Basic Neurochemistry - 8th Edition - ISBN: 9780123749475, 9780080959016

Basic Neurochemistry

8th Edition

Principles of Molecular, Cellular, and Medical Neurobiology

Editor-in-Chiefs: Scott Brady
Editors: Scott Brady George Siegel R. Wayne Albers Donald Price
eBook ISBN: 9780080959016
Hardcover ISBN: 9780123749475
Imprint: Academic Press
Published Date: 7th December 2011
Page Count: 1120
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Basic Neurochemistry, Eighth Edition, is the updated version of the outstanding and comprehensive classic text on neurochemistry. For more than forty years, this text has been the worldwide standard for information on the biochemistry of the nervous system, serving as a resource for postgraduate trainees and teachers in neurology, psychiatry, and basic neuroscience, as well as for medical, graduate, and postgraduate students and instructors in the neurosciences.

The text has evolved, as intended, with the science. This new edition continues to cover the basics of neurochemistry as in the earlier editions, along with expanded and additional coverage of new research from intracellular trafficking, stem cells, adult neurogenesis, regeneration, and lipid messengers. It contains expanded coverage of all major neurodegenerative and psychiatric disorders, including the neurochemistry of addiction, pain, and hearing and balance; the neurobiology of learning and memory; sleep; myelin structure, development, and disease; autism; and neuroimmunology.

Key Features

  • Completely updated text with new authors and material, and many entirely new chapters
  • Over 400 fully revised figures in splendid color
  • 61 chapters covering the range of cellular, molecular and medical neuroscience
  • Translational science boxes emphasizing the connections between basic and clinical neuroscience
  • Companion website at


Advanced undergraduates, graduate students and post-docs in neuroscience or biomedical science, medical students, clinical neuroscientists and neurologists

Table of Contents

Associate Editors

List of Boxes


List of Contributors

Eighth Edition Acknowledgments and History

Preface to the Eighth Edition

Part I: Cellular Neurochemistry and Neural Membranes

Chapter 1. Cell Biology of the Nervous System


Cellular Neuroscience is the Foundation of Modern Neuroscience

Neurons: Common Elements and Diversity

Macroglia: More than Meets the Eye


Blood–Brain Barriers and the Nervous System

The Neurovascular Unit Includes Multiple Components



Chapter 2. Cell Membrane Structures and Functions

Phospholipid Bilayers

Membrane Proteins

Biological Membranes


Chapter 3. Membrane Transport


Primary Active Transport (P-Type) Pumps

Na,K-Adenosinetriphosphatase (Na,K-ATPase)

Ca Adenosinetriphosphatases and Na,Ca Antiporters

The Primary Plasma Membrane Ca Transporter (PMCA)

Smooth Endoplasmic Reticulum Calcium Pumps (SERCA)

Other P-Type Transporters

V0V1 Proton Pumps

ATP-Binding Cassettes

Secondary Active Transport

General Physiology of Neurotransmitter Uptake and Storage

The Cation Antiporters

The Anion Antiporters

Facilitated Diffusion: Aquaporins and Diffusion of Water

Facilitated Diffusion of Glucose and Myoinositol


Chapter 4. Electrical Excitability and Ion Channels

Membrane Potentials and Electrical Signals in Excitable Cells

Action Potentials in Electrically Excitable Cells

Functional Properties of Voltage-Gated Ion Channels

The Voltage-Gated Ion Channel Superfamily

The Molecular Basis for Ion Channel Function

Ion Channel Diversity



Chapter 5. Lipids


Properties of Brain Lipids

Complex Lipids

Analysis of Brain Lipids

Brain Lipid Biosynthesis

Genes for Enzymes Catalyzing Synthesis and Degradation of Lipids

Lipids in the Cellular Milieu




Chapter 6. The Cytoskeleton of Neurons and Glia


Molecular Components of the Neuronal Cytoskeleton

Ultrastructure and Molecular Organization of Neurons and Glia

Cytoskeletal Structures in the Neuron Have Complementary Distributions and Functions



Chapter 7. Intracellular Trafficking


General Mechanisms of Intracellular Membrane Trafficking in Mammalian Cells Include Both Universal and Highly Specialized Processes

Fundamentals of Membrane Trafficking are Based on a set of Common Principles

The biosynthetic Secretory Pathway Includes Synthetic, Processing, Targeting and Secretory Steps

