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Part 2 Neurophysiology
Organization of the Brain: Cerebrum. . . . . . . 52 The Cerebral Cortex . . . . . . . . . . . . . . . . . . . . 70
Organization of the Brain: Cell Types. . . . . . . 53 Descending Motor Pathways . . . . . . . . . . . . . 71
Blood-Brain Barrier . . . . . . . . . . . . . . . . . . . . . 54 Cerebellum: Afferent Pathways. . . . . . . . . . . . 72
Synaptic Transmission: Cerebellum: Efferent Pathways . . . . . . . . . . . . 73
Morphology of Synapses . . . . . . . . . . . . . . . 55
Cutaneous Sensory Receptors . . . . . . . . . . . . . 74
Synaptic Transmission:
Neuromuscular Junction . . . . . . . . . . . . . . . 56 Cutaneous Receptors:
Pacinian Corpuscle. . . . . . . . . . . . . . . . . . . . 75
Synaptic Transmission:
Visceral Efferent Endings . . . . . . . . . . . . . . . 57 Proprioception and Reflex Pathways: I . . . . . . 76

Synaptic Transmission: Proprioception and Reflex Pathways: II . . . . . 77

Inhibitory Mechanisms . . . . . . . . . . . . . . . . 58 Proprioception and Reflex Pathways: III . . . . 78
Synaptic Transmission: Proprioception and Reflex Pathways: IV. . . . . 79
Chemical Synaptic Transmission . . . . . . . . . 59
Sensory Pathways: I . . . . . . . . . . . . . . . . . . . . . 80
Synaptic Transmission:
Temporal and Spatial Summation . . . . . . . . 60 Sensory Pathways: II . . . . . . . . . . . . . . . . . . . . 81

Cerebrospinal Fluid (CSF): Sensory Pathways: III. . . . . . . . . . . . . . . . . . . . 82

Brain Ventricles and CSF Composition . . . . 61
Visual System: Receptors. . . . . . . . . . . . . . . . . 83
Cerebrospinal Fluid (CSF):
Visual System: Visual Pathway . . . . . . . . . . . . 84
Circulation of CSF . . . . . . . . . . . . . . . . . . . . 62
Auditory System: Cochlea . . . . . . . . . . . . . . . . 85
Spinal Cord: Ventral Rami. . . . . . . . . . . . . . . . 63
Auditory System: Pathways . . . . . . . . . . . . . . . 86
Spinal Cord: Membranes and Nerve Roots . . 64
Vestibular System: Receptors . . . . . . . . . . . . . 87
Peripheral Nervous System . . . . . . . . . . . . . . . 65
Vestibular System: Vestibulospinal Tracts. . . . 88
Autonomic Nervous System: Schema . . . . . . . 66
Gustatory (Taste) System: Receptors . . . . . . . 89
Autonomic Nervous System:
Cholinergic and Adrenergic Synapses . . . . . 67 Gustatory (Taste) System: Pathways . . . . . . . . 90
Hypothalamus . . . . . . . . . . . . . . . . . . . . . . . . . 68 Olfactory System: Receptors . . . . . . . . . . . . . . 91
Limbic System . . . . . . . . . . . . . . . . . . . . . . . . . 69 Olfactory System: Pathway . . . . . . . . . . . . . . . 92
NEUROPHYSIOLOGY Organization of the Brain: Cerebrum

Central sulcus (Rolando) Postcentral gyrus

Precentral gyrus Postcentral sulcus
Precentral sulcus Superior parietal lobule

Inferior parietal lobule

Supramarginal gyrus
Angular gyrus

Parietooccipital
sulcus

Occipital pole
Frontal pole
Calcarine sulcus

Lateral sulcus
(Sylvius)

Temporal pole Superior temporal gyrus Inferior temporal gyrus

Middle temporal gyrus

Frontal lobe

Parietal lobe

Temporal lobe

Occipital lobe

Insula (island of Reil)

©

FIGURE 2.1 ORGANIZATION OF THE BRAIN: CEREBRUM•

The cerebral cortex represents the highest center for sensory and coordinates smooth motor activities and processes muscle position.
motor processing. In general, the frontal lobe processes motor, The brainstem (medulla, pons, midbrain) conveys motor and sensory
visual, speech, and personality modalities. The parietal lobe information and mediates important autonomic functions. The spinal
processes sensory information; the temporal lobe, auditory and cord receives sensory input from the body and conveys somatic and
memory modalities; and the occipital lobe, vision. The cerebellum autonomic motor information to peripheral targets (muscles, viscera).

52
Organization of the Brain: Cell Types NEUROPHYSIOLOGY

Multipolar (pyramidal) Bipolar cell of cranial n.

cell of cerebral Unipolar cell of
motor cortex sensory ganglia of
cranial nn.
Satellite cells
Astrocyte Schwann cell
Striated Interneurons
(somatic) Blood vessel Free nerve endings
muscle (unmyelinated fibers)
Encapsulated ending
Motor
endplate Specialized ending
Multipolar somatic Muscle spindle
motor cell of nuclei
Unipolar sensory cell
of cranial nn.
of dorsal spinal
Multipolar cell root ganglion
of lower brain Interneuron
Satellite cells
motor centers
Oligodendrocyte Astrocyte Myelinated afferent
Corticospinal fiber of spinal nerve
(pyramidal) fiber Multipolar visceral
Axodendritic ending motor (autonomic)
cell of spinal cord
Axosomatic ending Myelin sheath
Axoaxonic ending Autonomic preganglionic
Multipolar somatic (sympathetic or para- Red: Motor neuron
motor cell of sympathetic) nerve fiber Blue: Sensory neuron
anterior horn Myelin sheath Purple: Interneuron
of spinal cord Gray: Glial and
Autonomic postganglionic neurilemmal cells
Collateral neuron of sympathetic or and myelin
parasympathetic ganglion
Renshaw interneuron Note: Cerebellar cells
Satellite cells not shown here
(feedback)
Unmyelinated nerve fiber
Myelinated somatic motor Schwann cells
fiber of spinal nerve

Myelin sheath Schwann cells

Motor endplate with Endings on Unmyelinated fibers

Schwann cell cap cardiac muscle
or nodal cells
Striated Free nerve endings
(voluntary) Beaded
muscle varicosities
and endings on Encapsulated ending
smooth muscle
and gland cells Muscle spindle ©

FIGURE 2.2 ORGANIZATION OF THE BRAIN: CELL TYPES•

Neurons form the functional cellular units responsible for (e.g., astrocytes, oligodendrocytes), satellite cells, and other spe-
communication, and throughout the nervous system, they are cialized cells that optimize neuronal function, provide mainte-
characterized by their distinctive size and shapes (e.g., bipolar, nance functions, or protect the nervous system.
unipolar, multipolar). Supporting cells include the neuroglia

53
NEUROPHYSIOLOGY Blood-Brain Barrier

Cell
membrane
Basement
Tight membrane
junction
proteins

Cytoplasm

Red
blood
cell

Astrocyte
Capillary foot processes
lumen

Tight Capillary
junction endothelial
cell
Astrocyte
©

FIGURE 2.3 BLOOD-BRAIN BARRIER•

The blood-brain barrier (BBB) is the cellular interface between the other substances (including many drugs) from the blood to the
blood and the central nervous system (CNS; brain and spinal cord). interstitial space of the CNS is restricted by the BBB. CNS endothe-
It serves to maintain the interstitial fluid environment to ensure lial cells also exhibit a low level of pinocytotic activity across the
optimal functionality of the neurons. This barrier consists of the cell, so specific carrier systems for the transport of essential sub-
capillary endothelial cells with an elaborate network of tight junc- strates of energy and amino acid metabolism are characteristic of
tions and astrocytic foot processes that abut the endothelium and these cells. The astrocytes help transfer important metabolites from
its basement membrane. The movement of large molecules and the blood to the neurons and also remove excess K⫹ and neuro-
transmitters from the interstitial fluid.
54
Synaptic Transmission: Morphology of Synapses NEUROPHYSIOLOGY

Dendrite

Node Axon

Dendrites Myelin sheath

Numerous boutons (synaptic knobs)

of presynaptic neurons terminating
on a motor neuron and its dendrites

Enlarged section
of bouton

Axon (axoplasm)

Axolemma

Mitochondria
Glial process

Synaptic vesicles

Synaptic cleft

Presynaptic membrane
(densely staining)

Postsynaptic membrane
(densely staining)

Postsynaptic cell

FIGURE 2.4 MORPHOLOGY OF SYNAPSES•

Neurons communicate with each other and with effector targets at lower figure shows an enlargement of one such synaptic bouton.
specialized regions called synapses. The top figure shows a typical Chemical neurotransmitters are contained in synaptic vesicles,
motor neuron that receives numerous synaptic contacts on its cell which can fuse with the presynaptic membrane, release the trans-
body and associated dendrites. Incoming axons lose their myelin mitters into the synaptic cleft, and then bind to receptors situated
sheaths, exhibit extensive branching, and terminate as synaptic in the postsynaptic membrane. This synaptic transmission results in
boutons (synaptic terminals or knobs) on the motor neuron. The excitatory, inhibitory, or modulatory effects on the target cell.

