinical Correlation Box—Glucose 6-Phosphatase Deficiency

Thinking Critically—End of Chapter

Chapter 13. Metabolism of Fatty Acids, Acylglycerols, and Sphingolipids

Fatty Acid and Triacylglycerol Metabolism

β-Oxidation of Fatty Acids

Formation of Ketone Bodies from Acetyl-CoA in the Liver as a Fuel for Extrahepatic
Tissues

De Novo Synthesis of Fatty Acids from Acetyl-CoA

Synthesis of C20 and C22 Polyunsaturated Fatty Acids from Dietary Precursors
 Synthesis of Triacylglycerols

Synthesis, Secretion, and Metabolism of Triacylglycerol-Rich Lipoproteins

Mobilization of Stored Triacylglycerols

Phospholipids

Ether-Linked Glycerophospholipids

Sphingolipids

Proteolipids

Thinking Critically—Nutrition Insight BOX—The Carnitine Shuttle in Reverse

Thinking Critically—Clinical Correlation Box—Deficiencies in Beta-Oxidation

Thinking Critically—Clinical Correlation Box—Essential Fatty Acid Deficiency

Thinking Critically—Nutrition Insight Box—Fatty Acid Remodeling

Thinking Critically—Clinical Correlation Box—Inborn Errors

Thinking Critically—End of Chapter

Chapter 14. Cholesterol and Lipoproteins: Synthesis, Transport, and

Metabolism
Overview of Cholesterol and Lipoprotein Metabolism

Cholesterol and Isoprenoid Synthesis and its Regulation

The Role of Apolipoprotein B–Containing Lipoproteins in the Delivery of Cholesterol to

Tissues

The Role of Apolipoprotein A-I‒Containing Lipoproteins in the Efflux of Cholesterol and its
Delivery to the Liver

Cellular Metabolism of Cholesterol

Secretion of Cholesterol and Bile Acids in the Bile and their Enterohepatic Reabsorption
and Net Fecal Excretion

Role of Lipids in the Development of Atherosclerosis

Thinking Critically—Nutrition Insight Box—Switch-Like Control of SREBP2

Thinking Critically—Clinical Correlation Box—Apolipoprotein E Variants

 Thinking Critically—Clinical Correlation Box—Nature Points to New Methods of Lowering
Cholesterol

Thinking Critically—Clinical Correlation Box—Tangier Disease

Thinking Critically—End of Chapter

Chapter 15. Protein and Amino Acid Metabolism

Overview

Amino Acid Transport in and Out of Cells

Synthesis of Semi-Essential and Essential Amino Acids

Movement of Amino Groups from One Carbon Skeleton to Another

Protein Turnover

Use of Amino Acids for Protein Synthesis

Posttranslational Modification and Translocation or Trafficking of Newly Synthesized

Proteins

Protein Degradation

Use of Amino Acids for Synthesis of Nonprotein Nitrogenous Compounds and Sulfur-
Containing Compounds

Amino Acid Catabolism, Gluconeogenesis, and Excretion of Nitrogen

Changes in Protein Metabolism in Response to Growth, Undernutrition, Physical Activity,

and Hypermetabolic Catabolic Stresses

Thinking Critically – End of Chapter

Chapter 16. Metabolism of Individual Amino Acids

Metabolism of the Nonessential Amino Acids

Essential and Semi-Essential Amino Acids

Thinking Critically—Clinical Correlation Box—Glutamate Dehydrogenase and the

Hyperinsulinism/Hyperammonemia Syndrome

Thinking Critically—Clinical Correlation Box—Modification of Amino Acid Intake to Treat

Genetic Disorders of Amino Acid Metabolism
 Thinking Critically—End of Chapter

UNIT V. Macronutrient Requirements

Chapter 17. Total Energy Requirement: Energy Expenditure, Growth, and

Energy Stores
Energy, Heat, and Work

Energy Expenditure

Whole-Body Energy Expenditure

Use of Macronutrients as Substrates for Growth and Maintenance

Use of Macronutrients as Substrates for Energy Stores

Roles of White and Brown Adipose Tissues in Energy Storage and Energy Expenditure

Estimates of Total Energy Requirements and Physiological Energy Values of Foods

Energy Balance and Regulation of Energy Intake and Energy Expenditure

Thinking Critically—Clinical Correlation Box—Congenital Leptin Deficiency

Thinking Critically–End of Chapter

Chapter 18. Requirements or Recommended Intakes for Carbohydrates and

Lipids
Typical Intakes of Carbohydrates and Lipids

Recommendations for Amount and Type of Carbohydrate in the Diet

Essentiality of n−3 And n−6 Polyunsaturated Fatty Acids (PUFAs)

Recommendations for Amount and Type of Lipids in the Diet

Thinking Critically—Nutrition Insight Box—How Much Added Sugar Do You Consume?

Thinking Critically—End of Chapter

Chapter 19. Protein and Amino Acid Requirements

Classification of Dietary Essential and Nonessential Amino Acids
 Requirements for Dietary Protein

Requirements for Individual Amino Acids

Protein Quality: Amino Acid Composition of the Protein and the Digestibility, Absorbability,
and Utilizability of the Amino Acids in the Protein

Typical Intakes of Protein and Amino Acids and Significance of Protein/Energy Ratios

Effects of Inadequate Protein Intake and Assessment of Protein Status

How Much Protein is too Much Protein?

Other Factors for Consideration in Making Recommendations for Amino Acid and Protein
Intake

Thinking Critically—End of Chapter

Chapter 20. Consequences of Energy Imbalance: Obesity and

Undernutrition
Consequences of Excess Energy Intake: Overweight and Obesity

Metabolic Syndrome

Consequences of Inadequate Energy Intake: Protein-Energy Malnutrition and Starvation

Undernutrition Owing to Increased Energy Expenditure: Hypermetabolic States

Thinking Critically—End of Chapter

UNIT VI. The Vitamins

Chapter 21. Niacin, Riboflavin, and Thiamin

Niacin

Niacin and Pyridine Nucleotide Coenzyme Structure and Nomenclature

Sources, Digestion, and Absorption

Niacin and Tryptophan Metabolism to NAD Coenzymes

Niacin Catabolism and Excretion

Functions of Pyridine Nucleotide Coenzymes in Metabolism

Noncoenzymatic Functions of Niacin

Health Effects of Niacin

Niacin-Responsive Genetic Disorders

Dietary Recommendations, Intake, and Assessment

Toxicity

Riboflavin

Structure, Nomenclature, and Chemistry

Riboflavin Sources, Digestion, Absorption, and Excretion

Conversion of Riboflavin to Coenzymes

Functions of Flavocoenzymes in Metabolism

Health Effects of Riboflavin

Riboflavin-Responsive Genetic Disorders

Dietary Recommendations, Intake, Assessment, and Toxicity

Thiamin

Thiamin and Thiamin Coenzyme Structure and Nomenclature

Food Sources, Digestion, Absorption, and Transport

Conversion of Thiamin to Coenzyme Form

Functions of Coenzymatic Thiamin in Metabolism

Health Effects of Thiamin

Thiamin-Responsive Genetic Disorders

Dietary Recommendations, Intake, Assessment, and Toxicity

Interdependence of B3, B2, and B1

Thinking Critically—Clinical Correlation Box—Case Study of Pellagra in an Adult Woman

Thinking Critically—Clinical Correlation Box—Multiple Acyl-CoA Dehydrogenase

Disorders and Riboflavin

Thinking Critically—Clinical Correlation Box—Infantile Thiamine Deficiency in

Industrialized Nations
 Thinking Critically—End of Chapter

Chapter 22. Folate, Choline, Vitamin B12, and Vitamin B6

Folate

Nomenclature and Structure

Food Sources of Folate

Absorption Of Folate

Bioavailability of Folate

Tissue Uptake and Transport

Excretion

Metabolism and Biological Functions of Folate

Health Effects

Folate Dietary Recommendations

Folate Intake

Assessment and Status Indicators

Choline

Nomenclature and Structure

Food sources

Digestion and Absorption of Choline

Bioavailablity of Choline

Tissue Uptake of Choline

Metabolism of Choline

Excretion of Choline

Biological Functions

Choline Deficiency

Health Effects
Dietary Recommendations for Choline

Factors Affecting Choline Requirements

Dietary Intake of Choline

Assessment and Status Indicators

Toxicity

Vitamin B12

Nomenclature and Structure

Food Sources

Digestion of Vitamin B12

Absorption of Vitamin B12

Bioavailability of Vitamin B12

Transport, Tissue Uptake, Metabolism, and Excretion

Biological Functions of Vitamin B12

Vitamin B12 Deficiency

Dietary Recommendations and Intake

Assessment of Vitamin B12

Vitamin B6

Nomenclature and Structure

Food Sources

Digestion and Absorption

Bioavailablity

Metabolism of Vitamin B6

Distribution

Storage and Turnover

Biological Functions

Vitamin B6 Deficiency
 Health Effects

Dietary Recommendations

Vitamin B6 Intake

Assessment and Status Indicators

Toxicity

Thinking Critically—End of Chapter

Chapter 23. Biotin and Pantothenic Acid

Biotin

Nomenclature and Structure

Food Sources, Digestion, and Absorption

Synthesis of Biotin-Containing Carboxylases (Holoenzymes) and Their Mechanism of

Action

Functions of Biotin-Dependent Carboxylases in Metabolism

Biotin and Gene Regulation

Biotin Catabolism and Excretion

Health Effects Of Biotin

Inborn Errors of Biotin Metabolism

Dietary Recommendations, Intake, and Assessment

Pantothenic Acid

Nomenclature and Structure

Dietary Sources, Digestion, Tissue Uptake, and Excretion

Metabolism of Pantothenic Acid to CoA and ACP

Biological Functions of CoA and ACP

Health Effects of Pantothenic Acid

Dietary Recommendations, Intake, and Toxicity

Thinking Critically—End of Chapter

Chapter 24. Vitamin C
Nomenclature, Structure, and Chemical Properties

Vitamin C Synthesis

Dietary Sources

Supplemental Sources

Intestinal Absorption

Tissue Distribution

Transport of Vitamin C

Enzymatic Functions of Vitamin C

Dioxygenases

Other Enzymes

Nonenzymatic Functions of Vitamin C

Vitamin C Deficiency: Scurvy

Vitamin C in Disease Prevention

Vitamin C in Disease Treatment

Dietary Recommendations and Status Assessment

Vitamin C Intake in the United States

Toxicity

Thinking Critically—End of Chapter

Chapter 25. Vitamin K

Nomenclature and Structure of Vitamin K

Food Sources, Digestion, and Absorption

Tissue Uptake, Metabolism, Transport, and Excretion

Biological Functions of Vitamin K

Vitamin K Antagonists and Clinical Uses

 Health Effects of Vitamin K

Dietary Recommendations for Vitamin K

Vitamin K Deficiency

Assessment of Vitamin K Status

Thinking Critically—Clinical Correlation Box—Warfarin and Pharmacogenetics

Chapter 26. Vitamin E

Nomenclature and Structure of Vitamin E

Food Sources, Digestion, and Absorption

Transport and Tissue Uptake

Hepatic Export and Metabolism of Vitamin E

Biological Functions of Vitamin E

Health Effects of Vitamin E

Dietary Recommendations For Vitamin E

Dietary Intake of Vitamin E

Status Indicators

Toxicity of Vitamin E

Thinking Critically—Nutrition Insight Box—Vitamin E Requirement in Relation to

Polyunsaturated Fatty Acid Intake

Thinking Critically—Clinical Correlation Box—Familial Isolated Vitamin E Deficiency and

