Core Ideas in Neuroscience

Core Ideas in Neuroscience

William R. Klemm, DVM, PhD

Professor of Neuroscience

Texas A&M University

College Station, Texas 77843-4458

Second Edition

Published by W. R. Klemm at Smashwords

Copyright 2013, 2016, W. R. Klemm

All rights reserved. No part of this book may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopying, recording, or by any information storage and retrieval system, without written permission from the author, except for the inclusion of brief quotations in a review.

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Table of Contents

Introduction

Overview

Cell Biology

Development/Trophism

Senses

Information Processing

States of Consciousness

Emotions

Learning and Memory

Motor Output and Control

Glossary

Afterword

About the Author

Introduction

Science is not a collection of facts or of unquestionable generalizations, but a logically connected network of hypotheses that represent our current opinion about what the real world is like.

P. B. Medawar

In other words, Medawar is saying that our understanding of science should be driven by provisional theory and core ideas. Details either support or refute those ideas, showing us how reliable our ideas are. Core ideas drive new research and discovery. Therefore, I think it is the first duty of teachers and learners alike to make certain they identify and understand the core ideas of their discipline. This book is dedicated to that end.

Rationale for a Core-Ideas Book

What the brain produces is a kind of mental model of the world, a system for handling the information that flows from sense organs to the generation of appropriate responses. The integration of the sensory data is central to monitoring the world out there and to creating a model of it in here. The in here becomes the real world as far as animals and people experience it. To explain what we know about how all this occurs in the brain is no trivial task.

Too many books, in my opinion, fall short of providing the big picture of how nervous systems work. A typical neuroscience text is so heavily laden with factual and highly technical detail that the essence of understanding can be obscured. Students are easily confused over what they must know as opposed to what is nice to know.

Even when nice to know is deemed must know, there are over-arching ideas that ought to be mastered first. What a student-oriented text should be is one that focuses on central concepts and principles: the BIG ideas! If you get the big picture first, then it is easier to integrate the details into that conceptual framework. Learning the other way around is harder to do.

The standard science textbooks have gotten progressively larger, serving better a role as reference books than as textbooks. This trend is especially pervasive in such active research areas as neuroscience, where students and teachers alike are swamped with new information.

How bad have things gotten? I can best illustrate the problem by the annual meeting for the Society of Neuroscience. For years now, the attendance has been running around 30,000 scientists and 15,000 research report presentations. Who can digest all that? No professor can keep up. How can students do so? Probably no other field of science is in such dire need for condensation and integration into core ideas.

This learning problem is especially acute for professionals who are not neuroscientists, but whose work needs to be informed by neuroscience. These professionals include physicians, osteopaths, veterinarians, dentists, clinical and experimental psychologists, computer scientists, bioengineers, animal behaviorists, biologists, nurses, and allied health workers. For them, a book like this one can serve as their primary source of neuroscience understanding.

Certainly, the typical neuroscience textbook does a poor job of condensing the oceans of neuroscience information into drinkable amounts. Even one of the early, well-known neuroscience principles book, Elements of Neurophysiology (by Ochs, 1965, Wiley & Sons), took 621 pages to specify the Elements (principles). (This Core Ideas book is about 330 printed pages). At the time of the writing of the first edition of this Core Ideas book, the most popular book, Principles of Neural Science (4rd Ed.), was that by Kandel et al., 2000, Elsevier. This book is 1,414 pages long and has 44 authors. A competing book, Fundamental Neuroscience, by Zigmond et al. 1999, is 1600 pages long, has 15 editors and 150 authors. Clearly, such books tell most readers far more than they want or need to know.

The practical value of texts should lie in their pedagogical approach, which is the opposite from the tack taken in many textbooks on biological and medical subjects. Students don’t need encyclopedic texts to learn, though they can certainly find them useful as references. Students and professors alike are tired of an educational process devoted to pouring information into one ear while it spills out the other. The exponential expansion of new knowledge is causing cognitive overload in both students and professors, short-circuiting their ability to sustain perspective about the whole of biomedical science and to think coherently about the details of how the body works in both health and disease. We are learning more and more about less and less, and that causes a progressive loss of capacity for synthesis and ability to think about the larger meanings of biomedical science.

In recognition of the cognitive overload problem, several medical schools (McMasters, Harvard, Southern Illinois, Bowman Gray, U. New Mexico) pioneered in the now popular teaching approach of converting the traditional lecture-based curriculum to a tutorial, group-based learning format where critical thinking and information management skills are emphasized rather than rote memory. Some veterinary colleges are also making similar curricular changes. This trend will surely grow, because it is aimed at teaching students to manage and integrate an ever-expanding biomedical data base.