55
NEUROPHYSIOLOGY Synaptic Transmission: Neuromuscular Junction

Structure of Neuromuscular Junction

Active zone
Myelin sheath
Schwann cell process Neurilemma
Axoplasm
Acetylcholine
Schwann cell
receptor sites
Mitochondria
Basement membrane
Nucleus of Schwann cell
Presynaptic membrane
Active zone
Synaptic vesicles
Synaptic trough
Basement membrane
Sarcolemma
Nucleus of
muscle cell

Myofibrils

Synaptic cleft

Postsynaptic
membrane

Junctional fold

Sarcoplasm

Acetylcholine receptor sites

FIGURE 2.5 STRUCTURE OF THE NEUROMUSCULAR JUNCTION•

Motor axons that synapse on skeletal muscle form expanded termi- ately stimulated, release their neurotransmitter into the synaptic
nals called neuromuscular junctions (motor endplates). The motor cleft. The transmitter then binds to receptors that mediate depolar-
axon loses its myelin sheath and expands into a Schwann ization of the muscle sarcolemma and initiate a muscle action
cell–invested synaptic terminal that resides within a trough in the potential. A single muscle fiber has only one neuromuscular junc-
muscle fiber. Acetylcholine-containing synaptic vesicles accumu- tion, but a motor axon can innervate multiple muscle fibers.
late adjacent to the presynaptic membrane and, when appropri-

56
Synaptic Transmission: Visceral Efferent Endings NEUROPHYSIOLOGY

Visceral Efferent Endings






















A. Smooth muscle B. Gland (submandibular) C. Neurosecretory
(posterior pituitary)
Sympathetic terminal
ending Pituicyte processes
Mucous cells
Smooth muscle cells (cut) Varicosity Axon Axon
Schwann cell
cap enclosing Schwann
Schwann cell cap enclosing
nerve axons cell cap
nerve axons

Fibroblast
Schwann cell cap
Capillary
Schwann cell Neurosecretory
Smooth muscle cells Serous cells
cap enclosing vesicles
Endothelium
Varicosities Parasympathetic nerve axons
terminal ending Collagen space
Terminal endings Mast cell
Varicosity Basement membrane

FIGURE 2.6 VISCERAL EFFERENT ENDINGS•

Neuronal efferent endings on smooth muscle (A) and glands (B rosecretion) from expanded nerve terminal endings. This arrange-
and C) exhibit unique endings unlike the presynaptic and postsy- ment allows for the stimulation of numerous target cells over a
naptic terminals observed in neuronal and neuromuscular junc- wide area. Not all smooth muscle cells are innervated. They are
tion synapses. Rather, neurotransmitter substances are released connected to adjacent cells by gap junctions and can therefore
into interstitial spaces (A and B) or into the bloodstream (C, neu- contract together with the innervated cells.

57
NEUROPHYSIOLOGY Synaptic Transmission: Inhibitory Mechanisms

I E
E (Inhibitory (Excitatory
(Excitatory fiber) fiber)
fiber)

Motor
neuron Motor
neuron

I
Axon (Inhibitory Axon
fiber)

mV
⫹20
mV
A. Only E fires
90-mV spike 90 mV A′. Only E fires
in E terminal
⫺70 EPSP in
⫺60 motor ⫺60
EPSP in neuron
motor neuron ⫺70 ⫺70

B. Only I fires B′. Only I fires

Long-lasting
partial Motor
⫺60
depolarization ⫺70 neuron
in E terminal hyper- ⫺70
No response in polarized
⫺80
motor neuron ⫺70

C. I fires before E C′. I fires before E

Partial ⫺60
depolarization Depolariza-
⫹20
of E terminal tion of motor ⫺70
reduces spike neuron less
to 80 mV, thus 80 mV than if only ⫺80
releasing less E fires
transmitter ⫺70
substance ⫺60
Smaller EPSP in
motor neuron ⫺70 ©

FIGURE 2.7 SYNAPTIC INHIBITORY MECHANISMS•

Inhibitory synapses modulate neuronal activity. Illustrated here is presynaptic inhibition (left panel) and postsynaptic inhibition (right
panel) at a motor neuron.

58
Synaptic Transmission: Chemical Synaptic Transmission NEUROPHYSIOLOGY















Excitatory Inhibitory
Synaptic
vesicles
in synaptic
bouton
Presynaptic
membrane
Transmitter
Na
substances









  

Synaptic cleft  
   
 Cl
K
Postsynaptic
membrane

When impulse reaches excitatory synaptic At inhibitory synapse, transmitter substance re-
bouton, it causes release of a transmitter leased by an impulse increases permeability of
substance into synaptic cleft. This increases the postsynaptic membrane to Cl. K
moves
permeability of postsynaptic membrane to out of post-synaptic cell but no net flow of Cl
Na
and K
. More Na
moves into postsy- occurs at resting membrane potential
naptic cell than K
moves out, due to great-
er electrochemical gradient

Synaptic bouton

Resultant net ionic current flow is in a direc- Resultant ionic current flow is in direction that tends to
tion that tends to depolarize postsynaptic cell. hyperpolarize postsynaptic cell. This makes depolariza-
If depolarization reaches firing threshold, an tion by excitatory synapses more difficult—more depolari-
impulse is generated in postsynaptic cell zation is required to reach threshold
msec
Current 0 4 8 12 16
70
Potential (mV)
Potential (mV)

65 Potential
Potential

75 Current
70
0 4 8 12 16
msec
Current flow and potential change Current flow and potential change ©

FIGURE 2.8 CHEMICAL SYNAPTIC TRANSMISSION•

Chemical synaptic transmission between neurons may be excita- threshold for an action potential. During inhibition (right column),
tory or inhibitory. During excitation (left column), a net increase in the opening of K
and Cl channels drives the membrane potential
the inward flow of Na
compared with the outward flow of K
away from threshold (hyperpolarization) and decreases the proba-
results in a depolarizing potential change (excitatory postsynaptic bility that the neuron will reach threshold (inhibitory postsynaptic
potential [EPSP]) that drives the postsynaptic cell closer to its potential [IPSP]) for an action potential.