α-Tocopherol Transfer Protein

Thinking Critically—End of Chapter

Chapter 27. Vitamin A

Nomenclature, Structure, and Chemical Properties

Food Sources of VITAMIN A

Digestion and Absorption

 Hepatic Metabolism, Export, Transport, and Excretion

Extrahepatic Metabolism of Vitamin A

Intracellular Vitamin A–Binding Proteins

Physiological Functions of Vitamin A

Extranuclear and Nontranscriptional Effects

Health Effects Of Vitamin A

Dietary Recommendations and Intake

Assessment and Status Indicators

Toxicity

Thinking Critically—End of Chapter

Chapter 28. Vitamin D

Nomenclature and Structure

Endogenous Synthesis

Food Sources and Absorption

Metabolism of Vitamin D

Classic Endocrine Functions of Vitamin D

Molecular Actions of Vitamin D

Vitamin D and Health Outcomes

Dietary Recommendations and Intake

Assessment of Vitamin D Status

Toxicity

Thinking Critically—Clinical Correlation Box—Vitamin D–Dependent Rickets Type II

Thinking Critically—End of Chapter

UNIT VII. The Minerals

Chapter 29. Calcium and Phosphorus
Chemistry

Food Sources

Bioavailablity

Intestinal Absorption

Excretion

Non-Skeletal Functions

Calcium and Phosphorus as Components of Mineralized Tissue

Regulation of Calcium and Phosphorus Homeostasis

Health Effects

Medications that May Interfere with Calcium or Phosphate Homeostasis

Dietary Recommendations

Dietary Intake

Assessment and Status Indicators

Toxicity

Thinking Critically—End of Chapter

Chapter 30. Magnesium

Chemistry of Magnesium

Food Sources

Absorption

Magnesium Distribution, Transport, and Excretion

Physiological Roles of Magnesium

Health Effects of Magnesium

Magnesium Deficiency

Recommended Dietary Allowances

 Dietary Intake of Magnesium

Magnesium Toxicity

Thinking Critically—Clinical Correlation Box—Magnesium Depletion in Patients With

Diabetes Mellitus

Thinking Critically—Clinical Correlation Box—Magnesium Deficiency in Chronic Alcohol

Abuse

Chapter 31. Sodium, Chloride, and Potassium

Chemistry

Food Sources

Absorption

Distribution and Compartmentation

Regulation of Sodium, Chloride, and Potassium Balance

Sodium and Chloride Imbalances and their Consequences

Potassium Imbalance and its Consequences

Biological Functions

Chronic Disease Risk

Recommendations and Intake

Thinking Critically—Clinical Correlation Box—Hypertension Genetics: High and Low

Penetrance

Thinking Critically—Clinical Correlation Box—Diarrhea

Thinking Critically—End of Chapter

Chapter 32. Body Fluids and Water Balance

Physiological Functions of Water

Body Water Content And Forces Governing Its Distribution

Forces Determining The Distribution Of Water Between The Intracellular And

Extracellular Fluid Compartments
 Water Balance

Renal Mechanisms For Regulating Loss Of Water In Urine

Regulation of Water Balance

Water Imbalance and Its Consequences

Thinking Critically—Clinical Correlation Box—Heat Acclimatization

Thinking Critically—Clinical Correlation Box—Edema Formation

Chapter 33. Iron

Chemistry of Iron

Proteins Involved in Iron Transport, Storage, and Recycling

Food Sources of Iron

Absorption of Iron

Regulation of Iron Absorption

Cellular Iron Uptake and Export

Intracellular Iron Trafficking, Role of Mitochondria in Heme and Iron-Sulfur Cluster

Cofactor Synthesis, and Intracellular Iron Storage

Biological Functions of Iron

Body Iron Compartments and Daily Iron Exchange

Regulation of Cellular Iron Homeostasis

Systemic Iron Homeostasis

Body Iron Balance

Health Effects: Iron Deficiency

Health Effects: Iron Overload

Dietary Recommendations and Intake

Laboratory Evaluation of Iron Status

Thinking Critically—Clinical Correlation Box—Iron Overload

Chapter 34. Zinc, Copper, and Manganese
Chemistry

Food Sources

Absorption

Tissue Uptake, Metabolism, and Transport

Storage

Excretion

Biological Functions

Health Effects

Dietary Recommendations and Intake

Assessment and Status Indicators

Toxicity

Thinking Critically—Clinical Correlation Box—Genetic Disturbances in Copper

Metabolism

Thinking Critically—Clinical Correlation Box—The Role of Ceruloplasmin in Iron

Metabolism

Chapter 35. Iodine

Chemistry Of Iodine

Sources of Iodine

Absorption, Storage, and Excretion

Thyroid Hormone Synthesis, Metabolism, and Transport

Activation and Deactivation of Thyroid Hormones

Regulation of Thyroid Hormone Status

Mechanism of Action of Thyroid Hormones

Nonnuclear Pathways

Physiological Functions of Thyroid Hormones

 Iodine Deficiency

Effects of Iodine Excess

Dietary Recommendations for Iodine

Dietary Intake

Toxicity

Thinking Critically—End of Chapter

Chapter 36. Selenium

Chemistry of Selenium

Food Sources and Absorption

Tissue Uptake and Distribution

Metabolism OF Selenium

Excretion

Biological Functions

Health Effects

Dietary Recommendations and Intake

Status Indicators and Assessment

Toxicity

Thinking Critically—End of Chapter

Chapter 37. Fluoride

Chemistry of Fluoride

Sources and Absorption

Fluoride Metabolism and Excretion

Health Effects of Fluoride

Dietary Recommendations For Fluoride

 Fluoride Intake

Toxicity

Thinking Critically—End of Chapter

Chapter 38. Molybdenum and the Beneficial Bioactive Trace Elements

Molybdenum

Food Sources

Absorption

Transport and Excretion

Metabolism

Biological Functions

Health Effects

Dietary Recommendations and Intake

Assessment and Status Indicators

Toxicity

Boron

Food Sources

Absorption

Tissue Uptake, Metabolism, Transport, and Excretion

Biological Functions

Health Effects

Dietary Considerations

Assessment and Status Indicators

Toxicity

Chromium

Food Sources and Absorption

 Tissue Uptake, Transport, and Excretion

Biological Functions

Health Effects

Dietary Recommendations and Intake

Assessment and Status Indicators

Toxicity

Silicon

Food Sources and Absorption

Tissue Uptake, Distribution, Transport, and Excretion

Biological Functions

Health Effects

Dietary Considerations

Assessment and Status Indicators

Toxicity

Other Trace Elements

Nickel

Strontium

Vanadium

Other Elements

Thinking Critically—Clinical Correlation Box—Molybdenum Cofactor Deficiencies And

Therapeutic Interventions

Thinking Critically—Clinical Correlation Box—Nutritional Approaches to Mitigate Arsenic

Poisoning

Thinking Critically—Nutritional Insight Box—Using Chronic Disease Risk Reduction As A

Basis For Setting DRIs

Thinking Critically—End of Chapter

Index
Common Abbreviations
Copyright

3251 Riverport Lane

St. Louis, Missouri 63043

Biochemical, Physiological, and Molecular Aspects of Human Nutrition,

Fourth Edition ISBN: 978-0-323-44181-0

Copyright © 2019 by Elsevier, Inc. All rights reserved.

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Notices
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new research and experience broaden our understanding, changes in
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Practitioners and researchers must always rely on their own experience

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or methods they should be mindful of their own safety and the safety of
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With respect to any drug or pharmaceutical products identified, readers