The Endocytic Pathway Plays Multiple Roles in Cells of the Nervous System

Synaptic Vesicle Trafficking is a Specialized Form of Regulated Secretion and Recycling Optimized for Speed and Efficiency



Chapter 8. Axonal Transport


Neuronal Organelles in Motion

Discovery and Development of the Concept of Fast and Slow Components of Axonal Transport

Fast Axonal Transport

Slow Axonal Transport

Molecular Motors: Kinesin, Dynein and Myosin

AXONAL Transport and Neuropathology



Chapter 9. Cell Adhesion Molecules


Immunoglobulin Superfamily



Cooperation and Crosstalk between Cell Adhesion Molecules



Chapter 10. Myelin Structure and Biochemistry

The Myelin Sheath

Characteristic Composition of Myelin



Chapter 11. Energy Metabolism of the Brain


Substrates for Cerebral Energy Metabolism

Age and Development Influence Cerebral Energy Metabolism

Fueling Brain: Supply–Demand Relationships and Cerebral Metabolic Rate

Metabolism in the Brain is Highly Compartmentalized

Glycolysis: Conversion of Glucose to Pyruvate

Glycogen is Actively Synthesized and Degraded in Astrocytes

The Pentose Phosphate Shunt has Essential Roles in Brain

The Malate–Aspartate Shuttle has a key Role in Brain Metabolism

There is Active Metabolism of Lactate in Brain

Major Functions of the Tricarboxylic Acid (TCA) Cycle: Pyruvate Oxidation to CO2, NADH/FADH2 Formation for ATP Generation and Synthesis of Glutamate and Aspartate

Mitochondrial Heterogeneity: Differential Distribution of Many TCA Cycle Enzymes and Components of Oxidative Phosphorylation in Neuronal and Glial Mitochondria

Glutamate–Glutamine Metabolism is Linked to Energy Metabolism

Metabolic Studies in Brain: Imaging and Spectroscopy

Relation of Energy Metabolism to Pathological Conditions in the Brain



Part II: Intercellular Signaling

Chapter 12. Synaptic Transmission and Cellular Signaling: An Overview

Synaptic Transmission

Cellular Signaling Mechanisms



Chapter 13. Acetylcholine


Synthesis, Storage and Release of Acetylcholine: Distribution of Cholinergic Pathways

Enzymatic Breakdown of Acetylcholine

Nicotinic Cholinergic Receptors

Muscarinic Cholinergic Receptors


Chapter 14. Catecholamines

Overview of Catecholamines


Adrenergic Receptors

Agonist-Induced Downregulation

Repeated Antagonist Treatment


Chapter 15. Serotonin

Serotonin, The Neurotransmitter

Serotonin Receptors


Chapter 16. Histamine


Histamine: The Molecule and the Messenger

Histaminergic Cells of the Central Nervous System: Anatomy and Morphology

Dynamics of Histamine in the Brain

Molecular Sites of Histamine Action

Histamine Actions on the Nervous System

Significance of Brain Histamine for Drug Action


Chapter 17. Glutamate and Glutamate Receptors

The Amino Acid Glutamate is the Major Excitatory Neurotransmitter in the Brain

Brain Glutamate is Derived from Blood-Borne Glucose and Amino Acids that Cross the Blood–Brain Barrier

Glutamine is an Important Immediate Precursor for Glutamate: The Glutamine Cycle

Synaptic Vesicles Accumulate Transmitter Glutamate by Vesicular Glutamate Transporters

Is Aspartate a Neurotransmitter?

Long-Term Potentiation or Depression of Glutamatergic Synapses May Underlie Learning

The Neuronal Pathways of the Hippocampus are Essential Structures for Memory Formation

Ionotropic and Metabotropic Glutamate Receptors are Principal Proteins at the Postsynaptic Density

Three Classes of Ionotropic Glutamate Receptors are Identified

Glutamate Produces Excitatory Postsynaptic Potentials

Metabotropic Glutamate Receptors Modulate Synaptic Transmission

Glutamate Receptors Differ in their Postsynaptic Distribution

Proteins of the Postsynaptic Density Mediate Intracellular Effects of Glutamate Receptor Activation

Dendritic Spines are Motile, Changing their Shape and Size in Response to Synaptic Activity within Minutes

Sodium-Dependent Symporters in the Plasma Membranes Clear Glutamate from the Extracellular Space