59
NEUROPHYSIOLOGY Synaptic Transmission: Temporal and Spatial Summation
Temporal and Spatial Summation
of Excitation and Inhibition

Excitatory fibers mV Excitatory fibers mV

–70 –70

Axon Axon

Inhibitory fibers Inhibitory fibers

A. Resting state: motor nerve cell shown with synaptic boutons of B. Partial depolarization: impulse from one excitatory fiber has
excitatory and inhibitory nerve fibers ending close to it caused partial (below firing threshold) depolarization of motor
neuron

Excitatory fibers mV
Excitatory fibers mV
–70
–70

Axon
Axon
Inhibitory fibers
Inhibitory fibers
C. Temporal excitatory summation: a series of impulses in one D. Spatial excitatory summation: impulses in two excitatory fibers
excitatory fiber together produce a suprathreshold depolarization cause two synaptic depolarizations that together reach firing
that triggers an action potential threshold triggering an action potential

Excitatory fibers mV Excitatory fibers mV

–70 –70

Axon Axon

Inhibitory fibers Inhibitory fibers

E. Spatial excitatory summation with inhibition: impulses from E. (continued): motor neuron now receives additional excitatory
two excitatory fibers reach motor neuron but impulses from impulses and reaches firing threshold despite a simultaneous
inhibitory fiber prevent depolarization from reaching threshold inhibitory impulse; additional inhibitory impulses might still
prevent firing ©

CHART 2.1 SUMMARY OF SOME NEUROTRANSMITTERS AND WHERE WITHIN THE CENTRAL AND PERIPHERAL NERVOUS
SYSTEM THEY ARE FOUND

Transmitter Location Transmitter Location

Acetylcholine Neuromuscular junction, autonomic end- Gas
ings and ganglia, CNS Nitric oxide CNS, GI tract
Biogenic amines Peptides
Norepinephrine Sympathetic endings, CNS ␤-Endorphins CNS, GI tract
Dopamine CNS Enkephalins CNS
Serotonin CNS, GI tract Antidiuretic CNS (hypothalamus/posterior
Amino acids hormone pituitary)
␥-Aminobutyric CNS Pituitary-releasing CNS (hypothalamus/anterior
acid (GABA) hormones pituitary)
Glutamate CNS Somatostatin CNS, GI tract
Purines Neuropeptide Y CNS
Adenosine CNS Vasoactive
Adenosine CNS intestinal peptide CNS, GI tract
triphosphate (ATP)

CNS, Central nervous system; GI, gastrointestinal.

FIGURE 2.9 TEMPORAL AND SPATIAL SUMMATION •

Neurons receive multiple excitatory and inhibitory inputs. Tempo- tial (panel D). Both temporal and spatial summation can be modu-
ral summation occurs when a series of subthreshold impulses in lated by simultaneous inhibitory input (panel E). Inhibitory and
one excitatory fiber produces an action potential in the postsynap- excitatory neurons use a wide variety of neurotransmitters, some of
tic cell (panel C). Spatial summation occurs when subthreshold which are summarized here.
impulses from two or more different fibers trigger an action poten-

60
Cerebrospinal Fluid (CSF): Brain Ventricles and CSF Composition NEUROPHYSIOLOGY
Left lateral phantom view
Right lateral ventricle Frontal (anterior) horn
Central part Left
lateral
Temporal (inferior) horn ventricle
Occipital (posterior) horn

Cerebral aqueduct (Sylvius)

4th ventricle

Left lateral aperture

(foramen of Luschka)

Left lateral recess

Left interventricular
foramen (Monro) Median aperture
(foramen of Magendie)
3rd ventricle

Central canal of spinal cord

CHART 2.2 CSF COMPOSITION

CSF Blood Plasma

Na⫹ (mEq/L) 140–145 135–147
K⫹ (mEq/L) 3 3.5–5.0
Cl− (mEq/L) 115–120 95–105
HCO3− (mEq/L) 20 22–28
Glucose (mg/dL) 50–75 70–110
Protein (g/dL) 0.05–0.07 6.0–7.8
pH 7.3 7.35–7.45

FIGURE 2.10 BRAIN VENTRICLES AND CSF COMPOSITION•

CSF circulates through the four brain ventricles (two lateral ventri- secretes the CSF. Importantly, the CSF has a lower [HCO3⫺] than
cles and a third and fourth ventricle) and in the subarachnoid plasma and therefore a lower pH. This allows small changes in
space surrounding the brain and spinal cord. The electrolyte com- blood PCO2 to cause changes in CSF pH, which in turn regulates
position of the CSF is regulated by the choroid plexus, which the rate of respiration (see Chapter 5).

61
NEUROPHYSIOLOGY Cerebrospinal Fluid (CSF): Circulation of CSF

Choroid plexus of lateral

ventricle (phantom) Superior sagittal sinus

Cistern of corpus callosum Subarachnoid space

Arachnoid granulations
Dura mater

Arachnoid

Interventricular
foramen (Monro)

Choroid plexus of 3rd ventricle

Cerebral aqueduct (Sylvius)

Lateral aperture (foramen of Luschka)

Choroid plexus of 4th ventricle

Median aperture
(foramen of Magendie)
Dura mater

Arachnoid

Subarachnoid space

Central canal of spinal cord

FIGURE 2.11 CIRCULATION OF CEREBROSPINAL FLUID•

CSF circulates through the four brain ventricles (two lateral ventri- reabsorbed into the venous system through the arachnoid granula-
cles and a third and fourth ventricle) and in the subarachnoid tions and through the walls of the capillaries of the central nervous
space surrounding the brain and spinal cord. Most of the CSF is system and pia mater.

62
Spinal Cord: Ventral Rami NEUROPHYSIOLOGY

Base of skull
C1 vertebra (atlas)
C1 spinal nerve
Cervical plexus
C8 spinal nerve
Brachial plexus
T1 spinal nerve

Spinal dura mater

1st rib
Filaments
of spinal
nerve roots
Intercostal (T7 and T8)
nerves

T12 spinal
nerve

L1
12th rib vertebra

Conus medullaris Lumbar plexus

L1 spinal nerve
L5 vertebra
Cauda equina

S1 spinal nerve
Sacral plexus
Sacrum (cut away)
Termination of
dural sac
Sciatic nerve
Coccygeal nerve

Coccyx

FIGURE 2.12 SPINAL CORD AND VENTRAL RAMI IN SITU•

The spinal cord gives rise to 31 pairs of spinal nerves that distribute fibers of these spinal nerves innervate skeletal muscle, and sensory
segmentally to the body. These nerves are organized into plexuses fibers convey information back to the central nervous system from
that distribute to the neck (cervical plexus), upper limb (brachial the skin, skeletal muscles, and joints.
plexus), and pelvis and lower limb (lumbosacral plexus). Motor

63
NEUROPHYSIOLOGY Spinal Cord: Membranes and Nerve Roots

Posterior view
Ventral root of spinal nerve
Dorsal root of spinal nerve
Spinal sensory (dorsal root) ganglion

Ventral ramus of spinal nerve

Dorsal ramus of spinal nerve
Dura mater

Arachnoid mater

Subarachnoid space
Pia mater overlying spinal cord
Filaments of dorsal root

Membranes removed: anterior view

(greatly magnified)

Gray matter
White matter Filaments of dorsal root
Dorsal root of spinal nerve
Filaments of ventral root
Spinal sensory (dorsal
root) ganglion
Dorsal ramus of
spinal nerve
Ventral ramus of
spinal nerve
Ventral root of spinal nerve
Spinal nerve
Gray and white rami
communicantes
©

FIGURE 2.13 SPINAL MEMBRANES AND NERVE ROOTS•

The spinal cord gives rise to 31 pairs of spinal nerves that distribute The spinal cord is ensheathed in three meningeal coverings: the
segmentally to the body. Motor fibers of these spinal nerves inner- outer, tough dura mater; the arachnoid mater; and the pia mater,
vate skeletal muscle, and sensory fibers convey information back which intimately ensheaths the cord itself. CSF bathes the cord and
to the central nervous system from the skin, skeletal muscles, and is found in the subarachnoid space.
joints.

64
Peripheral Nervous System NEUROPHYSIOLOGY

Posterior horn Dorsal root ganglion

Sensory neuron cell body

Dorsal root

Anterior horn
Motor neuron cell body
Ventral root

Peripheral nerve

Axon

Myelin sheath

Motor neuron Sensory neuron

Neuromuscular
junction
Skin

Muscle

with

FIGURE 2.14 PERIPHERAL NERVOUS SYSTEM•

The peripheral nervous system (PNS) consists of all of the neural shown here (see Figure 2.15 for the autonomic nervous system).
elements outside of the CNS (brain and spinal cord) and provides The somatic component of the peripheral nerves contains both
the connections between the CNS and all other body organ sys- motor and sensory axons. Cell bodies of the motor neurons are
tems. The PNS consists of somatic and autonomic components. The found in the anterior horn gray matter, whereas the cell bodies of
somatic component innervates skeletal muscle and skin and is sensory neurons are located in the dorsal root ganglia.

