Neuroscience

Second Edition
 Section K – Lipid metabolism

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Neuroscience
A. Longstaff
Science writer and freelance lecturer in neuroscience
© BIOS Scientific Publishers, 2000

First published 2000

Reprinted 2003

All rights reserved. No part of this book may be reproduced or transmitted, in any form or
by any means, without permission.

A CIP catalogue record for this book is available from the British Library.

ISBN 1 85996 082 0

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C ONTENTS

Abbreviations viii
Preface xi

Section A – Brain cells 1

A1 Neuron structure 1
A2 Classes and numbers of neurons 4
A3 Morphology of chemical synapses 7
A4 Glial cells and myelination 10
A5 Blood–brain barrier 13

Section B – Foundations of electrophysiology 15

B1 Resting potentials 15
B2 Action potentials 19
B3 Voltage-dependent ion channels 22
B4 Molecular biology of sodium and potassium channels 26
B5 Action potential conduction 31

Section C – Synapse function 35

C1 Overview of synaptic function 35
C2 Fast neurotransmission 38
C3 Slow neurotransmission 43
C4 Receptor molecular biology 48
C5 Neurotransmitter release 53
C6 Voltage-dependent calcium channels 59
C7 Neurotransmitter inactivation 61
C8 Autoreceptors 64

Section D – Neural integration 67

D1 Properties of neurites 67
D2 Spatial and temporal summation 72

Section E – Elements of neuroanatomy 75

E1 Organization of the peripheral nervous system 75
E2 Organization of the central nervous system 81
E3 Neuroanatomical techniques 88
E4 Brain imaging 91
E5 Meninges and cerebrospinal fluid 95

Section F – Neural coding 99

F1 Information representation by neurons 99
F2 Intensity and time coding 101
F3 Stimulus localization 104
F4 Stimulus quality 107
vi Contents

Section G – Somatosensory systems 111

G1 Cutaneous sensory receptors 111
G2 Dorsal column pathways for touch sensations 116
G3 Anterolateral systems and descending control of pain 120
G4 Sense of balance 126

Section H – Vision 131

H1 Attributes of vision 131
H2 Eye and visual pathways 135
H3 Retina 141
H4 Phototransduction 146
H5 Retinal processing 150
H6 Early visual processing 156
H7 Parallel processing in the visual system 161

Section I – Auditory neuroscience 167

I1 Acoustics and audition 167
I2 Anatomy and physiology of the ear 169
I3 Peripheral auditory processing 173
I4 Central auditory processing 175

Section J – Chemical senses: smell and taste 181

J1 Olfactory receptor neurons 181
J2 Olfactory pathways 183
J3 Taste 186
J4 Gustatory pathways 189

Section K – Motor function: muscle and cortex 191

K1 Skeletal muscles and excitation–contraction coupling 191
K2 Motor units and motor pools 197
K3 Elementary motor reflexes 202
K4 Spinal motor function 209
K5 Brainstem postural reflexes 215
K6 Cortical control of voluntary movement 221
K7 Motor disorders 229

Section L – Cerebellum and basal ganglia 233

L1 Anatomy of the cerebellum 233
L2 Cerebellar cortical circuitry 237
L3 Functional subdivisions of the cerebellum 240
L4 Cerebellar function 246
L5 Anatomy of the basal ganglia 250
L6 Basal ganglia function 254
L7 Oculomotor control 258

Section M – Neuroendocrinology and autonomic functions 265

M1 Anatomy and connections of the hypothalamus 265
M2 Posterior pituitary function 270
M3 Neuroendocrine control of metabolism and growth 274
M4 Neuroendocrine control of reproduction 283
M5 Smooth and cardiac muscle 288
M6 Autonomic nervous system function 292
M7 Central control of autonomic function 299
Contents vii