Sodium-Dependent Glutamine Transporters in Plasma Membranes Mediate the Transfer of Glutamine from Astrocytes to Neurons

Excessive Glutamate Receptor Activation may Mediate Certain Neurological Disorders


Chapter 18. GABA


GABA Synthesis, Release and Uptake

GABA Receptor Physiology and Pharmacology

Structure and Function of GABA Receptors

GABA is the Major Rapidly Acting Inhibitory Neurotransmitter in Brain


Chapter 19. Purinergic Signaling

Nomenclature of Purines and Pyrimidines

Purine Release

Purinergic Receptors

Effects of Purines in the Nervous System

Disorders of the Nervous System—Purines and Pain: A1R, P2X and P2Y Receptors

Disorders of the Nervous System: Adenosine Kinase and the Adenosine Hypothesis of Epilepsy

Disorders of the Nervous System: Parkinson’s Disease and A2A Antagonists

Concluding Comments


Chapter 20. Peptides


Neuropeptide Receptors

Neuropeptide Functions and Regulation

Peptidergic Systems in Disease


Part III: Intracellular Signaling

Chapter 21. G Proteins

Heterotrimeric G Proteins

Small G Proteins

Other Features of G Proteins


Chapter 22. Cyclic Nucleotides in the Nervous System

Introduction: Second Messengers

Adenylyl Cylcases

Guanylyl Cyclases


Spatiotemporal Integration and Regulation of Cyclic Nucleotide Signaling in Neurons

Conclusion and Future Perspective


Chapter 23. Phosphoinositides


The Inositol Lipids

The Inositol Phosphates


Phosphoinositides and Cell Regulation


Chapter 24. Calcium

The Calcium Signal in Context

Calcium Measurement

Calcium Homeostasis at the Plasma Membrane

Cellular Organelles and Calcium Pools

Ca2+ Signaling Begins in Microdomains

Local and Global Ca2+ Signaling: Integrative Roles for Astrocytes?



Chapter 25. Serine and Threonine Phosphorylation

Protein Phosphorylation is a Fundamental Mechanism Regulating Cellular Functions

Protein Ser/Thr Kinases

Protein Ser/Thr Phosphatases

Common strategies used for the evaluation of neuronal functions of protein kinases and phosphatases

Neuronal Phosphoproteins

Protein Phosphorylation is a Fundamental Mechanism Underlying Synaptic Plasticity and Memory Functions

Protein Phosphorylation in Human Neuronal Disorders



Chapter 26. Tyrosine Phosphorylation

Tyrosine Phosphorylation in the Nervous System

Protein Tyrosine Kinases

Protein Tyrosine Phosphatases

Role of Tyrosine Phosphorylation in the Nervous System


Chapter 27. Transcription Factors in the Central Nervous System

The Transcriptional Process

Regulation of Transcription by Transcription Factors

Glucocorticoid and Mineralocorticoid Receptors as Transcription Factors

cAMP Regulation of Transcription

The Role of Transcription Factors in Cellular Phenotype

The Transcriptome Dictates Cellular Phenotype

Transcription as a Target for Drug Development


Part IV: Growth, Development and Differentiation

Chapter 28. Development of the Nervous System


Early Embryology of the Nervous System

Spatial Regionalization

Neurogenesis and Gliogenesis

PNS Development and Target Interactions

Axon Guidance Contributes to Correct Connections

Synapse Formation

Activity and Experience Shape Long-Lasting Connections



Chapter 29. Growth Factors

Introduction: What is a Growth Factor?


Regulation of Neurotrophin Expression


Neurotrophin Receptors

Glial Cell line–Derived Neurotrophic Factor (GDNF)

GFL Receptors


Neurotrophic Cytokines

Summary and Conclusions


Chapter 30. Stem Cells in the Nervous System


Stem Cells are Multipotent and Self-Renewing

Neural Stem Cells Contribute to Neurons and Glia During Normal Development

Stem Cells can be Identified Antigenically and Functionally

Stem Cells Offer Potential for Repair in the Adult Nervous System


Chapter 31. Formation and Maintenance of Myelin


Schwann Cell Development

Oligodendrocyte Development

Regulation of Myelination

Developmental and Metabolic Aspects of Myelin

Genetic Disorders of Myelination

Myelin Maintenance




Chapter 32. Axonal Growth in the Adult Mammalian Nervous System: Regeneration and Compensatory Plasticity