65
NEUROPHYSIOLOGY Autonomic Nervous System: Schema

Oculomotor nerve (III) Intracranial vessels

Facial nerve (VII) Ciliary ganglion
Glossopharyngeal nerve (IX) Eye
Pterygopalatine ganglion
Medulla oblongata
Lacrimal glands
Vagus nerve (X) Otic ganglion
Parotid glands
C1

Gray ra ntes
Submandibular ganglion
Sublingual and

mi
C2

nica
submandibular glands
C3

m u
C4 Peripheral cranial

m
and facial vessels

o
C5
c
C6 Larynx
C7 Trachea
C8 Bronchi
Lungs
Sweat T1
gland T2 Pulmonary plexus
T3
tes

Heart Cardiac plexus

Gray and white rami communican

T4 Greater  Splanchnic
Peripheral T5 Lesser  nerves
blood vessel Celiac ganglion
T6 Least 
Stomach
T7 Aorticorenal ganglion
Liver
T8 Gallbladder
Bile ducts
T9
Lumbar Pancreas
T10 splanchnic Superior mesenteric
nerves ganglion
Suprarenal glands
T11
Arrector (smooth) Kidneys
T12 Inferior mesenteric
muscle of hair follicle
ganglion
Note: Above three L1
structures are shown at
L2 Intestines
only one level but Superior hypogastric
occur at all levels L3 plexus
cantes

L4 Descending colon
L5 Sigmoid colon
m uni

Note: Blue-shaded S1 Rectum

Inferior
co m

areas indicate zones of hypogastric

parasympathetic S2 Urinary bladder
plexus
a mi

outflow from CNS S3

ay r

S4 Pelvic
S5 splanchnic Prostate
Gr

Coccygeal nerves External genitalia

©
Presynaptic Presynaptic
Sympathetic Parasympathetic Antidromic
fibers Postsynaptic fibers Postsynaptic conduction

FIGURE 2.15 AUTONOMIC NERVOUS SYSTEM: SCHEMA•

The autonomic nervous system is composed of two divisions: the a peripheral autonomic ganglion. Postganglionic axons of the auto-
parasympathetic division derived from four of the cranial nerves nomic nervous system innervate smooth muscle, cardiac muscle, and
(CN III, VII, IX, and X) and the S2-S4 sacral spinal cord levels, and the glands. Basically, the sympathetic division mobilizes our body (“fight
sympathetic division associated with the thoracic and upper lumbar or flight”) while the parasympathetic division regulates digestive and
spinal cord levels (T1-L2). The autonomic nervous system is a two- homeostatic functions. Normally, both divisions work in concert to
neuron chain, with the preganglionic neuron arising from the central regulate visceral activity (respiration, cardiovascular function, diges-
nervous system and synapsing on a postganglionic neuron located in tion, and associated glandular activity).

66
Autonomic Nervous System: Cholinergic and Adrenergic Synapses NEUROPHYSIOLOGY

Parotid
Glossopharyngeal gland
nerve (IX)
Larynx
Medulla Internal carotid nerve Trachea
oblongata Bronchi
Lungs
Vagus nerve (X)

Heart
Cervical
sympathetic
ganglia Striated muscle

Sweat glands

White ramus Hair follicles

Thoracic Gray ramus communicans
part of communicans
Celiac
spinal cord
ganglion Peripheral arteries

Superior
mesenteric Visceral
Upper ganglion arteries
lumbar
part of Gastrointestinal
spinal cord tract
(L1-2 [3]) Suprarenal
gland

Inferior
mesenteric ganglion

Sacral Pelvic splanchnic nerves

part of Urinary bladder
spinal cord
C Cholinergic synapses Urethra
Prostate
A Adrenergic synapses

Presynaptic Presynaptic
Sympathetic Parasympathetic
fibers Postsynaptic fibers Postsynaptic

Somatic fibers

Antidromic conduction
©

FIGURE 2.16 CHOLINERGIC AND ADRENERGIC SYNAPSES: SCHEMA•

The autonomic nervous system (ANS) is a two-neuron chain, with aline (norepinephrine) at its postganglionic synapses, causing
the preganglionic neuron arising from the central nervous system adrenergic (A) effects (except on sweat glands, where acetylcholine
and synapsing on a postganglionic neuron located in a peripheral is released). Although acetylcholine and noradrenaline are the
autonomic ganglion. Acetylcholine is the neurotransmitter in both chief transmitter substances, other neuroactive peptides often are
the sympathetic and parasympathetic ganglia. The parasympathetic colocalized with them and include such substances as gamma-
division of the ANS releases acetylcholine at its postganglionic aminobutyric acid (GABA), substance P, enkephalins, histamine,
synapses and is characterized as having cholinergic (C) effects, glutamic acid, neuropeptide Y, and others.
whereas the sympathetic division releases predominantly noradren-
67
NEUROPHYSIOLOGY Hypothalamus

Corpus callosum
Septum Fornix
pellucidum
Lateral
ventricle

From
Thalamus hippocampal
formation
Interthalamic
Lateral adhesion
hypothalamic area
Paraventricular nucleus
Medial Anterior Anterior hypothalamic area
forebrain commissure Dorsal hypothalamic area
bundle
Dorsomedial nucleus
Mamillothalamic tract
Posterior area
Lateral Periventricular
preoptic Medial nucleus
nucleus preoptic Nucleus
nucleus intercalatus

Olfactory
tract

Fornix Cerebral
Red nucleus peduncle
Ventromedial
nucleus Mamillary
Optic (II) complex
nerve Tuberohypophyseal tract Dorsal
Optic chiasm longitudinal
Oculomotor (III) nerve fasciculus
Supraoptic nucleus Descending
hypothalamic
Supraopticohypophyseal connections
tract
Pons
Posterior lobe of pituitary
Anterior Reticular
lobe of formation
pituitary ©

CHART 2.3 MAJOR FUNCTIONS OF THE HYPOTHALAMUS

Hypothalamic Area Major Functions*

Preoptic and anterior Heat loss center: cutaneous vasodilation and sweating
Posterior Heat conservation center: cutaneous vasoconstriction and shivering
Lateral Feeding center: eating behavior
Ventromedial Satiety center: inhibits eating behavior
Supraoptic (subfornical organ and organum vasculosum) ADH and oxytocin secretion (sensation of thirst)
Paraventricular ADH and oxytocin secretion
Periventricular Releasing hormones for the anterior pituitary
*Stimulation of the center causes the responses listed.

FIGURE 2.17 SCHEMATIC RECONSTRUCTION OF THE HYPOTHALAMUS•

The hypothalamus, part of the diencephalon, controls a number of information), the thalamus (pain), the limbic system (emotion, fear,
important homeostatic systems within the body, including tempera- anger, smell), the medulla oblongata (blood pressure and heart
ture regulation, food intake, water intake, many of the endocrine rate), and the optic system, and it integrates these inputs for regula-
systems (see Chapter 8), motivation, and emotional behavior. It tion of the functions listed.
receives inputs from the reticular formation (sleep/wake cycle

68
Limbic System NEUROPHYSIOLOGY

Genu of corpus callosum

Head of caudate nucleus

Columns of fornix

Body of fornix

Thalamus

Uncus

Crura of fornix

Fimbria of hippocampus

Hippocampus

Commissure of fornix

Splenium of corpus callosum

Lateral ventricle

Body of fornix
Columns
of fornix
Commissure
of fornix

Crura of fornix

Mamillary
bodies
Hippocampus
Amygdaloid bodies with fimbria ©

FIGURE 2.18 HIPPOCAMPUS AND FORNIX•

The limbic system includes the hypothalamus and a collection of pocampal formation. The limbic system functions in linking emo-
interconnected structures in the telencephalon (cingulate, parahip- tion and motivation (amygdala), learning and memory (hippocam-
pocampal, and subcallosal gyri), as well as the amygdala and hip- pal formation), and sexual behavior (hypothalamus).

69
NEUROPHYSIOLOGY The Cerebral Cortex

Ms I Sm I
Motor Sensory
Premotor; orientation; Ms II Sm II
eye and head Sensory
movements analysis

Prefrontal;
inhibitory control
of behavior;
higher
intelligence Visual III

Visual II

Visual I

Motor control Language;

of speech reading; speech
Auditory I
Auditory II

Ms I Sm I
Motor Sensory
Ms II Sm III
Premotor ?
Prefrontal;
inhibitory control
of behavior;
higher
intelligence
Visual III

Visual II

Visual I

Cingulate gyrus ©
(emotional behavior) Corpus callosum
and cingulum Hippocampal commissure
Olfactory Anterior commissure

FIGURE 2.19 CEREBRAL CORTEX: LOCALIZATION OF FUNCTION AND ASSOCIATION PATHWAYS•

The cerebral cortex is organized into functional regions. In addi- including aspects of memory storage and recall, language, higher
tion to specific areas devoted to sensory and motor functions, there cognitive functions, conscious perception, sensory integration, and
are areas that integrate information from multiple sources. The planning/execution of complex motor activity. General cortical
cerebral cortex participates in advanced intellectual functions, areas associated with these functions are illustrated.