Section N – Diffuse monoamine transmission 305

N1 Dopamine neurotransmission 305
N2 Norepinephrine neurotransmission 309
N3 Serotonin neurotransmission 313
N4 Acetylcholine neurotransmission 316
Section O – Brain and behavior 319
O1 Motivation 319
O2 Control of feeding 324
O3 Brain biological clocks 329
O4 Sleep 333
Section P – Developmental neurobiology 339
P1 Early patterning of the nervous system 339
P2 Cell determination 346
P3 Cortical development 351
P4 Axon pathfinding 355
P5 Synaptogenesis and developmental plasticity 360
P6 Neurotrophic factors 365
P7 Brain sexual differentiation 370
Section Q – Learning and memory 375
Q1 Types of learning 375
Q2 Invertebrate procedural learning 380
Q3 Memory circuitry in mammals 384
Q4 Hippocampal learning 390
Q5 Cerebellar motor learning 397
Section R – Aspects of neuropathology 401
R1 Strokes and excitotoxicity 401
R2 Epilepsy 405
R3 Parkinson’s disease 409
R4 Alzheimer’s disease 413

Further reading 421

Index 427
A BBREVIATIONS

ACh acetylcholine CVLM caudal ventrolateral medulla

AChE acetylcholinesterase CVO circumventricular organ
ACTH adrenocorticotrophic hormone DAG diacylglycerol
AD Alzheimer’s disease DBL dorsal blastopore lip
AII angiotensin II DCML dorsal column–medial lemniscal
AMPA α-amino-3-hydroxy-5-methyl-4- system
isoxazole-proprionic acid DCN dorsal column nuclei
ANS autonomic nervous system 2-DG 2-deoxyglucose
AP action potential DHC dorsal horn cell
apoE apolipoprotein E DI diabetes insipidus
APP amyloid precursor protein DLPN dorsolateral pontine nucleus
APV D-2-amino-5-phosphonovalerate DOPAC dihydroxyphenyl acetic acid
ATN anterior thalamic nuclei DRG dorsal root ganglion
ATP adenosine 5’-triphosphate DYN dynorphin
AVP arginine vasopressin ECT electroconvulsive therapy
βA β-amyloid EEG electroencephalography
βAR β adrenoceptors EGF epidermal growth factor
BAT brown adipose tissue EGL external granular layer
BDNF brain derived neurotrophic factor EMG electromyography
bl basal lamina ENK encephalin
BMP bone morphogenetic protein ENS enteric nervous system
α-BTX α-bungarotoxin epp endplate potential
CA cornu ammonis epsp excitatory postsynaptic potential
CaM calmodulin ER endoplasmic reticulum
CAM cell adhesion molecule F-actin filamentous actin
CaMKII calcium–calmodulin-dependent FEF frontal eye field
protein kinase II FF fast fatiguing
cAMP cyclic adenosine monophosphate FM frequency modulation
CAT computer assisted tomography fMRI functional magnetic resonance
cbf cerebral blood flow imaging
CC cingulate cortex FR fatigue resistant
CCK cholecystokinin FRA flexor reflex afferents
CF characteristic frequency FSH follicle stimulating hormone
cGMP 3’,5’-cyclic guanosine Gi inhibitory G protein
monophosphate Gq G protein coupled to
ChAT choline acetylesterase phospholipase
CL central laminar nucleus (of Gs stimulatory G protein
thalamus) GABA γ-aminobutyrate
CNG cyclic-nucleotide-gated channel GC guanylyl cyclase
CNS central nervous system GDP guanosine 5’-diphosphate
CoA coenzyme A GFAP glial fibrillary acidic protein
CPG central pattern generators GH growth hormone
CRH corticotrophin releasing hormone GHRH growth hormone releasing
CRO cathode ray oscilloscope hormone
CS conditioned stimulus GnRH gonadotrophin releasing hormone
CSF cerebrospinal fluid GPe globus pallidus pars externa
CVA cerebrovascular accident GPi globus pallidus pars interna
Abbreviations ix