Regeneration in the Peripheral Nervous System

Regeneration in the Central Nervous System

Central Nervous System Injury and Compensatory Plasticity




Part V: Cell Injury and Inflammation

Chapter 33. Molecular Mechanisms and Consequences of Immune and Nervous System Interactions


Distinguishing Friend from FOE

The Nervous System Regulates Both Innate and Adaptive Immunity

Immune Privilege Is Not Immune Isolation: The CNS as an Immune-Active Organ

Immune-Regulated Changes in Neuronal Function and Mammalian Behavior

Summary: Manipulating Neuroimmune Interactions


Chapter 34. Neuroinflammation

Neuroinflammation: Introduction

The Highly Regulated Activation of Microglia and Phagocytosis

Microglial Dysfunction During Aging

Protein Aggregation


Lipid Mediator Pathways in Neuroinflammation

Ischemia-Reperfusion Damage

The Interface Between Inflammation and the Immune System in the CNS

Mitochondria: A Connection Between Inflammation and Neurodegeneration

Neuroprotective Signaling Circuits


Chapter 35. Brain Ischemia and Reperfusion: Cellular and Molecular Mechanisms in Stroke Injury

Brain Responses to Ischemia

Injury in the Ischemic Phase

Brain Injury During the Reperfusion Phase: Free Radicals in Ischemia–Reperfusion Injury

Breakdown of the Neurovascular Unit and Brain Edema

Neuroprotection Signaling and Resolution of Inflammation: Mechanisms

Potential Therapeutic Strategies for Acute Ischemic Stroke



Chapter 36. Lipid Mediators: Eicosanoids, Docosanoids and Platelet-Activating Factor

Storage of Lipid Messengers in Neural Membrane Phospholipids

Phospholipases A2


Platelet-Activating Factor



Diacylglycerol Kinases

Lipid Signaling in Neuroinflammation

Docosahexaenoic Acid

Lipid Peroxidation and Oxidative Stress


Neuroprotectin D1: A Docosahexaenoic-Acid–Derived Mediator

The Future of Neurolipidomic Signaling


Chapter 37. Apoptosis and Necrosis

Distinguishing Features of Apoptosis and Necrosis



Targeting Apoptosis and Necrosis in Neurological Disorders


Part VI: Inherited and Neurodegenerative Diseases

Chapter 38. Peripheral Neuropathy: Neurochemical and Molecular Mechanisms


Peripheral Nerve Organization

Genetically Determined Neuropathies

Diabetic Neuropathy

Autoimmune Neuropathies

Other Causes of Peripheral Nerve Disorders

Axon Degeneration and Protection


Chapter 39. Diseases Involving Myelin

General Classification

Acquired Immune-Mediated and/or Infectious Diseases of Myelin

Genetically Determined Disorders of Myelin

Other Diseases Primarily Involving Myelin

Disorders Primarily Affecting Neurons with Secondary Involvement of Myelin

Repair in Demyelinating Diseases



Chapter 40. The Epilepsies: Phenotypes and Mechanisms

Epilepsy is a Common Neurological Disorder

Terminology and Classification

Mechanisms of Antiseizure Drugs

Genetics of Epilepsy


Chapter 41. Genetics of Neurodegenerative Diseases

Genetic Aspects of Common Neurodegenerative Diseases

Alzheimer’s Disease

Parkinson’s Disease

Dementia with Lewy Bodies

Frontotemporal Dementia

Amyotrophic Lateral Sclerosis

Neurodegenerative Triplet Repeat Disorders

Creutzfeld-JaKob Disease and other Prion Diseases

Concluding Remarks


Chapter 42. Disorders of Amino Acid Metabolism


Disorders of Branched-Chain Amino Acids: Maple Syrup Urine Disease

Disorders of Phenylalanine Metabolism: Phenylketonuria

Disorders of Glycine Metabolism: Nonketotic Hyperglycinemia

Disorders of Sulfur Amino Acid Metabolism: Homocystinuria

The Urea Cycle Defects

Disorders of Glutathione Metabolism

Disorders of -Aminobutyric Acid Metabolism

Disorders of N-Acetyl Aspartate Metabolism


Chapter 43. Inborn Metabolic Defects of Lysosomes, Peroxisomes, Carbohydrates, Fatty Acids and Mitochondria