70
Descending Motor Pathways NEUROPHYSIOLOGY

Motor Hip

Trunk
Shoulder
Elbow
cortex

Fin rist
Knee

rs
Ankle

ge
b
um
Internal
Toes Th eck
capsule N
Browd
Eyeli
Nares Lateral aspect of
Lips
cerebral cortex to show
Tongue
Larynx topographic projection
of motor centers on
precentral gyrus

Midbrain

Motor system
Basis
pedunculi Fibers originate in motor cortex and
descend via posterior limb of internal
capsule to basis pedunculi of midbrain
Pons
Longitudinal bundles branch upon
entering basis pontis and rejoin to
Basis enter pyramids of medulla
pontis
At lower medulla, bulk of fibers cross
median plane to form lateral
corticospinal tract; some fibers
Medulla continue downward in ipsilateral
lateral corticospinal tract; others
descending ipsilateral anterior
Pyramids corticospinal tract
Synapse occurs at spinal level: Lateral
corticospinal fibers synapse on
ipsilateral anterior horn cells; anterior
Medulla corticospinal fibers synapse on
contralateral anterior horn cells
Decussation
of pyramids
Above mid-
thoracic
level
Motor
endplate
Spinal Anterior corticospinal tract
cord Below mid-
thoracic Lateral corticospinal tract
level
Motor
endplate
©

FIGURE 2.20 CORTICOSPINAL TRACTS•

The corticospinal, or pyramidal, tract is the major motor tract that cord constitute the upper portion of the system (upper motor neu-
controls voluntary movement of the skeletal muscles, especially ron). The anterior horn cells and their associated axons constitute
skilled movements of distal muscles of the limbs. All structures the lower portion of the system (lower motor neuron).
from the cerebral cortex to the anterior horn cells in the spinal

71
NEUROPHYSIOLOGY Cerebellum: Afferent Pathways

Superior cerebellar peduncle

Middle cerebellar peduncle

To contralateral cerebellar cortex

Cortical input
Leg
Nucleus Arm
reticularis Fac Primary fissure
tegmenti e
pontis
Pontine nuclei
(contralateral)
Spinal input
Inferior olive
Upper part
of medulla To nodule and flocculus
oblongata Vestibular Inferior cerebellar peduncle
Spinal input nuclei
Functional Subdivisions of Cerebellum
Hemisphere Vermis
Reticulocerebellar
Inter-
tract Lateral mediate
Vestibular nerve Cuneocerebellar part part
and ganglion Anterior lobe
tract Leg zone Primary
Lower part Gracile nucleus Arm zone fissure
of medulla Face zone
Main cuneate
oblongata
nucleus (relay Middle
Cortical input for cutaneous (posterior)
2nd spinal
information) lobe
Lateral reticular projection
nucleus External cuneate nucleus area (gracile
(relay for proprioceptive lobule) Posterolateral
Spinal input information) fissure
Archi-
Cervical part From skin (touch Lingula
cerebellum Flocculo-
of spinal cord and pressure) Flocculus
(vestibulo- nodular lobe
Motor interneuron From muscle (spindles cerebellum) Nodule
Rostral
and Golgi tendon organs) Paleocerebellum Uvula
spinocerebellar From skin and (spinocerebellum) Pyramid
Vermis
tract deep tissues
Neocerebellum Middle vermis
(pain and Golgi
Spinal border cells (pontocerebellum) Hemisphere
tendon organs)
Schema of
Motor interneuron From skin (touch theoretical
Lumbar part and pressure) “unfolding”
of spinal cord and from muscle of cerebellar
(spindles and surface in
Clarke’s column Golgi tendon derivation of
Ventral spino- organs) above diagram
cerebellar tract Dorsal spinocerebellar tract

FIGURE 2.21 CEREBELLAR AFFERENT PATHWAYS•

The cerebellum plays an important role in coordinating movement. It bellum primarily controls movement of the proximal portions of the
receives sensory information and then influences descending motor limbs. It receives sensory information on limb position and muscle
pathways to produce fine, smooth, and coordinated motion. The tone and then modifies and coordinates these movements through
cerebellum is divided into three general areas: archicerebellum (also efferent pathways to the appropriate descending motor pathways. The
called vestibulocerebellum) paleocerebellum (also called spinocere- neocerebellum is the largest portion of the cerebellum, and it coordi-
bellum) and the neocerebellum (also called the cerebrocerebellum). nates the movement of the distal portions of the limbs. It receives
The archicerebellum is primarily involved in controlling posture and input from the cerebral cortex and thus helps in the planning of
balance, as well as the movement of the head and eyes. It receives motor activity (e.g., seeing a pencil and then planning and executing
afferent signals from the vestibular apparatus and then sends efferent the movement of the arm and hand to pick it up).
fibers to the appropriate descending motor pathways. The paleocere-

72
Cerebellum: Efferent Pathways NEUROPHYSIOLOGY

Excitatory
endings
Motor and
Inhibitory premotor
endings of cerebral
Purkinje cortex
cells

Ventral anterior and Internal capsule

ventral lateral nuclei
of thalamus Cerebral peduncle
Mesencephalic reticular formation Decussation of superior
cerebellar peduncles
Red nucleus
Descending fibers from
Fastigial nucleus superior cerebellar
peduncles
Globose nuclei
Hook bundle of Russell
Emboliform nucleus

Dentate nucleus Section

A–B
Cerebellar viewed
cortex from
below

Section
B–C
viewed
from
above
Vestibular
nuclei
Inferior cerebellar
peduncle A
Planes of
Inferior olive section: B
Lateral reticular nucleus red arrows
Medulla oblongata indicate
C
direction
Pontomedullary reticular formation of view ©

FIGURE 2.22 CEREBELLAR EFFERENT PATHWAYS•

The cerebellum plays an important role in coordinating movement. the proximal portions of the limbs. It modifies and coordinates
It influences descending motor pathways to produce fine, smooth, these movements through efferent pathways to the appropriate
and coordinated motion. The archicerebellum is primarily involved descending motor pathways. The neocerebellum coordinates the
in controlling posture and balance and movement of the head and movement of the distal portions of the limbs. It helps in the plan-
eyes. It sends efferent fibers to the appropriate descending motor ning of motor activity (e.g., seeing a pencil and then planning and
pathways. The paleocerebellum primarily controls movement of executing the movement of the arm and hand to pick it up).

73
NEUROPHYSIOLOGY Cutaneous Sensory Receptors

Free nerve endings Meissner’s corpuscle Stratum corneum

Hair shaft Pore of sweat gland Stratum
lucidum Stratum
Melanocyte

Epidermis
granulosum
Arrector muscle of hair
Sebaceous gland Stratum
Cuticle spinosum
Stratum
Internal sheath basale
Hair follicle

External sheath

Glassy Dermal
membrane papilla
(of papillary
Connective
layer)

Dermis
tissue layer

Reticular
layer
Hair cuticle

Sweat gland

Subcutaneous tissue
Hair matrix

Papilla of hair follicle

Pacinian
corpuscle
Artery Subcutaneous
Vein artery and vein
Sensory nerves Basement membrane
Elastic fibers Axon terminal
Cutaneous Mitochondrion
Detail of Merkel’s disc Skin ligaments nerve Schwann cell
(retinacula cutis)
Motor
(autonomic)
Basal nerve
epithelial Desmosomes Cross section
cells

Merkel
Cytoplasmic cell
protrusion
Lobulated
Mitochondria nucleus

Granulated
Expanded
vesicles Axon ©
axon terminal
Schwann cells
Schwann
cell Detail of free nerve ending