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Contributors
Tracy G. Anthony, PhD, Associate Professor, Department of
Nutritional Sciences, Rutgers University, New Brunswick, New Jersey
Sarah L. Booth, PhD, Senior Scientist, Jean Mayer USDA Human
Nutrition Research Center on Aging, Tufts University, Boston,
Massachusetts
Patsy M. Brannon, PhD, RD, Professor and Director of Cornell
Dietetic Internship, Division of Nutritional Sciences, Cornell University,
Ithaca, New York
Marie A. Caudill, PhD, RD, Professor, Division of Nutritional
Sciences, Cornell University, Ithaca, New York
Gerald F. Combs Jr. PhD
Senior Scientist, Jean Mayer USDA Human Nutrition Research Center on
Aging, Tufts University, Boston, Massachusetts
Professor of Nutrition Emeritus, Cornell University, Ithaca, New York
Robert R. Crichton, PhD, FRSC, Professor, Unite’ Biochimie,
Universite’ Catholique de Louvain, Louvain-la-Neuve, Belgium
Balz Frei, PhD, Director Emeritus and Distinguished Professor
Emeritus, Linus Pauling Institute and Department of Biochemistry and
Biophysics, Oregon State University, Corvallis, Oregon
Xueyan Fu, PhD, Scientist II, Jean Mayer USDA Human Nutrition
Research Center on Aging, Tufts University, Boston, Massachusetts
Arthur Grider, PhD, Associate Professor, Department of Foods and
Nutrition, University of Georgia, Athens, Georgia
Kevin C. Klatt, BA, PhD Candidate, Division of Nutritional Sciences,
Cornell University, Ithaca, New York
Martin Kolisek, RNDr, PhD, Biomedical Center Martin, Department
of Neurosciences, Jessenius Faculty of Medicine in Martin, Comenius
University in Bratislava, Martin, Slovak Republic
Angela M. Leung, MD, MSc, Assistant Professor of Medicine and
Endocrinologist, UCLA David Geffen School of Medicine,
Endocrinologist, Division of Endocrinology, Diabetes, and Metabolism,
VA Greater Los Angeles Healthcare System, Los Angeles, California
Nell I. Matthews, BA, Research Associate (Retired), Department of
Biochemistry and Molecular Biology, University of Arkansas for Medical
Sciences, Little Rock, Arkansas
Alexander Michels, PhD, Research Associate, Linus Pauling Institute,
Oregon State University, Corvallis, Oregon
Donald M. Mock, MD, PhD, Professor, Department of Biochemistry
and Molecular Biology, University of Arkansas for Medical Sciences,
Little Rock, Arkansas
Paul J. Moughan, PhD, DSc, Hon DSc, FRSC, FRSNZ, Distinguished
Professor and Co-Director, Riddet Institute, Massey University,
Palmerston North, New Zealand
Forrest H. Nielsen, PhD, Research Nutritionist Consultant,
Independent (Retired), Grand Forks, North Dakota
Heyjun Park, PhD, Graduate Student, Division of Nutritional Sciences,
Cornell University, Ithaca, New York
Robert S. Parker, PhD, Professor, Division of Nutritional Sciences,
Cornell University, Ithaca, New York
Elizabeth N. Pearce, MD, MSc, Associate Professor of Medicine,
Boston University School of Medicine, Boston, Massachusetts
W. Todd Penberthy, PhD, Continuing Medical Education Writer,
Orlando, Florida
Sue A. Shapses, PhD, RDN, Professor, Department of Nutritional
Sciences, Rutgers University, New Brunswick, New Jersey
Hwai-Ping Sheng, PhD, Honorary Associate Professor (Retired),
Department of Physiology and Department of Pharmacology and
Pharmacy, The University of Hong Kong, Hong Kong SAR, China
Christina M. Stark, MS, RD, CDN, Nutrition Specialist, Division of
Nutritional Sciences, Cornell University, Ithaca, New York
Bruce R. Stevens, PhD, Professor, Department of Physiology and
Functional Genomics, College of Medicine, University of Florida,
Gainesville, Florida
Martha H. Stipanuk, PhD, Professor, Division of Nutritional Sciences,
Cornell University, Ithaca, New York
Jürgen Vormann, PhD, Professor, Institute for Prevention and
Nutrition, Ismaning, Germany
Xiang-Dong Wang, MD, PhD, Professor and Senior Scientist, Jean
Mayer USDA Human Nutrition Research Center on Aging, Tufts
University, Boston, Massachusetts
Gary M. Whitford, PhD, DMD, Regents’ Professor, Department of
Oral Biology, College of Dental Medicine, Augusta University, Augusta,
Georgia
Reviewers
Alison G. Borkowska, PhD, Instructor, Department of Nutritional
Sciences, Pennsylvania State University, University Park, Pennsylvania
Kimberly Buhman, PhD, Associate Professor, Department of Nutrition
Science, Purdue University, West Lafayette, Indiana
Gale B. Carey, PhD, Professor, Department of Molecular, Cellular, and
Biomedical Sciences, University of New Hampshire, Durham, New
Hampshire
Mark A. Kantor, PhD, Office of Nutrition and Food Labeling (ONFL),
U. S. Food and Drug Administration, College Park, Maryland
David Levitsky, PhD, Professor, Division of Nutritional Sciences,
Cornell University, Ithaca, New York
Alfred Merrill, PhD, Professor, School of Biological Sciences, Georgia
Institute of Technology, Atlanta, Georgia
Angela D. Myracle, MPH, PhD, Assistant Professor of Human
Nutrition, School of Food and Agriculture, University of Maine, Orono,
Maine
Eve Essery Stoody, PhD, Lead Nutritionist, USDA Center for Nutrition
Policy and Promotion, Alexandria, Virginia
Hei Sook Sul, PhD, Professor, Doris H. Calloway Chair in Human
Nutrition, Department of Nutritional Sciences and Toxicology, University
of California, Berkeley, California
Jennifer A. Tomesko, DCN, RD, CNSC, Assistant Professor, School of
Health Professions, Rutgers Biomedical and Health Sciences, Newark,
New Jersey
Malcolm Watford, DPhil, Professor, Department of Nutritional
Sciences, Rutgers University, New Brunswick, New Jersey
Michaelann S. Wilke, PhD, Instructor and Laboratory Coordinator,
Department of Agricultural, Food and Nutritional Science, University of
Alberta, Edmonton, Alberta
Jane E. Ziegler, DCN, RD, LDN, Associate Professor, Graduate
Programs in Clinical Nutrition, School of Health Professions, Rutgers
Biomedical and Health Sciences, Newark, New Jersey
Preface
We are pleased to present the fourth edition of Biochemical, Physiological,
and Molecular Aspects of Human Nutrition. Our understanding of nutrition
and its role in health and disease continues to expand rapidly, and much of
this progress has been made at the biochemical, physiological, and
molecular levels. Recognizing that it is difficult for students, instructors,
clinicians, and practitioners to obtain and sustain a deep understanding of
the biology of human nutrition, we have worked to develop a textbook that
provides a comprehensive, accessible, scientifically accurate, and up-to-
date overview of the current understanding of the biological bases of
human nutrition. Individuals specializing in nutrition and its effects on
health will want to have the fourth edition of Biochemical, Physiological,
and Molecular Aspects of Human Nutrition on their shelves as a
convenient resource.
The fourth edition has been extensively rewritten in order to provide a
structure and format that reflects current knowledge. Summary headings
are used throughout the text, which will enable readers to efficiently
review information and recognize the major messages. Many new
illustrations and helpful tables have been included. Chapters on
macronutrient metabolism are now preceded by two new chapters that
present a more integrated view of macronutrient metabolism—an
overview of how macronutrient metabolism is integrated with central
pathways of metabolism and an overview of how macronutrient
metabolism is regulated at the whole-body and cellular levels. In addition,
a new unit covering the nutritional requirements for energy, carbohydrate,
fats, and protein has been added. As editors, we have done our best to
ensure consistency of style and approach so that the individual chapters
and units work together as a whole, especially for those who will use this
text to teach the biology of nutrition or to acquire a solid understanding of
this topic for themselves.
Included within each chapter are feature boxes with Nutrition Insights,
Clinical Correlations, and Historical Tidbits. Significant disease-related
aspects of nutrition are incorporated into the individual chapters and also
are highlighted in many of the feature boxes scattered throughout the
book. In addition, all chapters and many of the feature boxes contain
Thinking Critically questions to encourage application of concepts to
problems and clinical cases. Sample answers to these questions are
available to readers on the Evolve website.

Audience for the Book

Students in nutrition, metabolism, and other life sciences—This book is
written largely for upper-level undergraduate students and graduate
students who have completed studies in organic chemistry, biochemistry,
molecular biology, and physiology. Hence topics are covered at a more
advanced level than in introductory and intermediate textbooks that
assume less background in these supporting disciplines. Nevertheless, we
have made our best effort to make complex material accessible to readers
who are unfamiliar with any topic and to cover topics in a way that allows
the reader to gain an overview of a topic as well as a thorough
understanding of the essential concepts. Particular attention has been
given to the design of figures and choice of tabular material to ensure that
illustrations and tables clarify, extend, and enrich the text.
Instructors in nutrition, metabolism, and other life sciences—Although
the topics are logically arranged for reading from first to last, each chapter
is also written to be a self-contained unit, making it possible for
instructors to use a subset of chapters or a different sequence of
presentation of chapters according to instructional needs. The depth and
breadth of coverage make this text somewhat unique among nutrition
texts. It is an especially good choice for courses on macronutrient
metabolism and for courses on micronutrient (vitamin and mineral)
metabolism. Teaching resources for instructors are on the Evolve website
and include all images from the text.
 Dietitians, clinicians, and other health professionals—Given the broad
availability of scientific and pseudoscientific information to the general
public, it is important that dietitians and other health practitioners have a
solid understanding of the biology of human nutrition and health.
Biochemical, Physiological, and Molecular Aspects of Human Nutrition
provides insights into recommended nutritional practices as well as the
science behind the advice. Many of the Nutrition Insights address
contemporary issues and common misunderstandings, and the Clinical
Correlations explore nutrition and disease connections and medical
nutrition therapy.

Organization of the Book

The text consists of seven units that encompass a traditional coverage of
nutrients by classification (macronutrients, vitamins, and minerals) as
well as the integrated metabolism and use of these nutrients. In
recognition of new paradigms in thinking about nutrition, Unit I considers
the historical foundations of nutrition, changes in how nutrients are being
defined and in how dietary recommendations are being made, and the
potentially beneficial nonnutrient components of food. The macronutrients
or energy-yielding nutrients (carbohydrates, proteins, and lipids) are
discussed in Units II through V. Unit II provides an overview of the
structure and properties of the macronutrients. The digestion and
absorption of the macronutrients are discussed in Unit III, and the
metabolism of the macronutrients is the topic of Unit IV. Unit V covers
current recommendations for intakes of energy, carbohydrates, lipids, and
protein, including how these were determined and ongoing controversies.
The vitamins are discussed in Unit VI. The B vitamins have been grouped
and discussed in three chapters in a manner that facilitates an
understanding of their functions in energy metabolism and other metabolic
processes. The unique functions of vitamins C, K, E, A, and D are
described in individual chapters. The minerals and water are the subjects
of Unit VII; those with well-characterized nutritional or health-related
roles are discussed in detail.
Acknowledgments
Working on the fourth edition of Biochemical, Physiological, and
Molecular Aspects of Human Nutrition has been a very positive experience
for us. All of those who helped us as we worked on this project contributed
to our enjoyment of this work, and we extend our deep gratitude to each of
you.
In particular, our deep appreciation goes to the chapter contributors. The
text is much enriched by the contributions of so many talented researchers
and teachers. Their commitment to making scientific advances available
and accessible to our audiences is clear from the willingness of these busy
individuals to accept the challenge and devote the time and effort required
to see their chapters through the entire process. Their willingness to
respond to queries and to allow the editorial flexibility needed to turn
individual chapters into a coherent and integrated text was superb.
It has been a delight to work with the amazing staff at Elsevier who
handled the publication process. We would like to thank Senior Content
Strategist Sandy Clark, Senior Content Development Specialist Laura
Selkirk, and Senior Project Manager David Stein for their help and support
while working on this edition.
THE EDITORS
Martha H. Stipanuk
Marie A. Caudill
UNIT I
Nutrients
O U T L IN E

Chapter 1. Nutrients: History and Definitions

Chapter 2. Guidelines for Food and Nutrient Intake
CHAPTER 1

Nutrients
History and Definitions
Martha H. Stipanuk, PhD

Nutrients are defined as chemical substances found in foods that are

necessary for human life and growth, maintenance, and repair of body
tissues. It is now commonly accepted that proteins, fats, carbohydrates,
vitamins, minerals, and water are the major nutritional constituents of
foods.