The situation argues for a new kind of textbook, one that is focused on the first-principles of the discipline. In addition to encyclopedic tomes for each discipline, we need small supplementary texts that give the big picture and explicitly describe the basic foundational ideas of the discipline: and no more! The time it takes to learn is important for students (if they work part time, for example, time is money). An inexpensive e-book like this will speed the time it takes to grasp instruction from a traditional, detail-oriented textbook.

Defining a Core Idea

We must have some kind of working definition of core idea or principle. While many ways can express the idea, this text uses the following working definition:

A principle goes beyond a collection of observations. Principles integrate multiple observations and help to explain these observations, providing understanding and insight. A principle embodies the underlying rules or mechanisms of structure, organization, or operation that give rise to the observations. We distinguish principles from concepts only in the sense that concepts often embody more than one principle.

In identifying these core ideas, we must recognize that they are commonly held beliefs about what is true and fundamental. Not everyone will agree that each of the more than 75 core ideas in this book deserves such lofty status. Never mind. We should not quibble over what ideas deserve to be included and what ought to be thrown out or merged. This can only lead to endless debate and interfere with what is a practical way to teach and learn.

Many of the statements of principle are incomplete. They may also lack sufficient qualification. Statements of principles often serve to inspire debate and more complete or precise exposition. The effort to identify principles comes at a cost: arbitrariness, uncertainty, and controversy. But the value is worth the price.

Rationale for An E-book

This book should be considered as a complement to a traditional text in neuroscience. When students have to pay hundreds of dollars for traditional textbooks, it is unrealistic to expect them to buy a supplementary text. Several leading neuroscience books cost over $150 each. E-books are inexpensive and at a price low enough that most students should be willing to pay. The e-book also has the advantage of hyperlinking, making it easy for students to find quickly what they seek. While many people prefer reading hard copy rather than a computer screen, it needs to be remembered that this book has a modular design and it only takes a few minutes to read (or print) a given module.

The modular construction of topics allows the reader to pick and choose to read a minimal amount of material to get prepared for understanding the more comprehensive and detailed traditional textbooks and research papers. Maybe even highly specialized expert neuroscientists may find some useful perspectives from this approach to explaining the subject of neuroscience from a holistic perspective.

Organization of the Book

Most of what everybody needs to know about the nervous system can be summarized in this list of principles or core ideas. Each core idea is treated as a distinct module, although there are links indicating other closely related ideas. Each module has the same basic format:

Core idea is stated succinctly.

Terms used in the module are defined.

Explanation of the idea/concept.

Examples that illustrate the idea/concept.

Related ideas found elsewhere in this book.

References (including so-called Citation Classics).

The Classic papers are highly cited publications that scientists generally agree have helped to define the field. A Classic is a highly cited publication that was identified by the Science Citation Index, published by ISI Press, in Philadelphia. For some of the classics, there is an associated publication that appeared in the ISI publication, Current Contents, that contains a history of the research that enabled the publication to have such a major impact. In some cases, I have taken the liberty of including some references in the Classic section that have not been officially annointed by ISI’s data. Sometimes, it is because these were landmark papers that were published before ISI began keeping citation records. In a few other cases, the choice is strictly my opinion.

Unfortunately, even these classics seem to have a short life span. One of the things I have noticed over the years as neuroscience information has accumulated is that there is much need for a sense of history of the discipline. Young scientists, who tend to be the most active in research, tend to work in the currently hot areas. Ideas and data over a few years old are often ignored. As a result, I see more and more research reports that have either reinvented the wheel, or that would have been greatly enriched had the authors been aware of older literature.

Most of the ideas in the book are presented in the form of the reductionistic research tradition in which the nervous system is understood in terms of molecules, single-cell physiology, anatomical pathways, and input-output relationships. Despite all the powerful insights that have accumulated, mostly in the last 10 years or so, we know that reductionistic research has it limits and cannot answer the most important questions about how the brain assigns meaning to life experience, generates intentionality, and makes choices and decisions. It is now increasingly clear that the more profound thinking actions, such as memory, emotions, and consciousness, derive from interactions of huge populations of neurons. Dynamic interactions of large neuronal ensembles are best revealed by monitoring concurrent activity in multiple neuronal populations, using such tools as functional MRI for metabolic indicators and quantitative electroencephalography for indications of extracellular voltage fluctuations at various frequencies and degrees of coherence. In short, brain functions are system functions, not functions of individual molecules or cells.