FIGURE 2.23 SKIN AND CUTANEOUS RECEPTORS•

Cutaneous receptors respond to touch (mechanoreceptors), pain respond best to high-frequency stimulation (i.e., vibration).
(nociceptors), and temperature (thermoreceptors). Several different Merkel’s discs have small receptive fields and respond to touch and
types of receptors are present in skin. Meissner’s corpuscles have pressure (i.e., indenting the skin). Ruffini’s corpuscles have large
small receptive fields and respond best to stimuli that are applied receptive fields, and they also respond to touch and pressure. Free
at low frequency (i.e., flutter). The pacinian corpuscles are located nerve endings respond to pain and temperature.
in the subcutaneous tissue and have large receptive fields. They

74
Cutaneous Receptors: Pacinian Corpuscle NEUROPHYSIOLOGY

Pacinian Corpuscle
as Pressure Transducer Pressure
To amplifier

Generator potential

Action potential

1st node A. Sharp “on and off” changes in pressure at start and end
of pulse applied to lamellated capsule are transmitted to
Myelin sheath central axon and provoke generator potentials, which in
Lamellated capsule turn may trigger action potentials; there is no response to a
slow change in pressure gradient. Pressure at central core
Central core and, accordingly, generator potentials are rapidly dissipated
by viscoelastic properties of capsule (Action potentials may
Unmyelinated axon terminal be blocked by pressure at a node or by drugs)

Pressure
To amplifier

Generator potential

B. In absence of capsule, axon responds to slow as well

as to rapid changes in pressure. Generator potential Action potential
dissipates slowly, and there is no “off” response
Pressure Na+
ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ
ⴙ ⴙ ⴚ ⴙ
ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ

Pressure applied to axon terminal directly or

via capsule causes increased permeability of
membrane to Na+, thus setting up ionic
generator current through 1st node
ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴙ ⴚ ⴙ ⴙ ⴙ
ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴚ ⴙ ⴚ ⴚ ⴚ ⴚ ⴚ

If resultant depolarization at 1st node is great

enough to reach threshold, an action potential
appears which is propagated along nerve fiber ©

FIGURE 2.24 PACINIAN CORPUSCLE•

Pacinian corpuscles are mechanoreceptors that transduce mechani- producing a “generator potential.” As demonstrated in the figure,
cal forces (displacement, pressure, vibration) into action potentials pacinian corpuscles respond to the beginning and end of a
that are conveyed centrally by afferent nerve fibers. As the vis- mechanical force while the concentric lamellae dissipate slow
coelastic lamellae are displaced, the unmyelinated axon terminal changes in pressure. In the absence of the capsule, the generator
membrane’s ionic permeability is increased until it is capable of potential decays slowly and yields only a single action potential.

75
NEUROPHYSIOLOGY Proprioception and Reflex Pathways: I

Spinal Effector Mechanisms

Dorsal horn
interneuron

From motor neuron

From
Proprioceptive fibers cutaneous
Dorsal horn interneuron receptor
From
muscle
spindle

Dorsal root
ganglion
Flexor reflex interneuron To motor neuron
Ventral root
Dorsal horn interneuron

To motor neuron
␣ motor axon

Schematic representation of motor neurons

In cervical
enlargement of
spinal cord

In lumbar
Flex enlargement
rs

o rs of spinal cord Fl e x o
rs

r s
n so
so

E xt e n
Ext e
©

FIGURE 2.25 PROPRIOCEPTION: SPINAL EFFECTOR MECHANISM•

Position sense or proprioception involves input from cutaneous panel) initiate muscle contraction reflexes. The lower panel shows
mechanoreceptors, Golgi tendon organs, and muscle spindles the somatotopic distribution of the motor neuron cell bodies in the
(middle figure of upper panel). Both monosynaptic reflex pathways ventral horn of the spinal cord that innervate limb muscles (flexor
(middle figure of upper panel) and polysynaptic pathways involv- and extensor muscles of upper and lower limbs).
ing several spinal cord segments (top and bottom figures of upper

76
Proprioception and Reflex Pathways: II NEUROPHYSIOLOGY

Alpha motor neurons to extrafusal

striated muscle end plates
Gamma motor neurons to intrafusal
striated muscle end plates
Ia (A␣) fibers from annulospiral
endings (proprioception)
II (A␤) fibers from flower spray
endings (proprioception);
from paciniform corpuscles (pressure)
and pacinian corpuscles (pressure)
III (A␦) fibers from free nerve endings
and from some specialized endings
(pain and some pressure)
IV (unmyelinated) fibers from free
nerve endings (pain)
Ib (A␣) fibers from Golgi tendon
organs (proprioception)
A␣ fibers from
Golgi-type
endings
A␤ fibers from
paciniform corpuscles
and Ruffini terminals
A␦ and C fibers
from free nerve
endings
Alpha motor neuron to extrafusal
muscle fiber end plates
Gamma motor neuron to intrafusal
muscle fiber and plates
II (A␤) fiber from flower spray endings
Extrafusal
Ia (A␣) fiber from annulospiral endings muscle fiber
Intrafusal
muscle fibers
Sheath
Lymph space
Nuclear bag fiber
Nuclear chain fiber ©

Detail of
Efferent fibers
muscle spindle
Afferent fibers

FIGURE 2.26 MUSCLE AND JOINT RECEPTORS•

Muscle spindles and Golgi tendon organs send afferent signals to forces, whereas the nuclear chain fibers respond to static forces.
the brain to convey the position of limbs and help coordinate mus- Intrafusal fibers maintain appropriate tension on the nuclear bag
cle movement. Muscle spindles convey information on muscle ten- and nuclear chain fibers. If the muscle tension is too great (e.g.,
sion and contraction (dynamic forces) and muscle length (static overstretching of muscle or too heavy a load), activation of the
forces). The nuclear bag fibers respond to both dynamic and static Golgi tendon organ causes a reflex relaxation of the muscle.

77
NEUROPHYSIOLOGY Proprioception and Reflex Pathways: III

Ib fibers

Ia fibers ⫹⫹⫹⫹

Extrafusal muscle fiber

Intrafusal muscle fiber

Alpha motor neurons ⫹⫹⫹

Gamma motor neurons

A. Passive stretch. Both intrafusal and extrafusal muscle fibers stretched;

Golgi spindles activated. Reflex via Ia fibers and alpha motor neurons causes
tendon secondary contraction (basis of stretch reflexes, such as knee jerk).
organ Stretch is too weak to activate Golgi tendon organs
Ib fibers ⫹⫹
Alpha activation
from brain
Ia fibers

Extrafusal muscle fiber

Intrafusal muscle fiber
Inhibitory interneuron

Alpha motor neurons ⫹⫹

Gamma motor neurons

B. Active contraction. Central excitation of alpha motor neurons

only causes contraction of extrafusal muscle fibers with consequent
relaxation of intrafusal fibers; spindles not activated. Tension is low;
Golgi does not adjust to increased resistance. Tendon organ activated,
tendon causing relaxation
organ Alpha and
Ib fibers ⫹⫹⫹ gamma
activation
from brain
Ia fibers ⫹⫹⫹⫹

Extrafusal muscle fiber

Intrafusal muscle fiber

Alpha motor neurons ⫹⫹⫹⫹

Gamma motor neurons ⫹⫹⫹⫹

C. Active contraction with gamma coactivation. Intrafusal as well as

Golgi extrafusal fibers contract; spindles activated, reinforcing contraction
tendon stimulus via Ia fibers in accord with resistance. Tendon organ
organ activated, causing relaxation if load is too great ©

FIGURE 2.27 PROPRIOCEPTIVE REFLEX CONTROL OF MUSCLE TENSION•

Interaction of the muscle spindle and Golgi tendon organ during passive stretch of a muscle (panel A) and during a contraction (panels B
and C).

78
 Proprioception and Reflex Pathways: IV NEUROPHYSIOLOGY

B. Stretch reflex
A. Afferent inhibition (reciprocal inhibition)

From extensor spindle

receptor (Ia, II fibers) From extensor spindle
receptor (Ia, II fibers)
From flexor spindle
(Ia, II fibers) Axosomatic or
axodendritic
inhibitory
synapse
Axoaxonic presynaptic
inhibitory synapse Excitatory
synapse
To extensors
To extensors
To flexors

C. Recurrent inhibition D. Tendon organ reflex

From extensor tendon

organ (Ib fibers)

Inhibitory synapse

Renshaw cells Excitatory synapse

Collaterals
To extensors
To synergistic
muscles To flexors

E. Flexor withdrawal reflex

Nociceptive fibers

Ipsilateral Contralateral
flexion extension

Inhibitory synapse Excitatory synapse

Excitatory synapse Inhibitory synapse

To extensors To extensors
To flexors To flexors ©

FIGURE 2.28 SPINAL REFLEX PATHWAYS•

Summary of the spinal reflex pathways.