Discovery of the Nutrients

Before the chemical nature of food was understood, food was believed to
be made up of nutriment, medicines, and poisons. In ancient Greece
(∼500–300 BC), differences in the physical properties of foods and in
their content of medicinal and toxic substances were recognized. The role
of diet in the causation and treatment of disease was recognized, as
evidenced by the use of liver to treat night blindness. However, the
physicians of this era had no understanding of the chemical nature of
foods and believed that foods contained only a single nutritional principle
that was called “aliment.” The belief that foods contained a single
nutritional principle persisted for more than two millennia up until the
19th century and impeded progress in understanding nutrition. In some
ways the ancient understanding of food as made up of nutriment,
medicines, and poisons is still appropriate in that foods do contain
nutrients, substances beneficial to health, and substances that have adverse
effects on health. However, we now know that each of the three principles
in fact includes many different chemical compounds.

Early Observations
Some recorded observations made during the 17th and 18th centuries
hinted at scientific progress to come. For example, during the 1670s
Thomas Sydenham, a British physician, observed that a tonic of iron
filings in wine produced a clinical response in patients with chlorosis, a
condition now recognized as hypochromic or iron-deficiency anemia
(McCollum, 1957). In 1747 James Lind of Scotland, while serving as a
naval surgeon, conducted a clinical trial of various proposed treatments of
sailors who were ill with scurvy. He observed that consumption of citrus
fruits (oranges and lemons), but not other typical foods and medicines,
cured the disease (Carpenter, 1986). Nevertheless, chlorosis and scurvy
were not viewed as nutritional diseases, and the concept that a disease
might be caused by a deficit of a substance that was nutritionally essential
did not exist. Before 1900, toxins, heredity, and infections, but not yet
nutrition, were recognized as causes of disease.

Recognition that Food is a Source of Specific

Nutrients
During the 1800s the concept that food was a source of several specific
nutrients gradually developed.
During the 19th century, food was recognized to contain several
components: protein, energy-yielding substances, and a few minerals.
By the early 1800s the elements carbon, nitrogen, hydrogen, and oxygen
were recognized as the primary components of food, and the need for the
carbon-containing components of food as a substrate for combustion (heat
production) was recognized (Carpenter, 2003a). Protein was identified as a
specific essential dietary component by the observations of François
Magendie in Paris in 1816. Magendie showed that dogs fed only
carbohydrate or fat lost considerable body protein and weight within a few
weeks, whereas dogs fed foods that contained nitrogen (protein) remained
healthy. In 1827 William Prout, a physician and scientist in London,
proposed that the nutrition of higher animals could be explained by their
need for proteins, carbohydrates, and fats, and this explanation was widely
accepted. During the next two decades, the need of animals for several
mineral elements was demonstrated, and at least six mineral elements
(calcium, phosphorus, sodium, potassium, chloride, and iron) had been
established as essential for higher animals by 1850 (Harper, 1999;
Carpenter et al., 1997).
The 19th-century German chemist Justus von Liebig postulated that
energy-yielding substances (carbohydrates and fats) and proteins, together
with a few minerals, represented the essentials of a nutritionally adequate
diet, and he proposed that the nutritive value of foods and feeds could be
predicted from knowledge of their gross chemical composition. Liebig
prepared tables of food values based on this concept—work that was
facilitated by the work of Wilhelm Henneberg, who devised the Weende
system, known as proximate analysis, for analyzing foods and feeds for
protein, fat, fiber, nitrogen-free extract (carbohydrate), and ash
(McCollum, 1957). Throughout the remainder of the 19th century,
nutritional thinking continued to be dominated by the belief that sources
of energy, protein, and a few minerals were the sole principles of a
nutritionally adequate diet.
The lack of the concept that nutrient deficiency could be a cause of
disease impeded the recognition that food contained additional
nutrients essential for health. Despite the dominance of Liebig’s views
during the mid to late 19th century, it should be noted that the validity of
his assumptions was challenged during this period (McCollum, 1957). In
1843 Jonathan Pereira in England stated that diets containing a wide
variety of foods were essential for human well-being, whereas diets
containing only a few foods were associated with the acquisition of
diseases such as scurvy. Jean-Baptiste Dumas, based on his observation
that an artificial milk formula that contained all of the known dietary
constituents failed to prevent deterioration of the health of children during
the siege of Paris (1870-1871), also questioned the validity of Liebig’s
assumptions. In addition, Nikolai Lunin (∼1881), who worked in Gustav
von Bunge’s laboratory in Dorpat, Estonia, conducting studies with mice
in an effort to identify inadequacies in the mineral component of purified
diets, demonstrated that mice fed a diet composed of purified proteins,
fats, carbohydrates, and a mineral mixture survived less than 5 weeks,
whereas mice that received milk or egg yolk in addition to the purified
components remained healthy throughout the experiment. Lunin
concluded that milk must contain small quantities of other unknown
substances essential to life, but von Bunge apparently did not encourage
him or subsequent students in his laboratory who made similar
observations to investigate what the active factor in milk might be. The
Liebig–von Bunge view that nutritional requirements consisted only of
protein, energy-yielding substances, and a few minerals still had such hold
on scientific thought that, rather than consider that these observations
might point to the presence of other essential nutrients in foods, the
inadequacies of the purified diets were attributed to mineral imbalances,
failure to supply minerals as organic complexes, or lack of palatability
(Wolf and Carpenter, 1997).

H ist o rica l Tid b it

The Connection between Combustion and Respiration: The
Experiments of Antoine Lavoisier
 Nearly 300 years before Lavoisier, during the 16th century, the artist
and scientist Leonardo da Vinci noted the part played by air in
combustion. The ancients realized that air was necessary for burning but
did not understand the nature of the combustion process. Leonardo
arranged deliberate experiments on enclosed combustion and arrived at
the correct answer to a problem that continued to worry experimenters
for years afterward. In manuscripts deposited as the Codex Leicester,
Leonardo noted that “air is consumed by the introduction of the fire.”
He also noted, in the Codice Atlantico, that “where flame cannot live,
no animal that draws breath can live,” clearly correlating the
phenomenon of combustion with the one of animal respiration. Like
Leonardo, Robert Fludd and John Mayow came to their own correct
Other documents randomly have
different content
on the cliff under the walls of Turnberry Castle, whence the annexed sketch
(Fig. 96) was taken. The lower andesite (a) is highly amygdaloidal towards the
top, and is traversed in all directions with irregular veins and nests of sandstone
which can be traced upward to the bed (b), consisting of sandstone, but so full
of lumps or slags of amygdaloidal andesite that one is here and there puzzled
whether to regard it as a sedimentary deposit, or as the upper layer of clinkers
of a lava-stream strewn with sand. Above this fragmentary layer lies another
bed of andesite (c) of a coarsely amygdaloidal structure, which encloses
patches of the underlying sandstone. It passes upward, in a space of from four
to six feet, into a mass of angular scoriaceous fragments (d) of all sizes up to
blocks 18 inches in length cemented in a vein-stuff of calcite, chalcedony and
quartz. This brecciated structure ascends for about 13 or 14 feet, and is then
succeeded by a greenish compact andesite (e), which further north becomes
amygdaloidal and much veined with sandstone, passing into a breccia of lava
fragments and sandstone.

Fig. 97.—Lenticular form of a brecciated andesite

(shown in Fig. 96), Turnberry, Ayrshire.

MAP OF THE VOLCANIC DISTRICTS OF THE LOWER OLD RED SANDSTONE

OF "LAKE CALEDONIA" IN CENTRAL SCOTLAND & NORTH EAST IRELAND

The Edinburgh Geographical Institute Copyright J. G. Bartholomew.

Click on map's left, middle, or right to view larger sized version.

The remarkable brecciated band (d) in this cliff, though 13 or 14 feet in the
centre, immediately thins out on either side, until in the course of a few yards it
completely disappears and allows the lavas c and e to come together, as shown
in Fig. 97. We may suppose that this section reveals the structure of the
terminal portion of a highly viscous lava which was shattered into fragments as
it moved along under water.
No clear evidence of the sites of any of the volcanic vents has yet been
detected in the Old Red Sandstone of Ayrshire. Possibly some of the numerous
felsitic bosses to the south-west of Dalmellington may partly mark their
positions. But the sills connected with the volcanic series are well exposed in
the 12 miles of hilly ground between Dalmellington and Barr. Two groups of
intrusive sheets may there be seen. The most numerous consist of pale or dark-
pink felsite, often full of crystals of mica. They form prominent hills, such as
Turgeny, Knockskae and Garleffin Fell. The second group comprises various
diabase-sheets which have been intruded near the base of the red sandstones
and conglomerates, over a distance of seven miles on the north side of the
Stinchar Valley above Barr. They attain their greatest development on Jedburgh
Hill, where they form a series of successive sills, the largest of which unite
northwards into one thick mass and die out southward among the sandstones
and conglomerates.
 CHAPTER XXI
VOLCANOES OF THE LOWER OLD RED
SANDSTONE OF THE CHEVIOT HILLS, LORNE,
"LAKE ORCADIE" AND KILLARNEY