GR glucocorticoid receptor MI primary motor cortex

GTO Golgi tendon organs MII secondary motor cortex
GTP guanosine 5’-triphosphate MLCK myosin light chain kinase
5-HIAA 5-hydroxyindoleacetic acid MLR mesencephalic locomotor region
HPA hypothalamic–pituitary–adrenal MOPEG 3-methoxy,4-hydroxy
(axis) phenylglycol
HPG hypothalamic–pituitary–gonadal MPOA medial preoptic area
(axis) MPP+ 1-methyl-4-phenyl pyridinium
HPT hypothalamic–pituitary–thyroid mpsp miniature postsynaptic potential
(axis) MPTP 1-methyl-4-phenyl-1,2,3,6-
HRP horseradish peroxidase tetrahydropyridin
5-HT 5-hydroxytryptamine (serotonin) MR mineralocorticoid receptor
5-HTP 5-hydroxytryptophan MRI magnetic resonance imaging
HVA high voltage activated MSO medial superior olivary complex
IaIN Ia inhibitory interneurons MST medial superior temporal cortex
IbIN Ib inhibitory neurons NA noradrenaline
IC inferior colliculus nAChR nicotinic cholinergic receptor
ICSS intracranial self-stimulation NGF nerve growth factor
Ig immunoglobulin NMDA N-methyl-D-aspartate
IGF-1 insulin-like growth factor 1 NMDAR N-methyl-D-aspartate receptor
IGL internal granular layer nmj neuromuscular junction
iGluR ionotrophic glutamate receptor NMR nuclear magnetic resonance
ILD interaural level differences NPY neuropeptide Y
IP3 inositol 1,4,5-trisphosphate NREM nonrapid eye movement sleep
ipsps inhibitory postsynaptic potential NRM nucleus raphe magnus
IT inferotemporal cortex NST nucleus of the solitary tract
JGA juxtoglomerular apparatus NT3-6 neurotrophins 3-6
L-DOPA L-3,4-dihydroxyphenylalanine OC olivocochlear
LC locus cerulus OCD obsessive–compulsive disorder
LCN lateral cervical nucleus OHC outer hair cells
LDCV large dense-core vesicle 6-OHDA 6-hydroxydopamine
LGN lateral geniculate nucleus ORN olfactory receptor neurons
LH luteinizing hormone OVLT vascular organ of the lamina
LSO lateral superior olivary nucleus terminalis
LTD long-term depression P parvocellular pathway
LTM long-term memory PAD primary afferent depolarization
LTN lateral tegmental nucleus PAG periaqueductal gray matter
LTP long-term potentiation Pc Purkinje cells
LVA low voltage activated PD Parkinson’s disease
M magnocellular pathway PDE phosphodiesterase
M/T mitral/tufted cells PDS paroxysmal depolarizing shifts
mAChR muscarinic cholinergic receptor PET positron emission tomography
MAO monoamine oxidase pf parallel fibers
MAP mean arterial (blood) pressure PFC prefrontal cortex
MB mammillary bodies PGO pontine–geniculate–occipital
mepp miniature endplate potential spikes
MFB medial forebrain bundle PHF paired helical filaments
MFS mossy fiber sprouting PIP2 phosphatidyl inositol-4,5-
mGluR1 type 1 metabotropic glutamate bisphosphate
receptor PKA protein kinase A
MGN medial geniculate nucleus PM premotor cortex
x Abbreviations

PNS peripheral nervous system STN subthalamic nucleus

POA preoptic area STT spinothalamic tract
POM posterior complex (medial TB trapezoid body
nucleus) of thalamus TCA tricyclic antidepressants
POMC pro-opiomelanocortin TEA tetraethylammonium
PP posterior parietal complex TENS transcutaneous electrical nerve
PRL prolactin stimulation
PSNS parasympathetic nervous system TH tyrosine hydroxylase
psp postsynaptic potential TM transmembrane
PVN paraventricular nucleus TRH thyrotropin releasing hormone
RA retinoic acid trk tyrosine kinase receptors
REM rapid eye movement sleep TSH thyroid releasing hormone
RF receptive field TTX tetrodotoxin
RHT retinohypothalamic tract UR unconditioned response
S slow twitch fiber US unconditioned stimulus
SCG superior cervical ganglion VDCC voltage-dependent calcium
SCN suprachiasmatic nucleus channel
Sc Schaffer collateral VDKC voltage-dependent potassium
SDN–POA sexually dimorphic nucleus of the channel
preoptic area VDSC voltage-dependent sodium
SER smooth endoplasmic reticulum channel
SH2 src homology domain 2 VIP vasoactive intestinal peptide
SHH sonic hedgehog protein VLH ventrolateral hypothalamus
SMA supplementary motor area VLPO ventrolateral preoptic area
SNpc substantia nigra pars compacta VMAT vesicular monoamine transporter
SNzc substantia nigra zona compacta VMH ventromedial hypothalamus
SNS sympathetic nervous system VOR vestibulo-ocular reflexes
SON supraoptic nucleus VPL ventroposterolateral nucleus (of
SP substance P thalamus)
SPL sound pressure level VPM ventroposteromedial nucleus (of
SR sarcoplasmic reticulum thalamus)
SSRI selective serotonin reuptake VRG ventral respiratory group
inhibitors VST ventral spinocerebellar tract
SSV small clear synaptic vesicle VZ ventricular zone
STM short-term memory
P REFACE