79
NEUROPHYSIOLOGY Sensory Pathways: I

Cerebral cortex:
postcentral gyrus
Posterior limb of
internal capsule

Ventral posterolateral
(VPL) nucleus of
thalamus

Mesencephalon
(cerebral peduncles)

Medial lemniscus

Gracile nucleus
Spinothalamic tract
Cuneate nucleus

Lower part of
medulla oblongata
Fasciculus gracilis

Fasciculus cuneatus
Reticular formation
Dorsal (posterior) spinal
root ganglion

Proprioception,
position Large
Cervical part Touch, myelinated
of spinal cord pressure, fibers
vibration
Small
Lateral Pain, myelinated
spinothalamic tract: temperature and unmyelin-
pain, temperature ated fibers
Lateral cervical nucleus
Ventral (anterior)
spinothalamic tract: Spinocervical tract
touch, pressure

Lumbar part
of spinal cord ©

FIGURE 2.29 SOMESTHETIC SYSTEM OF THE BODY•

Pain, temperature, and pressure sensations below the head ulti- sations to the thalamus (ventral posterolateral nucleus), whereas
mately are conveyed to the primary somatosensory cortex (post- the lateral cervical system mediates some touch, vibratory, and
central gyrus) by the anterolateral system (spinothalamic and spin- proprioceptive sensations (blue and purple lines show these dual
oreticular tracts). The fasciculus gracilis and cuneatus of the spinal pathways). Ultimately, these fibers ascend as parallel pathways to
lemniscal system convey proprioceptive, vibratory, and tactile sen- the thalamus, synapse, and ascend to the cortex.

80
Sensory Pathways: II NEUROPHYSIOLOGY

Cerebral cortex:
postcentral gyrus
Ventral posteromedial (VPM)
nucleus of thalamus
Internal capsule

Midbrain
(cerebral Dorsal trigeminal lemniscus
peduncles) Trigeminal mesencephalic nucleus
Ventral Trigeminal motor nucleus
trigeminal Principal sensory trigeminal nucleus
lemniscus
Touch, pressure
Pontine Pain, temperature
reticular
formation Proprioception
Trigeminal (semilunar) ganglion
Ophthalmic n.
Maxillary n.
Pons
Sensory root
and
Motor root of mandibular n.

Medullary
reticular
formation
Spinal
trigeminal
tract
Spinal
trigeminal
nucleus

Cervical
part of
spinal Facial (VII) n.
cord
Vagus (X) n.
Dorsolateral fasciculus (of Lissauer)
Substantia gelatinosa (Iamina II)
©

FIGURE 2.30 SOMESTHETIC SYSTEM OF THE HEAD•

Nerve cells bodies for touch, pressure, pain, and temperature in of CN V (purple fibers). Most relay neurons project to the con-
the head are in the trigeminal (semilunar) ganglion of the trigemi- tralateral VPM nucleus of the thalamus and thence to the postcen-
nal (CN V) nerve (blue and red lines in figure). Neuronal cell bod- tral gyrus of the cerebral cortex, where they are somatotopically
ies mediating proprioception reside in the mesencephalic nucleus represented.

81
NEUROPHYSIOLOGY Sensory Pathways: III

Schematic demarcation of
dermatomes shown as distinct C2
segments. There is actually
considerable overlap between C3
C2 any two adjacent dermatomes C4
C5
C3 C6
C4
C5 C7
C8 T1
T1 T2
T2 T3
T3 T4
T4 C6 C6 T5
T6
T5 T1 T7
T6 T8
T9
T7 C5 T10
T8 C8 T11
T9 C7 T12
L1
T10 L2
L3
T11 L4
C6 L5
T12 S1
L1 C8 C7 C8 S2
C7
S2, 3 L2 S3
S4
L3 S5 S1
S2
L4 L5
L1
L2
L3

L5 S1 S2

L4

S1 S1
L5
L5
L4 L4

Levels of principal dermatomes T10 Level of umbilicus

C5 Clavicles T12 Inguinal or groin regions
C5, 6, 7 Lateral parts of upper limbs L1, 2, 3, 4 Anterior and inner surfaces of lower limbs
C8, T1 Medial sides of upper limbs L4, 5, S1 Foot
C6 Thumb L4 Medial side of great toe
C6, 7, 8 Hand S1, 2, L5 Posterior and outer surfaces of lower limbs
C8 Ring and little fingers S1 Lateral margin of foot and little toe
T4 Level of nipples S2, 3, 4 Perineum ©

FIGURE 2.31 DERMATOMES•

Sensory information below the head is localized to specific areas dorsal root is called a dermatome. This figure shows the der-
of the body, which reflect the distribution of peripheral sensory matome segments and lists key dermatome levels used by clini-
fibers that convey sensations to the spinal cord through the dorsal cians. Variability and overlap occur, so all dermatome segments
roots (sensory nerve cell bodies reside in the corresponding dorsal are only approximations.
root ganglion). The area of skin subserved by afferent fibers of one

82
Visual System: Receptors NEUROPHYSIOLOGY

A. Eyeball B. Section Inner limiting

through retina membrane
Axons at surface
Lens of retina passing
Iris Cornea via optic nerve,
Suspensory chiasm, and tract
ligament Ciliary
body to lateral
geniculate body
Ganglion cell
Anterior Posterior
chamber chamber Müller cell
(supporting glial
Ora containing cell)
serrata aqueous humor
Amacrine cell
Bipolar cell
Vitreous humor Horizontal cell
Rod
Retina Cone
Choroid Pigment cells
Sclera of choroid
Fovea
Optic nerve

Synaptic
Synaptic ending
ending fully
depolarized polarized

Synaptic bar
C. Rod in dark D. Rod in light

Photons Nucleus
of light

Rhodopsin Lumirhodopsin
Metabolic
energy Metarhodopsin
Centriole
Current Retinene Retinene (basal body)
flow + +
Opsin Opsin Na+
Na+ Vitamin A permeability
permeability Vitamin A decreased
increased
Circulation
©

FIGURE 2.32 VISUAL RECEPTORS•

The rods and cones of the retina transduce light into electrical sig- dark the cell is depolarized, but it is hyperpolarized in the light.
nals. As illustrated for the rod, light is absorbed by rhodopsin, and This electrical response to light is distinct from other receptor
through the second messenger cGMP (not shown), Na⫹ channels responses, in which the response to a stimulus results in a depolar-
in the membrane close and the cell hyperpolarizes. Thus, in the ization of the receptor cell membrane.

83
NEUROPHYSIOLOGY Visual System: Visual Pathway

Central
darker
circle
represents
G macular
G zone
Overlapping
visual fields Lightest
A shades
A
represent
B B monocular
H H fields
Each
quadrant
a different
color
Projection on Projection on
R RC C left retina right retina

P P Optic
Choroid Choroid (II) nerves
Optic chiasm
Periphery Macula
Structure of retina (schematic):
Projection on left Projection on right
A Amacrine cells
dorsal lateral dorsal lateral
B Bipolar cells
geniculate nucleus Optic tracts geniculate nucleus
C Cones
G Ganglion cells
Lateral
H Horizontal cells
geniculate
P Pigment cells
bodies
R Rods

Calcarine
fissure

Projection Projection
on left on right
occipital lobe occipital lobe ©

FIGURE 2.33 RETINOGENICULOSTRIATE VISUAL PATHWAY•

The retina has two types of photoreceptors: cones that mediate optic chiasm while information from the temporal retina remains in
color vision and rods that mediate light perception but with low the ipsilateral optic tract. Fibers synapse in the lateral geniculate
acuity. The greatest acuity is found in the region of the macula of nucleus (visual field is topographically represented here and
the retina, where only cones are found (upper left panel). Visual inverted), and signals are conveyed to the visual cortex on the
signals are conveyed by the ganglion cells whose axons course in medial surface of the occipital lobe.
the optic nerves. Visual signals from the nasal retina cross in the