THE CHEVIOT AND BERWICKSHIRE DISTRICT

In the south-east of Scotland, and extending thence into the north
of England, the remains of several distinct volcanic centres of the
Lower Old Red Sandstone may still be recognized. Of these the
largest and most interesting forms the mass of the Cheviot Hills; a
second has been partially dissected by the sea along the coast south
from St. Abb's Head; while possibly relics of others may survive in
detached bosses of eruptive rock which rise through the Silurian
formations of Berwickshire. The water-basin in which these volcanic
groups were active was named by me "Lake Cheviot,"[371] to
distinguish it from the other basins of the same geological period
(Map I.).
[371] Trans. Roy. Soc. Edin. xxviii. (1878), p. 354.
The volcanic rocks of the Cheviot Hills, though their limits have been
reduced by faults, unconformable overlap of younger formations and
severe denudation, still cover about 230 square miles of ground, and
rise to a height of 2676 feet above the sea. As they have been
mapped in detail by the Geological Survey, both on the English and
the Scottish sides of the Border, their structure is now known.[372]
No good horizontal section, however, has yet been constructed to
show this structure—a deficiency which, it is hoped, may before long
be supplied.
 [372] The Geology of the Cheviot Hills is comprised in Sheets 108 N.E.,
109 N.W., and 110 S.W. of the Geological Survey of England and
Wales, and in Sheets 17, 18 and 26 of the Geological Survey of
Scotland. For descriptive accounts the Memoirs to some of these
Sheets may be consulted, particularly "Geology of the Cheviot Hills"
(English side), by C. T. Clough (Mem. Geol. Surv. 1888); "Geology of
Otterburn and Elsdon," by H. Miller and C. T. Clough (Mem. Geol.
Surv. 1887); "Geology of Part of Northumberland between Wooler
and Coldstream," by W. Gunn and C. T. Clough, with Petrographical
Notes by W. W. Watts (Mem. Geol. Surv. 1895). Other descriptions
have been published by Professor James Geikie, Good Words, vol.
xvii. (1876), reprinted in Fragments of Earth-lore (1893), and by Prof.
Lebour, Outlines of the Geology of Northumberland, 2nd edit. 1886.
For the petrography of the rocks consult Mr. J. J. H. Teall, Geol. Mag.
1883, pp. 100, 145, 252, 344; 1884, p. 226; 1885, p. 106; Proc.
Geol. Assoc. ix. (1886) p. 575; and his British Petrography, 1888; Dr.
J. Petersen, Mikroskopische und chemische Untersuchungen am
Enstatit-porphyrit aus den Cheviot Hills, Inaugural Dissertation, Kiel,
1884.
This volcanic pile, consisting mainly of bedded andesites which rest
unconformably on the upturned edges of Wenlock shales and grits,
presents a most typical display of the lavas of the Lower Old Red
Sandstone. These rocks range from vitreous or resinous pitchstone-
like varieties to coarsely porphyritic forms, on the one hand, and to
highly vesicular and amygdaloidal kinds, on the other. Analyses of
some of these rocks, and an account of their petrography, have
already been given.
The lavas are often separated by thin partings of tuff, and their
upper surfaces show the fissured character with sandstone infillings,
so characteristic among the lavas of "Lake Caledonia."[373] Tuffs form
a very subordinate part of the whole volcanic series. One of the
most important bands is a thick mass at the base of the series, lying
immediately on the highly inclined Silurian shales. The fragments are
generally of a fine-grained purple mica-andesite, often two or three
feet and sometimes at least five feet long. For a few feet near the
bottom of this mass of tuff, pieces of Silurian shale an inch in length
may be noticed. Mr. Clough remarks that distinct bedding is not
usual among the tuffs. Though no doubt most of the fragmental
materials really lie intercalated between successive lava-streams, yet
some of the isolated patches of coarse volcanic breccia may mark
the sites of eruptive vents. One such probable neck has been
mapped on the Scottish side between Cocklawfoot, at the head of
the Bowmont Water, and King's Seat, while others may perhaps
occur among the detached patches that have been observed on the
Northumbrian side. No thick conglomerates or sandstones have been
noticed in the Cheviot District. The volcanic eruptions appear to have
usually succeeded each other without the spread of any notable
amount of ordinary detritus over the floor of the water-basin. It is
difficult to estimate the total thickness of volcanic material here piled
up, but it probably amounts to several thousand feet. The top of the
series is not visible, having been partly removed by denudation and
partly buried under the Carboniferous formations.
[373] Clough, Geology of the Cheviot Hills, p. 15.
It will thus be seen that the Cheviot area stands apart from the
other volcanic districts of the Lower Old Red Sandstone in the great
relative thickness of its accumulated lavas, the comparative thinness
of its tuffs, and the absence of the thick intercalations of coarse
conglomerate so abundantly developed among the volcanic series all
over Central Scotland. But there is yet another characteristic in
which this area is pre-eminently conspicuous. In the heart of the
andesites lies a core of augite-granitite, around which these rocks
are traversed with dykes.
This interesting granitic boss rises into the highest summit of the
whole Cheviot range, and covers an area of rather more than 20
square miles. While its petrographical characters have been
described by Mr. Teall, its boundary has been mapped by Mr. Clough,
who found the line difficult to trace, owing partly to the prevalent
covering of peat, and partly to the jagged and irregular junction
caused by the protrusion of dykes from and into the boss. He
obtained evidence that the granite has broken through the bedded
andesites, and that it is in turn traversed by dykes composed of a
material indistinguishable from that of some of the flows. He
therefore considered that it is essentially of the same age as the rest
of the volcanic series, and "not improbably the deep-seated source
of it."[374] Mr. Teall also, from a chemical and microscopical
examination of the rocks, drew a similar conclusion.[375]
[374] Op. cit. p. 24.
[375] Geol. Mag. 1885, p. 106.
The andesites around the granite have undergone contact-
metamorphism, but the nature and extent of the change have not
yet been studied. There occur around the granite many dykes of
felsite and quartz-felsite, to the petrographical character of which
reference has already been made. But the most abundant and
remarkable dykes of the district are those of a reddish mica-
porphyrite, of which Mr. Clough has mapped no fewer than forty,
besides those in the granitic area. He has called attention to the
significant manner in which all the dykes of the district tend to point
in a general way to the great core of granite, as if that were the
nucleus from which they had radiated.[376]
[376] Op. cit. pp. 26-28.
The central granite of the Cheviot Hills, with its peripheral dykes, has
no accompanying agglomerates nor any decided proof that it ever
communicated with the surface. When, however, we consider its
petrographical and chemical constitution, its position as a core
among the bedded lavas, and the intimate way in which it is linked
with these rocks by the network of dykes, we are, I think, justified in
accepting the inference that it belongs to the volcanic series. It
possesses some curious and interesting features in common with the
great granophyre bosses of Tertiary age in the Inner Hebrides. Like
these it has no visible accompaniment of superficial discharges. Yet
it may have ascended by means of some central vent or group of
vents which, offering to it a weak part of the crust, allowed it to
communicate with the surface and give rise to the outflow of lavas
and fragmental ejections. In any case, it affords us a most
interesting and instructive insight into one of the deeper-seated
ducts of a volcanic region, and the relation of a volcanic focus to the
ascent of the granitic magma.

About twenty miles to the north of the Cheviot Hills, and separated
from them by the Carboniferous and Upper Old Red Sandstones
which spread across the broad plain of the Merse, a group of
volcanic rocks has been laid open in a singularly instructive manner
along the coast of Berwickshire, between the village of Eyemouth
and the promontory of St. Abb's Head. Not only the actual vents, but
the lavas and tuffs connected with them, have there been admirably
dissected by the forces of denudation.
That this volcanic area was quite distinct from that of the Cheviot
Hills may be inferred from its coarse agglomerates, and from the fact
that when the rocks are followed inland in a south-westerly
direction, that is, towards the Cheviot area, they are found to
diminish in thickness and to disappear among the ordinary
sediments. For the same reason we may regard the area as
independent of any vents which may have risen further west about
Cockburn Law and the Dirrington Laws. Unfortunately, however, only
a small part of the area comes into view, the rest of it lying beneath
the waters of the North Sea.[377]
[377]This area lies in Sheet 34 Geological Survey of Scotland, and
was described by myself in the Memoir to accompany that Sheet
("Geology of Eastern Berwickshire," 1864, p. 20). More recently the
shore between St. Abb's Head and Coldingham has been re-mapped
by Professor James Geikie who has also studied the microscopic
character of the rocks, Proc. Roy. Soc. Edin. xiv. (1887).
Of the several vents dissected along this coast-line, one may be seen
at Eyemouth, filled with a very coarse tumultuous agglomerate of
andesite fragments embedded in a compact felspathic matrix,
through which are scattered broken crystals of felspar, and imperfect
tabular crystals of black mica. Another of similar character is
exposed for more than a mile and a half along the shore at
Coldingham. It contains blocks, sometimes more than a yard in
diameter, of different varieties of andesite, and, as at Eyemouth, is
much invaded by veins and bosses of intrusive andesite.