Core ideas are grouped into categories to make it easier to organize and remember them. The categories are:

Overview

Cell Biology

Senses

Information Processing

States of Consciousness

Emotions

Learning and Memory

Motor Output

Control Development/Trophism

The Overview section contains certain ideas that transcend more than one of the other categories. Each of the book’s sections begins with a short introduction that presents an overview of the big ideas in that category. There is a concept map that helps to display graphically the relationships among the various ideas. Then there follows a succession of modules, each of which explains one of the ideas in that category. Each module has a name, reflective of the idea.

Suggested Uses by Teachers

This book is not meant to be read front to back like a textbook. Its modular construction allows the teacher to pick and choose what topics need to be covered at any particular time. This design makes the e-book useful for traditional neuroscience courses or for other kinds of courses where only a portion of the neuroscience discipline needs to be taught to students. The idea is to use these core ideas as an introduction that should precede what is planned for a given day’s class. Teachers know that students typically won’t study ahead of time, but there are ways to make that happen. The first approach should be to convince students that it is in their interest to read about the relevant core ideas before coming to a class where those ideas are explored. If gentle persuasion does not work, there are always the options of pop quizzes or advance assignments, where, for example, the student writes a summary of the relevant core idea or submits written questions or insights about the core idea before class.

If students will come to class thus prepared, the students and teachers alike will find this book very liberating from the stifling tedium of traditional lectures. Now class time can be spent in more sophisticated and interesting ways. Thus primed before class with appropriate information and understanding, students can engage in such enrichment class-time activities as:

Discussion of questions at the back of each chapter.

Discussion of other questions that emerge from the text (instructor can project the text on a screen and highlight statements in Adobe that trigger other ideas or questions.

Critique of related research papers, presented either by the teacher or the students.

Participation in assignments, problems, projects, or case studies.

Socratic dialog to explore the implications of the core ideas as they relate to neuroscience theory or to medical or psychiatric situations.

Presentation and discuss information of related neurological disorders.

Presentation and discuss related news items from television, newspapers, magazines or the Internet.

Less can be more. By condensing information that students must have a working understanding of, we increase the odds that students will remember what they really need to remember in order to think creatively and critically about subsequently presented information.

There is also the issue that professors increasingly are becoming disenchanted with traditional textbooks. There are not only the problems of cost and built-in obsolescence, but also the advent of the Internet is changing the way information is communicated and used. Students are going to use the Internet anyway to get neuroscience information. This book on core ideas is a quick way to prepare them to use the Internet in an informed and discriminating way.

Suggested Uses by Students

For the newcomer to neuroscience, the learning tactic should be to study this book first, focusing on each principle on an as-needed basis. Then the student is better prepared to understand traditional lectures or what is read in other textbooks, reviews, and primary literature. A basic learning strategy should always be to get the big picture first and then fill in the details. Students have great difficulty in approaching neuroscience in the other direction of learning details first and then trying to infer by inductive logic the core ideas. It is all too easy to get so bogged down in the massive detailed knowledge of neuroscience that the basic core ideas go undetected. Even professional neuroscientists may need the opportunity to step back from their myopic sub-specialty perspective and view the nervous system less reductionistically and more comprehensively. At least that is what I discovered as I developed this book.

W. R. (Bill) Klemm

Professor of Neuroscience

Texas A&M University

College Station, Texas 77843-4458

Web: thankyoubrain.com

1. Overview

Absolute truth is like a mirage; it tends to disappear when you approach it ... Passionately though I may seek certain answers, some will remain, like the mirage, forever beyond my reach.

Richard Leakey and Roger Lewin

Perhaps nowhere in science is this quote more appropriate than in neuroscience. Science has just emerged from the Decade of the Brain, so called because we believe that a critical mass of information and understanding now exists that tempt us to believe we can understand the great mysteries of brain and mind. Yet our search for full and absolute truth may well prove to be forever beyond our reach.

A beginning point in this search is to ask, What is a nervous system for? Plants do not have one, and plant species generally seem to survive just fine. Clearly, a nervous system is not necessary for evolutionary success - at least for organisms that do not move about in their environment. But creatures that move about in their environment have the opportunity to change their environment. That ability allows such organisms to have more options for survival. In short, there are ecological niches for organisms that are flexible enough to change their environment. And that is why those niches have been filled with organisms with nervous systems.