Neuroscience is one of the most rapidly advancing areas of science and, as a consequence, spawns a lit-
erature which is growing dramatically. At one level it attempts to provide a mechanistic account of the
most complex ‘device’ in the known Universe, the human brain. Moreover, neuroscience is multidisci-
plinary, having contributions from biochemistry and molecular biology, physiology, anatomy, psy-
chology and clinical medicine, to name the most obvious. For these reasons, it is becoming increasingly
difficult for lecturers and textbook authors to present neuroscience in a way that manages to be com-
prehensive, up-to-date and accessible, while still being sufficiently rigorous to prepare students to be
successful explorers of the literature for themselves. Instant Notes Neuroscience is not intended as a
replacement for lectures or the standard textbooks, but as an affordable text to supplement them, which
is of a manageable size and in a format which aids learning.
The text is designed to provide the core of the subject in 18 sections containing 93 topics. When com-
ing to a new subject, it is my experience that students commonly express two concerns: first, how to
sort out the important ideas and facts from the wealth of detail, and second, how to get to grips with
the unfamiliar terminology. Lecturers, in addition, will want students (especially later in their studies)
to be able to integrate their knowledge across the subject. Instant Notes Neuroscience attempts to address
each of these issues. Each topic is supported by a ‘Key Notes’ panel which gives a concise summary of
the crucial points. Whenever a term appears for the first time it is in bold and immediately followed by
a definition or explanation. Extensive cross-references are provided between topics so that students can
forge the links that are important for integration.
This is a much slimmer volume than most neuroscience texts, which can be dauntingly large. A num-
ber of features contribute to this. First, I have tried to minimize the amount of detail without compro-
mising the need for a database for further study. Second, while many of the methods used by
neuroscientists are included, individual experiments or items of evidence are included only where I
thought it essential to illustrate a point, or on matters that would need some justification to be con-
vincing. Third, with a few exceptions, I have restricted examples to those most appropriate to the human
condition. In so doing I have always qualified the species, since species differences matter. If not, then
rats and cats would behave as humans do, which clearly they do not!
Section A introduces the cells of the nervous system, showing how they are specialized for the func-
tions they serve. The next three sections are essentially cellular neuroscience. Section B is concerned
mostly with action potentials, Section C with synapses, while Section D deals with how nerve cells act
as information processors. These sections provide an introduction to the electrophysiological techniques
used to study nerve cells, and say something about the molecular biology of the ion channels and recep-
tors that govern their behavior. Section E takes a broad view of neuroanatomy and summarizes tech-
niques, such as brain imaging, used to investigate nervous system structure. How information is
encoded by the firing and connectivity of neurons is considered in Section F. All the material thus far
might reasonably be found in first-year courses.
The next seven sections (G–M) form the core of systems neuroscience. Section G reviews the body
senses, touch, pain and balance. Sections H and I deal with vision and hearing, respectively, while
Section J looks at the chemical senses, smell and taste. The properties of skeletal muscle, motor reflexes,
and the cortical control of voluntary movement are the subject of Section K, while the involvement of
the cerebellum (including proprioception) and the basal ganglia in movement is covered in Section L.
Neuroendocrinology, and both peripheral and central aspects of the autonomic nervous system appear
in Section M, which also (and unusually for standard neuroscience texts) includes the functions of
smooth and cardiac muscle, and the enteric nervous system. A short Section N describes the essential
features of amine transmission, the basis for much neuropharmacology, and paves the way for under-
standing aspects of behavior, such as motivation and sleep that are included in Section O. Section P is
xii Preface