Fig. 98.—Section across the volcanic area of St. Abb's

Head (after Prof. J. Geikie).
11. Silurian formations; 2. Lower Old Red Conglomerate
and Sandstone; 3 3. Sheets of andesitic lava; 4. Volcanic
tuffs, largely composed of scoriæ in the higher parts; 5.
Volcanic agglomerate of neck on shore; 6. Intrusive
andesites. f, Fault.
To the north of Coldingham, a series of bedded volcanic rocks which
form the picturesque headland of St. Abb's Head, are, according to
the estimate of Professor James Geikie, about 1000 to 1200 feet
thick, but neither their bottom nor their top is seen. The same
observer found them to consist of three groups of andesite sheets
separated and overlain by bedded tuffs. The lowest lavas have their
base concealed under the sea, and are covered by a thick band of
coarse agglomeratic tuff, above which lies the second group of
andesites, about 250 feet thick. An intercalation of various tuffs from
40 to 50 feet thick then succeeds, followed by the third lava-group,
250 or 300 feet in depth. The highest member of the series is a
mass of bedded tuffs some 400 feet thick.
The andesites lie in beds varying from about 15 to about 50 feet or
more in thickness. They are fine-grained, purplish-blue, or greyish-
blue, often reddish rocks, of the usual type. Generally rather close-
grained, they are not as a rule very porphyritic, but often highly
scoriaceous and amygdaloidal, especially towards the top and
bottom of each bed. The more slaggy portions are sometimes so
filled in with fine tuff that the rock might be mistaken for one of
fragmental origin.
The bedded tuffs are usually well stratified deposits. The most
important band of them is that which forms the highest member of
the volcanic series. It consists of successive beds that vary from fine
red mudstones up to volcanic breccias with blocks one foot or more
in diameter. The materials have been derived from the explosion of
andesitic lavas. Most of the lapilli are vesicular or amygdaloidal, and
many of them have evidently come from vitreous scoriaceous lavas.
Professor Geikie remarks that "from their highly vesicular character,
they might well have floated in water at the time of their ejection—
they are in short mere cinders." He could detect no trace of ordinary
sediment in the matrix, the whole material being thoroughly volcanic
in origin.
The lavas, tuffs and agglomerates have been abundantly invaded by
intrusive rocks, chiefly andesites.[378]
[378] See Prof. J. Geikie, op. cit.
The agglomerates of this Berwickshire coast extend for a short way
inland from the Coldingham and Eyemouth vents, but the fragmental
material soon becomes finer and more water-rolled, and assumes a
distinctly stratified structure, as it is gradually and increasingly
interleaved with layers of ordinary sediment. Hence in passing
towards the south-west, away from the coast-line, we are obviously
receding from the vents of eruption and entering into the usual non-
volcanic deposits of the time. That these deposits belong to the
Lower Old Red Sandstone was first ascertained during the progress
of the Geological Survey in this district by the discovery of abundant
plant-remains in the form of linear grass-like strips, and also pieces
of Pterygotus in some of the green shales interstratified among fine
tuffs and ashy sandstones.[379] Before the volcanic detritus
disappears from the strata as they are followed in a south-westerly
direction, the whole series is unconformably overlain by the Upper
Old Red Sandstone. The lower division of the formation is not again
seen until it rises from under the southern margin of the plain of the
Merse into the Cheviot Hills.
 [379]"Geology of Eastern Berwickshire," Mem. Geol. Surv. Scotland
(1864), pp. 26, 27, 57.
About ten miles to the south-west of the large Coldingham neck the
great boss of Cockburn Law and Stoneshiel Hill rises out of the
Silurian rocks.[380] Five miles still further in the same direction the
group of the beautiful cones of Dirrington (Fig. 70) overlooks the
wide Merse of Berwickshire,[381] and six miles to the north of these
hills, in the very heart of Lammermuir, lies the solitary boss of the
Priestlaw granite.[382] To these protrusions of igneous material
reference has already been made as possible volcanic vents
connected with the eruptions of the Lower Old Red Sandstone. As
regards their age they must certainly be younger than the
Llandovery rocks which they disrupt, and older than the Upper Old
Red Sandstone, of which the conglomerates, largely made from their
debris, lie on them unconformably. It seems therefore probable that
these great bosses may form a part of the volcanic history of the
Lower Old Red Sandstone period. But no positive proof has yet been
obtained that any one of them was the site of an eruptive vent, and
no trace has been detected around them of any lavas or tuffs which
might have proceeded from them.
[380]See "The Geology of Eastern Berwickshire" (Sheet 34), Mem.
Geol. Surv. Scotland (1864), p. 29.
[381]These hills are chiefly represented in Sheet 25. But see "The
Geology of East Lothian," Mem. Geol. Surv. Scotland (1866), p. 26.
"Geology of East Lothian," Mem. Geol. Surv. Scotland, p. 15, and
[382]
authorities there cited.

"THE LAKE OF LORNE"

 Fig. 99.—View of terraced andesite hills resting on massive
conglomerate, south of Oban.
The basin of Lorne has not yet been carefully examined and
described, though various writers have referred to different parts of
it (Map I.). My own observations have been too few to enable me to
give an adequate account of it. The volcanic sheets of this area form
a notable feature in the scenery of the West Highlands, for they are
the materials out of which the remarkable terraced hills have been
carved, which stretch from Loch Melfort to Loch Creran (Fig. 99),
and which reappear in picturesque outliers among the mountains
traversed by Glen Coe. Between the ancient schists that form the
foundation-rocks of this district and the base of the volcanic series,
lies a group of sedimentary strata which in the western part of the
district must be 500 feet thick. This group consists of exceedingly
coarse breccias at the bottom, above which come massive
conglomerates, volcanic grits or tuffs, fine sandstones and courses of
shale. While the basement-breccias are composed mainly of detritus
of the underlying schists, including blocks six feet long, they pass up
into coarse conglomerates made up almost entirely of fragments of
different lavas (andesites, diabases, etc.), and more than 100 feet
thick, which often show little or no trace of stratification, but break
up into large quadrangular blocks by means of joints which cut
across the imbedded boulders. These volcanic conglomerates form
some of the more conspicuous features of the coast to the south
and north of Oban, and are well exposed in the Isle of Kerrera. They
offer many points of resemblance to those of Lake Caledonia, but no
certain proof has been noted that they belong to the Lower Old Red
Sandstone. They have obviously been derived from lava-sheets that
were exposed to strong breaker-action, which at the same time
transported and rounded blocks of granite, schist and other
crystalline rocks derived from the adjacent hills. During the intervals
of quieter sedimentation indicated by the fine sandstones and
shales, volcanic explosions continued, as may be seen by the
occurrence of occasional large bombs which have fallen upon and
pressed down the fine ashy silt that was gathering on the bottom.
It would seem from the characters of some of the strata in this
sedimentary series that over the area of deposit portions of the
shallower waters were occasionally laid bare to the sun and air.
Among the conglomerates there lie certain bands of reddish sandy,
ripple-marked, sun-cracked and rain-pitted shales and fine
sandstones. Such accumulations, indicative of the ultimate exposure
of fine sediment that silted up hollows in the great banks of coarse
shingle, may be noticed at the south end of the Island of Kerrera, on
at least two horizons which are separated from each other by thick
masses of conglomerate and fine felspathic grit. We may infer,
therefore, that the fine littoral mud, which gathered during pauses in
the heaping up of the coarse gravel and shingle, was occasionally
laid dry. Similar strata may be observed behind Oban, where the
alternation of exceedingly fine sediment and granular ashy bands is
well exhibited.

Fig. 100.—Section of lava-escarpment

at Beinn Lora, north side of mouth
of Loch Etive, Argyllshire.
1. Phyllites; 2. Thick conglomerate;
3. Successive sheets of andesite.
But the explosions that gave rise to the volcanic materials so largely
represented in these lower conglomerates, were merely preliminary
to those which led to the outflow of the great sheets of lava that
now constitute so large a part of the hills of Lorne. In the few
traverses which I have made across different parts of this district I
have noted the general resemblance of the lavas to those of the
Lower Old Red Sandstone of the Midland Valley of Scotland, their
bedded character, and the occurrence of occasional layers of
stratified material between them. The prominent features of these
rocks, and their relation to the volcanic conglomerates below them,
and to the underlying slates and schists are well displayed on Beinn
Lora at the mouth of Loch Etive (Fig. 100). There the black slates of
the district are unconformably covered by the coarse volcanic
conglomerate, formed chiefly of blocks of andesite, cemented in a
hard matrix of similar composition. About 150 or 200 feet of this
material underlie the great escarpment of the lavas, which here rise
in successive beds to the top of the hill, 1000 feet above its base.
On the south side of Loch Etive the base of the volcanic series, with
its underlying conglomerate, may be followed westward to Oban and
thence southward to Loch Feochan. The lavas cover most of the
ground from the western shore eastwards to near Loch Awe. But this
area is still very imperfectly known. The Geological Survey, however,
has now advanced to this part of the country, so that we shall before
long be in possession of more detailed information regarding the
character and sequence of its volcanic history and the geological age
of the eruptions.
Mr. H. Kynaston, who has begun the mapping of the eastern portion
of the district, finds that there, as further west, the bottom of the
volcanic series is generally a breccia or conglomerate. He has met
with two leading types among the lavas, the more abundant being
strongly vesicular, the other more compact. He has observed also
numerous dykes and sills of intrusive porphyrite, trending in a
general N.N.E. and S.S.W. direction, and pointing towards the great
granite mass of Ben Cruachan.[383]
 [383] Ann. Report Geol. Surv. (1895), p. 29 of reprint.
Mr. R. G. Symes has traced the volcanic series to the north and
south of Oban. While visiting with him part of his ground, I was
much struck with the evidence of an intrusive mass at the base of
the volcanic series in the Sound of Kerrera. A prominent feature on
the east side of the channel, known as Dun Uabairtich and 270 feet
high, consists of andesite which appears to combine both a central
boss and a sill. The rock breaks through the black slates and the
overlying conglomerates and sandstones, and has wedged itself into
the unconformable junction between the two formations. It is
beautifully columnar on its sea-covered face, some of the columns
being 120 feet or more in length, and gently curved.

"LAKE ORCADIE"
We now cross the whole breadth of the Scottish Highlands in order
to peruse the records of another of the great detached water-basins
of the Old Red Sandstone, which for the sake of brevity of reference
I have named and described as "Lake Orcadie" (Map I.). This area
has its southern limits along the base of the hills that enclose the
wide Moray Firth. It spreads northward over the Orkney and much of
the Shetland Islands, but its boundaries in that direction are lost
under the sea. In the extensive sheet of water which spread over all
that northern region the peculiar Caithness Flags, with their
associated sandstones and conglomerates, were deposited to a total
depth of 16,000 feet. A sigillaroid and lycopodiaceous vegetation
flourished on the surrounding land, together with ferns, Psilophyton
and conifers. The waters teemed with fishes of which many genera
and species have now been described. The remains of these
creatures lie crowded upon each other in the flagstones in such a
manner as to indicate that from time to time vast quantities of fish
were suddenly killed. Not impossibly, these destructions may have
been connected with the volcanic activity which has now to be
described.
In the year 1878 I called attention to the evidence for the existence
of contemporaneous volcanic rocks in the Old Red Sandstone north
of the range of the Grampians, and specially noted three localities
where this evidence could be seen—Strathbogie, Buckie and
Shetland.[384] Since that time Messrs. B. N. Peach and J. Horne have
added a fourth centre in the Orkney Islands. At present, therefore,
we are acquainted with the records of four separate groups of
volcanic vents in Lake Orcadie.
[384] Trans. Roy. Soc. Edin. xxviii. (1878).
The southern margin of this water-basin appears to have indented
the land with long fjord-like inlets. One of these now forms the vale
of Strathbogie, which runs into the hills for a distance of fully 20
miles beyond what seems to have been the general trend of the
coast-line. In this valley I found a bed of dark vesicular diabase
intercalated among the red sandstones and high above the base of
the formation, as exposed on the west side of the valley near Burn
of Craig. On the east side a similar band has since been mapped for
the Geological Survey by Mr. L. Hinxman who has traced it for some
miles down the Strath.[385] This latter band, as shown in Fig. 101,
lies not far above the bottom of the Old Red Sandstone of this
district, and is thus probably distinct from the Craig outcrop. There
would thus appear to be evidence of two separate outflows of basic
lava in this fjord of the Old Red Sandstone period.
[385] See Sheet 76 of the Geological Survey of Scotland.