To be able to move about and change the environment, organisms have certain special needs not required by plants. Obviously, they must have a mechanism to move their protoplasm around in the environment, and it certainly helps to have a control system for coordinating movements. Additionally, such organisms need an array of sensors that inform them of the conditions of the environment in which they have the option to move, as well as a processing network that decides whether the environment is optimal or whether better conditions should be sought.

Even the most primitive one-celled animals, such as Paramecia, have some of these capabilities. The evolution of higher life forms could be regarded as Nature’s way of evolving more efficient and powerful nervous systems. Indeed a progressive refinement of nervous systems is a major theme of animal evolution, culminating in the extraordinary mental powers of humans.

In this overview category of 12 nervous system core ideas, we begin by identifying the basic operational unit of the system: the Neuron. Next, we introduce Neuron Numbers and Types, the idea of differing kinds of neurons and the importance of their occurrence in large numbers in the higher animals and humans. Then, we deal with the perplexing issue of Brain SIze, which seems to be related, but only incompletely so, to the computational power of nervous systems.

Operations of these modules and the units within them paradoxically exhibit both Stochastic and Deterministic (random and non-random) properties that give animals a basis for self-generating function as well as a sensitive ability to change function in response to changing conditions. Neurons are organized into a variety of complex Circuit Designs, which route information flow. Organization of these circuits tends to have Symmetry and Hemispheric Lateralization, yet another paradox in which seemingly incompatible concepts co-exist. We describe how many of these circuits are organized to act with Modularity and thus constitute a system of interacting modules.

Then, we discuss Topographical Mapping, the idea that much of the world, including an organism’s own body, is mapped in the brain. By such mapping, the brain has a way to represent the world within its circuits and a way to issue output commands that are appropriate to that world. Next we identify Hierarchical Control in the nervous system, a property that provides efficiency of operation typical of hierarchies in general. But the hierarchical organization of complex nervous systems is adjustable to biological demands, and thus the nervous system can be flexible. Homeostasis is a process of servo-regulating control that keeps the various nervous operations in balance. A key to the ability to exert homeostatic control is Neurohormonal Control.

The collective influence of these various ideas leads to our last idea in the overview category, Behavior. For many purposes, we can regard behavior from a simple mechanistic perspective of patterned activity of glands and muscles.

Fig. 1.1 Concept map for the principles that provide an overview of the nervous system.

Ideas in this Category:

Neuron. The Operational Unit: The basic cellular unit that mediates the information processing actions of the nervous system is the single cell type called a neuron. The essence of neurons can be captured in three words: they are specialized, numerous, and hyper dense in their interconnections.

There are other, far more numerous, cells associated with the nervous system. These cells are called glia, and they provide multiple supporting functions.

Neuron Numbers and Types: Brains have enormous numbers of computing elements (neurons) that accomplish sophisticated computation because of their large numbers, extensive interconnections, and their high degree of specialization into different types of neurons. These types vary structurally and in the neurochemical ways by which they interact.

Brain Size: Brain size and neuron number are related to mental and behavioral capabilities, but not always in any clear, simple, or linear way. In humans, key differences in intellectual competencies seem to correlate with differences in a small number of discrete brain areas, rather than with overall brain size.

Stochastic and Deterministic Properties: The brain is a highly complex system that has both stochastic and nonlinear, deterministic properties. These are big words, loaded with meaning. But they provide a crucial perspective from which to comprehend how the brain operates.

Circuit Design: Neurons are organized in certain basic circuit designs: converging, diverging, parallel, and feedback. This provides an anatomical basis for distributed, parallel processing in large networks of neurons.

Symmetry and Hemispheric Lateralization: The brain is basically bilaterally symmetrical, which is a fundamental biological principle of vertebrate structure. Many functions in the brain are not bilaterally symmetrical, but rather are controlled by neuronal groups in one or the other hemisphere. These lateralizations seem to involve mostly higher nervous system functions and seem to be more pronounced with cortical regions of the brain.

Modularity: The nervous system is organized as interacting modular subsystems.

Topographical Mapping: Major sensory and motor systems are topographically mapped. That is, the body is mapped by the nervous system. Major sensory systems map the external world within their own circuitry. Likewise, the nervous system contains a mapped control over the muscles of the body. Mapped regions may have different inputs or outputs or may share the same ones. Maps are interconnected so that projections from one map to another trigger a back projection to the first map. Mapping can persist at all levels in a given sensory or motor pathway. Body mapping may be central in the ability of the brain to acquire a conscious sense of self.