an overview of how the embryonic nervous system develops, ranging from how the basic plan is genet-
ically specified, to how differences between male and female brains might arise. Section Q addresses
how the nervous system continues to rewire itself on the basis of experience (i.e., learning and mem-
ory). Finally, although quite a number of nervous system disorders are considered at appropriate places
throughout the book, section R takes the four most common neuropathologies, stroke, epilepsy,
Parkinson’s disease and Alzheimer’s dementia and looks in some detail at what has gone amiss and
what current and future treatments may do. Space precluded the inclusion of topics on two major psy-
chiatric disorders (schizophrenia and depression) in the text; it is intended that these topics will be made
available on the BIOS website free of charge. At the end of the book is a reading list for those who wish
to take their studies further.
As a student, how should you use this book? Restrict your reading only to the sections and topics
covered by your current course. That said, Sections A–F are likely to appear in, or be required knowl-
edge for, just about any neuroscience program; you will probably need to work through these first. Later
sections can be dipped into in any order. Read the main sections thoroughly first, making sure that you
understand the ideas, and use the ‘Related topics’ to make links, just as you would if you were surfing
the internet. Where appropriate, reference is made to areas that are covered in more detail in the com-
panion volumes, Instant Notes in Biochemistry, 2nd edn, and Instant Notes in Molecular Biology, 2nd edn.
At this stage you can incorporate additional material from lectures or other textbooks in the gaps at the
end of topics, or highlight things which seem to be particularly important for your course. Studying
Instant Notes Neuroscience ‘little but often’ is a good strategy. The information density in the text is high,
so many short, concentrated, bursts are much more effective than a few eight-hour stints. The more
times you work through a topic, the better your understanding, and the more likely you will be to
remember it clearly. When it comes to revision, use the ‘Key Notes’ as a prompt. In addition, you should
aim to be able to write, from memory, a few sentences about each of the terms that appears in bold in
the main text. Being able to reproduce the simpler diagrams is also an effective way of getting your
point across in an exam. Neuroscience is an extraordinary endeavor because it aims to reveal what, in
essence, it is to be human; how we behave, think and feel as we do. At the moment we are a long way
from being able to give a coherent account of any of these faculties; that there is so much still to be done
is one reason that this science is so exciting. This book is an account of the remarkable progress made
so far. I hope you find that it serves your needs well and that, like me, you enjoy discovering neuro-
science.

Acknowledgements

A number of colleagues – Barry Hunt, Vasanta Raman and John Wilkinson, all at the University of
Hertfordshire – kindly read through some individual topics and made very helpful suggestions. David
Hames (University of Leeds, UK), Kevin Alloway (Penn State University, USA), and Patricia Revest
(Queen Mary and Westfield College, London University, UK), were each brave enough to read through
the entire text and their thought-provoking comments have been important in shaping the final version.
I am very grateful to all of these people for their time and expertise. Finally, I thank Jonathan Ray,
Rachel Offord, Will Sansom and Fran Kingston at BIOS Scientific Publishers for their encouragement
and patience.

Alan Longstaff
Section A – Brain cells

A1 N EURON STRUCTURE

Key Notes

Cell body The neuron cell body contains all the subcellular organelles found in a
typical animal cell but it is specialized to maintain high rates of protein
synthesis, as shown by the ribosome-packed Nissl bodies.

Neurites Neurites are long projections from the cell body. There are two types of
neurite, dendrites and axons. Dendrites are large extensions of the cell
body and receive most of the synaptic inputs impinging onto the cell.
Neurons may have one or many dendrites. Neurons have a single axon
arising from the axon hillock. Axon terminals form the presynaptic
components of synapses.

Axon or dendrite? The two neurites can be distinguished on structural grounds. Dendrites
contain many organelles and are capable of protein synthesis. By
contrast, axons cannot synthesize protein, so axonal proteins are derived
from the cell body. Axons and dendrites both have mitochondria.
Organelles are transported into neurites via microtubules.