Fig. 101.—Section across Strathbogie, below Rhynie,

showing the position of the volcanic band.
1. Knotted schists; 2. Diorite intrusive in schists; 3. Old
Red Conglomerate; 4. Volcanic band; 5. Shales with
calcareous nodules; 6. Sandstones of Rhynie; 7. Shales
and sandstones. f, Fault.
No vestige has here been found of any vent, nor is the lava
accompanied with tuff. The eruptions took place some time after the
earlier sediments of the basin were accumulated, but ceased before
the thick mass of upper sandstones and shales was deposited. A
section across the valley gives the structure represented in the
accompanying diagram (Fig. 101).
Twenty-five miles further north a still smaller andesite band has been
detected by Mr. J. Grant Wilson among the sandstones and
conglomerates near Buckie.[386] It is a truly contemporaneous flow,
for pebbles of it occur in the overlying strata. But again it forms only
a solitary bed, and no trace of any accompanying tuff has been met
with, nor of the vent from which it came. Both this vent and that of
Strathbogie must have been situated near the southern coast-line of
the lake.
[386]See Sheet 95 of the Geological Survey of Scotland and Trans.
Roy. Soc. Edin. vol. xxviii. (1878), p. 435.
At a distance of some 90 miles northward from these Moray Firth
vents another volcanic district lies in the very heart of the Orkney
Islands.[387] The lavas which were there ejected occur at the south-
eastern corner of the island of Shapinshay, where they are regularly
bedded with the flagstones. They consist of dark green olivine-
diabases with highly amygdaloidal and vesicular upper surfaces.
Their thickness cannot be ascertained, as their base is not seen, but
they have been cut by the sea into trenches which show them to
exceed 30 feet in depth. The position of the vent from which they
came has not been ascertained. Neither here nor in the Moray Firth
area do any sills accompany the interbedded sheets, and in both
cases the volcanic action would seem to have been of a feeble and
short-lived character.
[387] Messrs. B. N. Peach and J. Horne, Proc. Roy. Phys. Soc. Edin.
vol. v. (1879), p. 80.
Much more important were the volcanoes that broke out nearly 100
miles still further north, where the Mainland of Shetland now lies. I
shall never forget the pleasure with which I first recognized the
traces of these eruptions, and found near the most northerly limits
of the British Isles proofs of volcanic activity in the Lower Old Red
Sandstone. Since my observations were published,[388] Mr. Peach,
who accompanied me in Shetland, has returned to the district, and,
in concert with his colleague Mr. Horne, has extended our knowledge
of the subject.[389] The chief vent or vents lay towards the west and
north-west of the Mainland and North Mavine; others of a less active
and persistent type were blown out some 25 miles to the east,
where the islands of Bressay and Noss now stand. In the western
district streams of slaggy andesite and diabase with showers of fine
tuff and coarse agglomerate were ejected, until the total
accumulation reached a thickness of not less than 500 feet. The
volcanic eruptions took place contemporaneously with the deposition
of the red sandstones, for the lavas and tuffs are intercalated in
these strata. The lavas and volcanic conglomerates are traceable
from the southern coast of Papa Stour across St. Magnus' Bay to the
western headlands of Esha Ness, a distance of more than 14 miles.
They have been cut by the Atlantic into a picturesque range of cliffs,
which exhibit in some places, as at the singular sea-stalk of
Doreholm, rough banks of andesitic lava with the conglomerate
deposited against and over them, and in other places, as along the
cliffs of Esha Ness, sheets of lava overlying the conglomerates.
[388] Trans. Roy. Soc. Edin. vol. xxviii. (1878), p. 418.
[389] Ibid. vol. xxxii. (1884), p. 359.
No trace of any vents has been found in the western and chief
volcanic district, but in Noss Sound a group of small necks occurs,
filled with a coarse agglomerate composed of pieces of sandstone,
flagstone and shale. Messrs. Peach and Horne infer that these little
orifices never discharged any streams of lava. More probably they
were opened by explosions which only gave forth vapours and
fragmentary discharges, such as a band of tuff which is intercalated
among the flagstones in their neighbourhood.
But one of the most striking features of the volcanic phenomena of
this remote region is the relative size and number of the sills and
dykes which here as elsewhere mark the latest phases of
subterranean activity. Messrs. Peach and Horne have shown us that
three great sheets of acid rocks (granites and spherulitic felsites, to
which reference has already been made, p. 292) have been injected
among the sandstones and basic lavas, that abundant veins of
granite, quartz-felsite and rhyolite radiate from these acid sills, and
that the latest phase of igneous action in this region was the
intrusion of a series of bosses and dykes of basic rocks (diabases)
which traverse the sills.

The Killarney District

In the south of Ireland the Upper Silurian strata are followed
upwards conformably by the great series of red sandstones and
conglomerates known as the "Dingle Beds." Lithologically these
rocks present the closest resemblance to the Lower Old Red
Sandstone of Central Scotland. They occupy a similar stratigraphical
position, and though they have not yielded any palæontological data
for comparison, there can, I think, be no hesitation in classing them
with the Scottish Lower Old Red Sandstone, and regarding them as
having been deposited under similar geographical conditions. They
offer one feature of special interest for the purpose of the present
inquiry, since they contain a well-marked group of contemporaneous
volcanic rocks, including nodular felsites, like those so characteristic
of the Silurian period.
The area where this remote and isolated volcanic group is best
developed forms a range of high rugged ground along the northern
front of the hills that stretch eastward from the Lakes of Killarney.
Their general distribution is shown on Sheets 184 and 185 of the
Geological Survey of Ireland;[390] though I may again remark that
petrography has made great strides during the thirty years and more
that have passed since these maps and their accompanying Memoirs
were published, and that, were the district to be surveyed now,
probably a considerable tract of ground coloured as ash would be
marked as felsite. At the same time the existence of both these
rocks here cannot be gainsaid.
[390]See the Memoir (by J. B. Jukes and G. V. Du Noyer) on Sheet
184, p. 15. Other volcanic rocks have been mapped at Valentia
Harbour in the Dingle Beds, but these I have not had an opportunity
of personally examining.
The felsite was long ago brought into notice by Dr. Haughton, who
published an analysis of it.[391] It is also referred to by Mr. Teall for
its spherulitic structure.[392] Seen on the ground it appears as a pale
greenish-grey close-grained rock, sometimes exhibiting flow-
structure in a remarkably clear manner, the laminæ of devitrification
following each other in wavy lines, sometimes twisted and delicately
puckered or frilled, as in some schists. Portions of the rock are
strongly nodular, the nodules varying in size from less than a pea to
that of a hen's egg.
[391] Trans. Roy. Irish Acad. vol. xxiii. (1859), p. 615.
[392] British Petrography, p. 349.
The close resemblance of this rock to many of the Lower Silurian
nodular felsites of Wales cannot but strike the geologist. It presents
analogies also to the Upper Silurian felsites of Dingle. But its chief
interest arises from the geological horizon on which it occurs. Lying
in the so-called "Dingle-Beds," which may be regarded as the
equivalents of the Lower Old Red Sandstone of England and
Scotland, it is, so far as my observations go, the only example of
such a nodular felsite of later date than the Silurian period. We
recognize in it a survival, as it were, of the peculiar Silurian type of
acid lava, the last preceding eruption of which took place not many
miles to the west, in the Dingle promontory. But elsewhere this type
does not appear to have survived the end of the Silurian period.
The detrital rocks accompanying the felsite, in the district east of
Killarney, vary from such closed-grained felsitic material as cannot
readily be distinguished from the felsite itself to unmistakable felsitic
breccias. Even in the finest parts of them, occasional rounded
quartz-pebbles may be detected, while here and there a reddish
shaly band, or a layer of fine pebbly conglomerate with quartz-
pebbles an inch in length, shows at once the bedding and the dip.
Mr. W. W. Watts, who, with Mr. A. M'Henry of the Irish Staff of the
Geological Survey, accompanied me over this ground, found that a
microscopic examination of the slides which were prepared from the
specimens we collected completely confirmed the conclusions
reached from inspection of the rocks in the field.[393] He detected
among the angular grains slightly damaged crystals of felspar, chiefly
orthoclase. Many portions of these felspathic grits much resemble
the detrital Cambrian rocks which in the Vale of Llanberis have been
made out of the pale felsite of that locality.
[393]Mr. Watts also examined the microscopic structure of the felsite
of Benaun More. He found that the spherulites appear to have a
micropegmatitic structure, owing to the intergrowth of quartz and
felspar. In some parts of the rock the spherulites, from ·02 to ·01
inch in diameter, are surrounded by exceedingly minute green
needles, possibly of hornblende, while inside some of them are small
quartz-grains. Larger porphyritic felspars occur outside the
spherulites, some being of plagioclase, but most of orthoclase. The
spherulitic structure is not so well developed near the felspars. A few
of the large nodules are hollow and lined with crystals, while some of
them show a finely concentric lamination like the successive layers of
an agate.
 CHAPTER XXII
VOLCANOES OF THE UPPER OLD RED
SANDSTONE—THE SOUTH-WEST OF IRELAND,
THE NORTH OF SCOTLAND

In the northern half of Britain, where the Old Red Sandstone is so

well displayed, the two great divisions into which this series of
sedimentary deposits is there divisible are separated from each other
by a strongly marked unconformability. The interval of time
represented by this break must have been of long duration, for it
witnessed the effacement of the old water-basins, the folding,
fracture, and elevation of their thick sedimentary and volcanic
accumulations, and the removal by denudation of, in some places,
several thousand feet of these rocks. The Upper Old Red Sandstone,
consisting so largely as it does of red sandstones and
conglomerates, indicates the return or persistence of geographical
conditions not unlike those that marked the deposition of the lower
subdivision. But in one important respect its history differs greatly
from that which I have sketched for the older part of the system.
Though the Upper Old Red Sandstone is well developed across the
southern districts of Scotland from the Ochil to the Cheviot Hills, and
appears in scattered areas over so much of England and Wales, no
trace has ever been there detected in it of any contemporaneously
erupted volcanic rocks. The topographical changes which preceded
its deposition must have involved no inconsiderable amount of
subterranean disturbance, yet the volcanic energy, which had died
out so completely long before the close of the time of the Lower Old
Red Sandstone, does not appear to have been rekindled until the
beginning of the Carboniferous period.
Two widely separated tracts in the British Isles have yielded traces of
contemporaneous volcanic rocks in the Upper Old Red Sandstone.
One of these lies in the south-west of Ireland, the other in the far
north of Scotland.