Neurohormonal Control: A major function of the nervous system is to release certain chemicals into the bloodstream that act as hormones to regulate various hormone-producing glands.

Hierarchical Control: The nervous system functions as a hierarchy of semiautonomous subsystems whose rank order is variable. There is no permanent supervisor neuron or population of neurons. Any subsystem may take part in many types of interrelationships. Whichever subsystem happens to dominate a situation, each subsystem is independent only to a certain extent, being subordinate to the subsystem above it and modulated by the inputs from its own subordinate subsystems and from other subsystems whose position in the hierarchy is ill-determined. This design feature of the mammalian nervous system provides maximum flexibility and is probably the basis for the brain’s marvelous effectiveness.

Homeostasis: The system has homeostatic control mechanisms over its own neural circuitry.

Behavior: Behavior is what emerges from the nervous system’s output to glands and muscles, particularly muscles. Behavior, in turn, has feedback actions on the brain that affect the brain’s ability to regulate behavior.

Neuron: The Operation Unit

Core Idea

The basic cellular unit that mediates the information processing actions of the nervous system is the single cell type called a neuron. The essence of neurons can be captured iln three words: 1) specialized, 2) numerous, 3) hyper-dense in their interconnections. There are other, far more numerous, cells associated with the nervous sytem. These cells are called glia, and they provide multiple support functions.

Terms

Axons: extensions of a neuron that provide an output, in the form of electrical pulses and/or chemical release, to target structures (other neurons, glands, or muscles).

Dendrites: cytoplasmic extensions of neurons that receive input, in the form of chemical or electrical stimuli, from other neurons. Some neurons that are specialized to act as sensory receptors for environmental stimuli have membrane specializations that are functional equivalents of dendrites.

Dendritic spines: small outgrowths on the membrane surfaces of dendrites. These are sites of synaptic contact with other axons and even dendrites

Glia: the other basic cell type found in the nervous system. These are not known to participate directly in the information processing reactions of the neurons.

Synapse: points of functional contact between the membranes of two or more neurons. It is here that electrochemical activity in one neuron affects the activity of a target neuron.

Explanation

In the early 20th Century the nervous system was thought to consist of a web of interconnected fibers. But a Spanish histologist, Ramon Cajal developed staining methods that indicated that there were individual cells. This gave rise to the neuron doctrine, the idea that individual nerve cells are the functionally distinct units of the nervous system. The advent of the electron microscope in 1954 provided convincing evidence that the cellular processes of neurons were distinct and separate.

We now know that this view has to be modified, and in some respects, we harken back to the original idea on interconnected webs of fibers. Higher resolution microscopy and refined electrical recording methods reveal that some neurons are coupled together in that the cytoplasm of adjacent cells can communicate directly through so-called gap junctions with the cytoplasm of neighbors. Ionic currents flow from one cell to another through these gaps with essentially no delay. This electrical coupling provides rapid communication among neurons and serves to synchronize groups of neurons as functional syncytia. These gap junctions are not always open and some are even under the control of what happens in chemical synapses in the same neurons.

The essence of neurons can be captured in three words: they are specialized, numerous, and hyper dense in their interconnections. Neurons are also polarized, both in the sense of their input/output relations and in terms of being electronegative, inside relative to the outside of the cell.

Neurons are liquid-state electronic devices in which electrical current is carried by flow of ions across the neuronal membrane. These currents may be graded, waxing and waning in response to excitatory or inhibitory influences. Under sufficiently intense excitation, neurons will discharge an all or none pulsatile voltage spike, known as a nerve impulse, which typically originates in dendrites, the cytoplasmic processes that give them the appearance of a bush or a tree. The electrical activity usually spreads to the cell body, whereupon it may reach a critical magnitude that generates a pulse of electrical current, which then propagates to the distal terminals of a process called the axon. Typically, axons carry electrochemical output away from the dendrites and cell body toward other target cells. The target cells of a neuron are muscles, glands, or - within the nervous system itself - other neurons. However, in some cases, the action potential can actually propagate backwards.

Another caveat to the neuron doctrine relates to the other cell types in the nervous system: the glial cells. Most of the cells in the nervous system are not neurons, but are supporting cells called glia. There are three types of glia: 1) oligodendrocytes, which wrap their membranes around neurons and help to insulate them electrically, 2) astrocytes, which help form synapses and attach their membranes to neurons and also to blood vessels, essentially creating a structural blood-brain barrier that makes it difficult for large molecules to move between blood and brain and spinal cord, 3) and microglia, which are phagocytic cells that ingest debris.