Lysosomal Storage Diseases

Peroxisomal Diseases

Classification of Peroxisomal Diseases

Therapy of Peroxisomal Diseases

Diseases of Carbohydrate and Fatty Acid Metabolism

Diseases of Mitochondrial Metabolism

Acknowledgments and Dedication


Chapter 44. Disorders of Muscle Excitability

Organization of the Neuromuscular Junction

Excitation and Contraction of the Muscle Fiber

Genetic Disorders of the Neuromuscular Junction

Hereditary Diseases of Muscle Membranes

Immune Diseases of Muscle Excitability

Toxins and Metabolites that Alter Muscular Excitation


Chapter 45. Motor Neuron Diseases

Amyotrophic Lateral Sclerosis Is the Most Common Adult-Onset Motor Neuron Disease

Models of Motor Neuron Disease Induced by Experimental Nerve Injury Have been Instructive

Selected Genetic Models of Relevance to ALS and Other Motor Neuron Diseases Have been Identified or Generated

Available Genetic Mouse Models Will Aid in Discovering Disease Mechanisms and Novel Means of Therapy



Chapter 46. Neurobiology of Alzheimer’s Disease

Alzheimer’s Disease is the Most Prevalent Neurodegenerative Disease of the Elderly




Chapter 47. Synucleinopathies and Tauopathies



Parkinson’s Disease and Other Lewy Body Diseases

Multiple System Atrophy

Synthetic α-Synuclein Filaments

Animal Models of Synucleinopathies


Microtubule-Associated Protein Tau

Tau and Alzheimer’s Disease

Other Tauopathies

MAPT Mutations Causing Tauopathy

Relevance for Other Tauopathies

Synthetic Tau Filaments

Animal Models of Human Tauopathies



Chapter 48. Cellular and Molecular Basis of Neurodegeneration in the CAG–Polyglutamine Repeat Diseases

Introduction to the CAG–Polyglutamine Repeat Diseases

Expanded Polyglutamine Tracts Promote Protein Misfolding to Drive Neurotoxicity

The Role of Protein turnover Pathways in Polyglutamine Disease Pathogenesis

The Importance of Normal Function in the Polyglutamine Repeat Diseases

RNA Toxicity in the Polyglutamine Repeat Diseases?