THE SOUTH-WEST OF IRELAND

The Irish locality is situated a few miles to the south of the town of
Limerick, where the Carboniferous Limestone has been thrown into
long folds, and where, along the anticlines, strips of the underlying
red sandstones have been brought up to the surface. Two such
ridges of Upper Old Red Sandstone bear, each on its crest, a small
but interesting relic of volcanic activity[394] (Map I.).
[394] See Sheet 153 of the Geological Survey of Ireland, and
Explanation to that Sheet (1861), by Messrs. G. H. Kinahan and J.
O'Kelly. The account of the ground above given is from notes which I
made during a personal visit.
The more northerly ridge rises in the conical eminence of
Knockfeerina to a height of 949 feet above the sea. Even from a
distance the resemblance of this hill (Fig. 102) to many of the
Carboniferous necks of Scotland at once attracts the eye of the
geologist. The resemblance is found to hold still more closely when
the internal structure of the ground is examined. The cone consists
mainly of a coarse agglomerate, with blocks generally somewhat
rounded and varying in size up to two feet in length. The most
prominent of these, on the lower eastern slopes, are pieces of a fine
flinty felsite weathering white, but there also occur fragments of grit
and baked shale. The matrix is dull-green in colour, and among its
ingredients are abundant small lapilli of a finely vesicular andesite or
diabase. These more basic ingredients increase in number towards
the top of the eminence, where much of the agglomerate is almost
wholly made up of them. No marked dip is observable over most of
the hill, the rock appearing as a tumultuous agglomerate, though
here and there, particularly near the top and on the south side, a
rude bedding may be detected dipping outwards. On the west side
the agglomerate is flanked with yellow sandstone baked into
quartzite, so that the line of junction there between the two rocks
not improbably gives us the actual wall of the vent. The induration of
the surrounding sandstones is a familiar feature among the
Carboniferous vents. Some intrusive dark flinty rock traverses the
agglomerate near the top on the north side.

Fig. 102.—View of Knockfeerina, Limerick, from

the north-east—a volcanic neck of Upper Old
Red Sandstone age.
Retiring eastwards from the cone, the observer finds evidence of the
intercalation of tuff among the surrounding Upper Old Red
Sandstone. At the east end of the village of Knockfeerina a red
nodular tuff, with rounded pieces of andesite, grit and sandstone,
sometimes 12 inches long, is seen to dip under yellow, grey and red
sandstones and shales, while other shales and sandstones underlie
this tuff and crop out between it and the agglomerate. There is thus
evidence of the intercalation of volcanic tuff in the Upper Old Red
Sandstone of this district. And there seems no reason to doubt that
the tuff was ejected from the adjoining vent of Knockfeerina.
On the next ridge of Old Red Sandstone, which runs parallel to that
of Knockfeerina at a distance of little more than a mile to the south,
another mass of volcanic material rises into a prominence at
Ballinleeny. On the north side it consists of agglomerate like that just
described, and is flanked by sandstone baked into quartzite. Here
again we probably see the edge of a volcanic funnel. There may
possibly be more than one vent in this area. But well-bedded tuffs
can be observed to dip away from the centre and to pass under
sandstones and shales which are full of fine ashy material.
Gradations can be traced from the tuff into ordinary sediment. In
this instance, therefore, there is additional proof of
contemporaneous volcanic action in the Upper Old Red Sandstone.
There can be no uncertainty as to the horizon of the strata in which
these records have been preserved, for they dip conformably under
the shales and limestones at the base of the Carboniferous
Limestone series. They are said to have yielded the characteristic
fern Palæopteris of Kiltorcan.[395]
[395] There may be some other examples of Upper Old Red
Sandstone volcanic rocks in Ireland which I have not yet been able
personally to examine. On the maps of the Geological Survey (Sheet
198, and Explanation, pp. 8, 17) contemporaneous rocks of this age
are marked as occurring at Cod's Head and Dursey Island, on the
south side of the mouth of the Kenmare estuary.

THE NORTH OF SCOTLAND

The only district in England or Scotland wherein traces of volcanic
action during the time of the Upper Old Red Sandstone have been
observed lies far to the north among the Orkney Islands, near the
centre of the scattered outliers which I have united as parts of the
deposits of "Lake Orcadie"[396] (Map I.). The thick group of yellow
and red sandstones which form most of the high island of Hoy, and
which, there can be little doubt, are correctly referred to the Upper
Old Red Sandstone, rest with a marked unconformability on the
edges of the Caithness flagstones (Fig. 103). At the base of these
pale sandstones, and regularly interstratified with them, lies a band
of lavas and tuffs which can be traced from the base of the rounded
hills to the edge of the cliffs at the Cam, along the face of which it
runs as a conspicuous feature, gradually sloping to a lower level, till
it reaches the sea. At the Cam of Hoy it is about 200 feet thick, and
consists of three or more sheets of andesite. The upper 50 or 60
feet show a strongly slaggy structure, the central portion is rudely
columnar, and the lower part presents a kind of horizontal jointing or
bedding. There can be no question that this rock is not a sill but a
group of contemporaneous lava-flows. Beneath it, and lying across
the edges of the flagstones below, there is a zone of dull-red, fine-
grained tuff, banded with seams of hard red and yellow sandstone.
This tuff zone dies out to the eastward of the Cam.
[396]First noticed in Geol. Mag. February 1878; and Trans. Roy. Soc.
Edin. xxviii. (1878), p. 411.

Fig. 103.—Section of the volcanic Fig. 104.—Section of the volcanic

zone in the Upper Old Red zone in the Upper Old Red
Sandstone, Cam of Hoy, Orkney. Sandstone at Black Ness,
1. Caithness flagstones; 2. Dull- Rackwick, Hoy.
red tuff and bands of sandstone;
3. Lava zone in three bands; 4.
Yellow and red sandstone.

A few miles south of the Cam the volcanic zone appears again as the
platform on which the picturesque natural obelisk of the Old Man of
Hoy stands. Here the lava runs out as a promontory from the base
of the cliff, and on this projection the Old Man has been left isolated
from the main precipice. The cliffs of Hoy are traversed by numerous
small faults which have shifted the volcanic zone. But on the great
face of rock behind the Old Man there appears to be a second
volcanic zone lying several hundred feet above that just described. It
is probably this upper zone which emerges from under the hills a
mile and a half to the south at Black Ness in the bay of Rackwick. A
good section is there visible, which is represented in Fig. 104. The
ordinary red and yellow sandstones (a) appear from under the
volcanic rocks at this locality, and stretch southwards to the most
southerly headland of Hoy. The lowest volcanic band in the section is
one of red sandy well-bedded tuffs (b). Some of the layers are
coarse and almost agglomeratic, while others are fine marly and
sandy, with dispersed bombs, blocks and lapilli of diabase and
andesite. Hard ribs of a sandy tufaceous material also occur. These
fragmental deposits are immediately overlain by a dark-blue rudely
prismatic diabase with slaggy top (c). It is about 150 feet thick at its
thickest part, but rapidly thins away in a westerly direction. It passes
under a zone of red tuff (d) like that below, and above this highest
member of the volcanic group comes the great overlying pile of
yellow and reddish sandstone of Hoy. Followed westward for a short
distance, the whole volcanic zone is found to die out and the
sandstones below and above it then come together.
The interest of this little volcanic centre in Hoy is heightened by the
fact that the progress of denudation has revealed some of the vents
belonging to it. On the low ground to the east of the Cam, and
immediately to the north of the volcanic escarpment, the flagstones
which there emerge from under the base of the unconformable
upper sandstones are pierced by three volcanic necks which we may
with little hesitation recognize as marking the sites of vents from
which this series of lavas and tuffs was discharged (Fig. 105). The
largest of them forms a conspicuous hill about 450 feet high, the
smallest is only a few yards in diameter, and rises from the surface
of a flagstone ridge. They are filled with a coarse, dull-green,
volcanic agglomerate, made up of fragments of the lavas with pieces
of hardened yellow sandstone and flagstone. Around the chief vent
the flagstones through which it has been opened have been greatly
hardened and blistered. The most easterly vent, which has been laid
bare on the beach at Bring, due east of Hoy Hill, can be seen to
pierce the flagstones, which, although their general dip is westerly at
from 10° to 15°, yet at their junction with the agglomerate are bent
in towards the neck, and are otherwise much jumbled and disturbed.

Fig. 105.—Section across the volcanic band and its

associated necks, Hoy, Orkney.
1. Caithness flagstones; 2. Volcanic band lying on
red sandstones and conglomerates and dying out
eastwards; 3 3. Two vents between the base of
the hills and the sea; their connection with the
volcanic band is shown by dotted lines; 4.
Overlying mass of Upper Old Red Sandstone
forming the hills of Hoy.
On the northern coast of Caithness I have described a remarkable
volcanic vent about 300 feet in diameter, which rises through the
uppermost group of the Caithness flagstones. It is filled with a
coarse agglomerate consisting of a dull-greenish diabase paste
crowded with blocks of diabase, sometimes three feet in diameter,
and others of red sandstone, flagstone, limestone, gneiss and lumps
of black cleavable augite (Fig. 106).[397] The sandstones around it
present the usual disrupted, indurated and jointed character, and are
traversed by a small diabase dyke close to the western margin of the
neck. Another similar neck has since been found by the officers of
the Geological Survey on the same coast. That these volcanic orifices
were active about the same time with those in the opposite island of
Hoy may be legitimately inferred.
[397]See Trans. Roy. Soc. Edin. xxviii. (1878), p. 405; also p. 482 of
the same volume for an account of the cleavable augite.
 Fig. 106.—Ground-plan of volcanic neck
piercing the Caithness Flagstone series
on the beach near John o' Groat's
House.
These northern volcanoes made their appearance in a district where
during the preceding Lower Old Red Sandstone period there had
been several widely separated groups of active volcanic vents. So far
as the fragmentary nature of the geological evidence permits an
opinion to be formed, they seem to have broken out at the
beginning, or at least at an early stage, of the deposition of the
Upper Old Red Sandstone, and to have become entirely extinct after
the lavas of Hoy were poured forth. No higher platform of volcanic
materials has been met with in that region. With these brief and
limited Orcadian explosions the long record of Old Red Sandstone
volcanic activity in the area of the British Isles comes to an end.[398]
[398] There appear to be traces of volcanic eruptions
contemporaneous with the Upper Old Red Sandstone of Berwickshire,
but as they merely formed a prelude to the great volcanic activity of
Carboniferous time, they are included in the account of the
Carboniferous plateau of Berwickshire in Chapters xxiv. and xxv.
 BOOK VI
THE CARBONIFEROUS VOLCANOES
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