The type that forms insulating sheaths of membrane, called myelin, serves to organize the ion channels through which current can flow along an axon into separated points, so that impulses seem to hop from one spot to another. These glial cells are sensitive to impulses in the neurons that they surround. This communication can occur at sites far removed from the axon terminals from chemical synapses and it propagates not only along the axon but also throughout the glia. Glia communicate with other glia by chemical signaling and by their own gap junctions.

Astroycytes communicate with each other by means of glial chemical transmitters and modulators and by gap junctions. Thus these glia seem to have their own parallel system of communication, operating at much slower speeds than occurs in neuron-to-neuron communication. Additionally, astrocytes detect and can absorb neurotransmitters released from neurons, thus acting as a repository for surplus released transmitter, which may then in turn be recycled back through axon terminals. Astrocytes are electrically active, but they do not discharge impulses. Rather their membrane potentials fluctuate slowly, typically in response to the ionic currents in adjacent neurons. There may be reciprocal influences of the glia potentials on the state of membrane polarization in nearby neurons. The role, if any, of these slow potential changes in brain function is not known. Recent studies indicate that astrocytes are instrumental in forming neuronal synapses.

These caveats notwithstanding, the neuron is a single cell type that is the basic functional building block of the nervous system (Fig. 1.2).

Axons terminate in branches that interact electrically or chemically with branches of other neurons. Functional contact points are called synapses. The active region of a synaptic membrane is presumed to be the isolated dark patches near the membrane which are seen in electron micrographs. These synaptic contacts permit communication between neurons, typically from one neuron’s axonal branches to the short dendrites of an adjacent neuron body or on the target cell body itself. Axons can also act on other axons.

Neuron organelles function similarly to those of any cell: chromosomes carry ancestral wisdom, mitochondria regulate energy supply, microsomal particles control biochemical synthesis, and cell membranes surround the cytoplasm and regulate transport of solutes. Mature neurons are unusual also in that they normally do not divide. We don’t know why mature neurons do not divide, but it may relate to one or more of the other unusual features of neurons: 1) they express a higher fraction of the genome than other cells, 2) RNA is clustered as Nissl substance on endoplasmic reticulum, 3) they are closely invested by supporting cells (glia), 4) they continuously exhibit pulsatile membrane voltage changes and associated ionic fluxes, and 5) they have an extensive cytoskeleton of tubules and filaments for transporting chemicals throughout an extensive proliferation of protoplasmic processes. Finally, a special pigment, lipofuscin, accumulates in the cytoplasm as the neuron ages or if its mitochondria or lysosomes are damaged.

The neuron is polarized, in more than one sense of the word. In one sense, the neuron is polarized so that it can more or less simultaneously respond to input and deliver an output. In an electrical sense, the membrane of a neuron is polarized, having a voltage difference (about 70 mV) between the inside of the neuron and the extracellular fluids that surround it. The inside is electronegative relative to the outside. Responsiveness is achieved by changes in voltage difference between the inside and outside of the cell. If an external stimulus produces depolarizing changes (inside less negative), the neuronal membrane becomes unstable, and at some point a threshold voltage is reached at which a pulsed voltage discharge occurs, serving as an output. Hyperpolarization may also occur in certain neurons. In this case, input causes the membrane voltage becomes more electronegative than usual, making it less likely that the neuron’s membrane voltage can be moved to the destabilization point where impulses can be triggered. If hyperpolarization occurs, it takes more than the usual amount of excitatory input to elicit impulse discharge.

Fig. 1.2 Diagrammatic, three-dimensional representation of a neuron constructed from a microscopic (Golgi) stain that stains all parts of a neuron. Dendrites have fuzzy extensions (spines) on their surface that under electron microscopy are seen to be enlarged and thickened patches of membrane that contain mitochondria and numerous vesicles containing neurotransmitter. Emerging from the bottom of the cell body is a single axon, which divides into many branches. From Cotman and McGaugh, 1980, with permission.

Examples

Neurons differ so much in structure that it is hard to describe a prototypical neuron. But most neurons share the same basic features as that in the diagram. There is always a distinct cell body, and typically there are extensions of cell membrane that can be visibly described as dendrites and axon. Sometimes, especially with nerves that come off the spinal cord, there are bipolar neurons (only two main processes) in which the identification of dendrite and axon is best confirmed by their function (i.e., a dendrite brings input into the cell body and an axon provides the output