Related topic Morphology of chemical synapses (A3)

Cell body The cell body (soma and perikaryon are synonyms) of a neuron (see Fig. 1)
contains the nucleus, Golgi apparatus, ribosomes and other subcellular
organelles, and is responsible for most of its routine metabolic ‘housekeeping’
functions. The neuronal perikaryon is not so different from non neuronal cells
although structurally it is specialized to maintain high levels of biosynthetic
activity. The rough endoplasmic reticulum, for example, is so densely packed as
to produce distinct structures called Nissl bodies which are extremely rich in
ribosomes. This reflects the high rates of protein synthesis of which neurons are
capable.
Because there is a great variety of types of neurons, their cell bodies vary in
size considerably. The smallest are some 5–8 µm in diameter, the largest 120 µm
across.

Neurites Neurons are distinguished from other cells by neurites, long (relative to the cell
body) cylindrical processes that come in two varieties, dendrites and axons.
Dendrites are highly branched extensions of the cell body, may be up to 1 mm
in length and account for up to 90% of the surface area of many neurons.
Dendrites on some neurons are covered with hundreds of tiny projections
termed dendritic spines on which synapses (see below) are made. Nerve cells
with spines are sometimes called spiny neurons, those lacking them aspiny
neurons. A neuron may have one or many dendrites, arranged in a pattern
which is cell typical and collectively referred to as the dendritic tree. The
majority of synaptic inputs from other neurons are made on dendrites.
2 Section A – Brain cells

Apical
dendrite

Cell body

Basal dendrite

"" ~Axon hillock

Axon collaterals

-----------~Axon entering
~ white matter
50!Lm

Fig. 1. Key features of a neuron. A drawing of a pyramidal cell showing the distribution of
neurites (dendrites and axon).

Nerve cells usually have only one axon which arises typically from the cell
body but may emerge from a proximal dendrite (the end of a dendrite closest to
the soma). In either case, the site of origin is termed the axon hillock. Axons
have diameters ranging from 0.2 to 20 µm in humans (though axons of inverte-
brates can reach 1 mm) and vary in length from a few µm to over a meter. They
may be encapsulated in a myelin sheath. Axons usually branch, particularly at
their distal end (furthest from the soma). These branches are referred to as axon
collaterals. The ends of an axon are swollen terminals (or boutons) and usually
contain mitochondria and vesicles. Some axons have a tuft of branches (a
terminal arbor) at their tip, each with its terminal bouton, some have boutons
along their length where they are described as varicosities. Axon terminals form
the presynaptic component of chemical synapses.

Axon or dendrite? Axons can be distinguished from dendrites on structural grounds. Axons tend
to be long, untapered, less highly branched, never spiny and may have a myelin
sheath, whereas dendrites are shorter, tapered, highly branched and may bear
spines. Dendrites are extensions of the cell body in that they contain Golgi appa-
ratus, rough endoplasmic reticulum and ribosomes – organelles not seen in
axons. By contrast both axons and dendrites have mitochondria. Since axons do
not possess protein synthetic machinery, proteins in axons must be made in the
cell body and subsequently moved into and along the axon by a mechanism
called axoplasmic transport. Axon terminals are often rich in mitochondria
which indicates their high requirement for metabolic energy.
A1 – Neuron structure 3

Differences in organelle composition are thought to result from the differing

arrangements of microtubules in the two types of neurite. Microtubules are
long protein polymers that are part of the internal scaffolding of cells called the
cytoskeleton. Microtubules act as ‘tramlines’ along which organelles are moved
within the cell. The two ends of a microtubule are different, designated + and – ;
thus microtubules have a distinct polarity which means that organelles move
along them in a specific direction. Mitochondria move in the – to + direction
whereas other organelles move in the + to – direction. Both dendrites and axons
have microtubules, but whereas dendrites have them orientated in either direc-
tion, those in axons are always arranged with their + ends away from the cell
body. Thus axonal microtubules can transport mitochondria out of the cell body
into the axon but cannot transport other organelles. Because microtubules are
orientated in both directions in dendrites, all organelles are transported into
these neurites.
When a neuron develops, it first grows a number of processes which are
indistinguishable. It is not yet known how one of these is subsequently selected
to differentiate into an axon. The first sign that a process will become an axon is
that it grows at a much faster rate than those which are destined to be dendrites.