Gene Silencing is a Promising Therapy for Polyglutamine Repeat Disease


Chapter 49. Neurotransmitters and Disorders of the Basal Ganglia

Anatomy and Physiology of the Basal Ganglia

Disorders that Involve Basal Ganglia Dysfunction



Chapter 50. Molecular Basis of Prion Diseases


Prion Diseases are Biologically Unique

Animal Prion Diseases

Human Prion Diseases

Prion Disease Pathology and Pathogenesis

The Protein-Only Hypothesis of Prion Propagation

Characterization of PrPC

Characterization of PrPSc

The Molecular Basis of Prion Strain Diversity

Prion Transmission Barriers

Future Perspectives


Part VII: Sensory Transduction

Chapter 51. Molecular Biology of Vision

Structure and Development of the Visual System

Photoreceptors and Phototransduction

Signaling Downstream of Photoreceptors

Recycling of Phototransduction Molecules

Retinal Neurodegeneration


Chapter 52. Molecular Basis of Olfaction and Taste





Chapter 53. Molecular Biology of Hearing and Balance

General Features of Mechanotransduction

Non-Vertebrate Model Systems

Hair Cells

Hair Cells in the Inner Ear

Balance: Vestibular Organs

Hearing: Cochlea



Chapter 54. Pain

Nociceptive Versus Clinical Pain

Nociceptors are First Responders

Pain Transmission in the Spinal Cord

Brainstem, Thalamus and Cortex

Opioid Analgesia


Inflammatory Pain

Neuropathic Pain

Genetic Factors




Part VIII: Neural Processing and Behavior

Chapter 55. Endocrine Effects on the Brain and Their Relationship to Behavior


Behavioral Control of Hormonal Secretion

Classification of Hormonal Effects

Biochemistry of Steroid and Thyroid Hormone Actions

Intracellular Steroid Receptors: Properties and Topography

Membrane Steroid Receptors and Signaling Pathways

Biochemistry of Thyroid Hormone Actions on Brain

Diversity of Steroid-Hormone Actions on the Brain



Chapter 56. Learning and Memory

Brief History of Memory Research in Humans

Divisions of Memory

Molecular Mechanisms of Learning

Molecular Mechanisms of Memory Consolidation and Storage

Neural Population-Level Memory Traces and Their Organizing Principles


Chapter 57. The Neurochemistry of Sleep and Wakefulness

Sleep Phenomenology and Function: The Search for Neurochemical Substrates

Development of Sleep Disorders Medicine and Sleep Neurobiology





Amino Acids


Conclusions and Future Directions


Chapter 58. The Neurochemistry of Schizophrenia

Clinical Aspects of Schizophrenia

Brain Imaging

Cellular and Molecular Studies



Chapter 59. The Neurochemistry of Autism

Clinical Aspects of Autism Spectrum Disorders (ASDs)

Genetic Studies

Neurochemical Studies



Chapter 60. Neurobiology of Severe Mood and Anxiety Disorders

Mood Disorders

Neurotransmitter and Neuropeptide Systems and the Pathophysiology of Mood Disorders

Neuroanatomical and Neuropathological Correlates of Mood Disorders

Intracellular Signaling Pathways

Anxiety Disorders

The Neurochemistry of Fear and Anxiety

Intracellular Targets for Anxiety Disorders

Future Directions and the Development of Novel Therapeutics


Chapter 61. Addiction

General Principles

Neuronal Circuitry of Addiction


Psychomotor Stimulants

Cannabinoids (Marijuana)


Ethanol, Sedatives and Anxiolytics

Hallucinogens and Dissociative Drugs

Addiction and Neuronal Plasticity Share Common Cellular Mechanisms






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About the Editor-in-Chief

Scott Brady

Affiliations and Expertise

Head, Department of Anatomy and Cell Biology, University of Illinois, Chicago, Illinois, USAUniversity of Illinois, Chicago IL

About the Editor

Scott Brady

Affiliations and Expertise

Head, Department of Anatomy and Cell Biology, University of Illinois, Chicago, Illinois, USAUniversity of Illinois, Chicago IL

George Siegel

Affiliations and Expertise

Emeritus Professor of Neruology, Loyola University Chicago Stritch School of Medicine; Chief of Neurology Service (retired), Edward Hines, Jr. Veterans Affairs Hospital, Chicago, IL, USA

R. Wayne Albers

R. Wayne Albers

OBITUARY FOR R. WAYNE ALBERS, August 5, 1928 - September 28, 2013

R. Wayne Albers, Ph.D., Scientist Emeritus, Chief of Section on Enzyme Chemistry (retired), Laboratory of Neurochemistry in the NINDS, NIH, Bethesda, MD, is a world-recognized neuroscientist most noted for his research in the field of membrane cation transport and neuronal excitability in the nervous system. Dr. Albers and physiologist R.L. Post performed the principal experiments leading to their now widely-held Albers-Post model for the mechanism of the cation transport enzyme, sodium-potassium-activated ATPase. Dr. Albers was one of the founding co-editors of the comprehensive text, Basic Neurochemistry: Molecular, Cellular and Medical Aspects, first published in 1972, continuing as co-editor for 8 editions, the latest having been published in 2012. After receiving his PhD at Washington University School of Medicine in 1954, Dr. Albers embarked on a distinguished career of research at the NIH, being a founding investigator in the first Laboratory of Neurochemistry.

Dr. Albers was one of the first members of the American Society for Neurochemistry at its inception, serving on its Council and its Committees on Publications and Education and on Electronic Publications. He has served as Professor of Biochemistry at George Washington University, Faculty Member of the NIH Foundation for Advanced Education in the Sciences, Associate Editor of the Journal of Neurochemistry and of Experimental Neurology, and on the editorial boards of several journals.

Dr. Albers passed away on September 28, 2013. He was 85 years old and is survived by his former wife, Frances Albers, their children Gail Morrell, Belinda Caron and Patricia Steinhoff, 6 grandchildren, and 8 great-grandchildren. He also had a son, the late Jonathan Albers.

Dr. Albers was considered a gentleman, an excellent scientific colleague with a keen intellect and friend by all who worked with him. He will be sorely missed, not only by his family, but also by the entire neurochemistry community.

George J. Siegel

October 1, 2013

Affiliations and Expertise

National Institutes of Health, Bethesda MD, USA

Donald Price

Affiliations and Expertise

The Johns Hopkins University School of Medicine, Baltimore MD

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