84
Auditory System: Cochlea NEUROPHYSIOLOGY

Cochlear nerve
A. Membranous Semicircular
Utricle
labyrinth within canals
bony labyrinth Saccule
(path of sound
waves)

Scala vestibuli
Cochlear duct
(scala media)
Scala tympani Round window Oval window and stapes
B. Section
through turn Vestibular (Reissner’s)
of cochlea Scala vestibuli membrane
(perilymph);
weakly Cochlear duct (scala media;
positive +80 mV endolymph)
Efferent
nerve fibers Tectorial membrane
Spiral ligament
Bone
Afferent Scala tympani Outer hair cells; ⫺60 mV
nerve fibers (perilymph); 0 mV
Basilar membrane
Spiral ganglion Inner hair cell; ⫺60 mV

C. Spiral organ
of Corti Hair cells Tectorial membrane
Inner Outer

Stereocilia

Rods
and
tunnel
of Corti

Basilar membrane
Supporting cells
Spiral lamina Afferent nerve fibers
Spiral ganglion Efferent nerve fibers
As basilar membrane moves up, hairs are deflected outward, causing
depolarization of hair cells and increased firing of afferent nerve fibers ©

FIGURE 2.34 COCHLEAR RECEPTORS•

The cochlea transduces sound into electrical signals. This is response to pressure changes imparted on the oval window of the
accomplished by the hair cells, which depolarize in response to cochlea in response to vibrations of the tympanic membrane.
vibration of the basilar membrane. The basilar membrane moves in

85
NEUROPHYSIOLOGY Auditory System: Pathways

Acoustic area of
temporal lobe cortex

Medial geniculate body

Brachium of
inferior
colliculus
Inferior
colliculus
Midbrain
Correspondence
between cochlea
and acoustic area
of cortex:
Lateral Low tones
lemnisci Middle tones
Nuclei of High tones
Medulla lateral
oblongata lemnisci

Dorsal cochlear nucleus

Inferior cerebellar peduncle
Ventral cochlear nucleus
Cochlear division of
vestibulocochlear nerve

Dorsal
acoustic stria
Reticular formation Inner Outer
Trapezoid body
Spiral ganglion Hair cells
Intermediate acoustic stria
Superior olivary complex
©

FIGURE 2.35 AUDITORY PATHWAYS•

The cochlea transduces sound into electrical signals. Axons convey geniculate bodies) and then the acoustic cortex in the transverse
these signals to the dorsal and ventral cochlear nuclei, where it is gyrus of the temporal lobe, where information is tonotopically rep-
tonotopically organized. Following a series of integrated relay resented (low, middle, and high tones).
pathways, the ascending pathway projects to the thalamus (medial

86
Vestibular System: Receptors NEUROPHYSIOLOGY

A. Membranous labyrinth Superior semi-

Vestibular ganglion
circular canal
Vestibular
and cochlear Cristae within
divisions of ampullae
vestibulocochlear n. Horizontal semi-
Maculae circular canal
Saccule
Posterior semi-
Utricle circular canal
Cochlear duct
(scala media)

B. Section of crista

Opposite wall
of ampulla
Gelatinous cupula C. Section of macula
Hair tufts Otoconia
Hair cells Gelatinous otolithic
Nerve fibers membrane
Basement Hair tuft
membrane Hair cells
Supporting cells
Basement membrane
Nerve fibers

D. Structure and innervation Excitation

of hair cells
Inhibition
Kinocilium
Kinocilium
Stereocilia
Basal body Stereocilia
Cuticle
Cuticle Basal body

Hair cell (type I) Hair cell (type II)

Supporting cells Supporting cell

Afferent nerve Efferent nerve

calyx endings

Efferent nerve Afferent nerve

ending endings

Basement Myelin sheath

membrane

Myelin sheath
©

FIGURE 2.36 VESTIBULAR RECEPTORS•

The vestibular apparatus detects movement of the head in the form such as the pull of gravity. The three semicircular canals are
of linear and angular acceleration. This information is important for aligned so that the angular movement of the head can be sensed in
the control of eye movements so that the retina can be provided all planes. The sensory hair cells are located in the maculae of the
with a stable visual image. It is also important for the control of utricle and saccule and in the cristae within each ampullae.
posture. The utricle and saccule respond to linear acceleration,

87
NEUROPHYSIOLOGY Vestibular System: Vestibulospinal Tracts

Excitatory
endings Superior
Inhibitory Medial
endings Vestibular nuclei
Lateral
Inferior
Ascending fibers in medial Rostral
longitudinal fasciculi Upper
limb Trunk

Ventral Dorsal
To Lower
cerebellum limb
Caudal
Somatotopical
pattern in lateral
vestibular nucleus
Vestibular
ganglion
and
nerve
Motor neuron
(controlling
neck muscles)

Medial vestibulo- Fibers from cristae

Lateral (rotational stimuli)
spinal fibers in medial vestibulospinal
longitudinal fasciculi tract
Excitatory Excitatory
endings interneuron
to back
Inhibitory
muscles ?
? interneuron Fibers from maculae
(gravitational stimuli)

Lower
part of To flexor muscles
cervical To extensor muscles
spinal cord
? Inhibitory ending
?
To axial To axial muscles
muscles Inhibitory Excitatory ending
ending
Lumbar part of Lateral vestibulospinal tract
spinal cord

Inhibitory interneuron

Excitatory synapse

To flexor muscles
To extensor muscles ©

FIGURE 2.37 VESTIBULOSPINAL TRACTS•

Sensory input from the vestibular apparatus is used to maintain sta- cord (muscle control), cerebellum (vermis), reticular formation
bility of the head and to maintain balance and posture. Axons con- (vomiting center), extraocular muscles, and cortex (conscious per-
vey vestibular information to the vestibular nuclei in the pons, and ception). This figure shows only the spinal cord pathways.
then secondary axons distribute this information to five sites: spinal

88
Gustatory (Taste) System: Receptors NEUROPHYSIOLOGY

A. Tongue B. Section
through
Foliate vallate
papillae papilla
Taste buds

Duct of
gustatory
(Ebner’s)
gland

C. Taste bud

Epithelium
Fungiform
Vallate papillae Basement membrane
papillae
Nerve plexus
Microvilli
Nerve fibers
Taste pore emerging from
taste buds
Taste cells

Desmosomes Large nerve fiber Basement membrane Fibroblast

Epithelium Granules Small nerve fiber Schwann cell
Microvilli Intercellular space Large nerve fiber Collagen
D. Detail of taste pore E. Detail of base of receptor cells ©

FIGURE 2.38 TASTE RECEPTORS•

Taste buds on the tongue respond to various chemical stimuli. Taste larize neurons connected to the taste cells. A single taste bud can
cells, like neurons, normally have a net negative charge internally respond to more than one stimulus. The four traditional taste qualities
and are depolarized by stimuli, thus releasing transmitters that depo- that are sensed are sweet, salty, sour, and bitter.

89
NEUROPHYSIOLOGY Gustatory (Taste) System: Pathways

Ventral posteromedial (VPM)

nucleus of thalamus
Sensory cortex (just below
face area)

Lateral hypothalamic area

Amygdala

Mesencephalic Pontine taste area

nucleus
and Trigeminal (V) n.
Motor nucleus
of trigeminal n. Maxillary n.
Mandibular n.

Pons
Pterygo-
palatine
Greater petrosal n. ganglion

Geniculate ganglion

Facial (VII) n. Otic

and ganglion Lingual n.
Nervus inermedius

Rostral part of nucleus Chorda

of solitary tract tympani
Glossopharyngeal (IX) n.
Fungiform papillae

Lower Foliate papillae

part of Valiate papillae
medulla
oblongata

Petrosal (inferior)
ganglion of Epiglottis
glossopharyngeal n. Larynx
Nodose (inferior)
ganglion of vagus n.
Vagus (X) n. Superior laryngeal n.
©

FIGURE 2.39 TASTE PATHWAYS•

Depicted here are the afferent pathways leading from the taste receptors to the brainstem and, ultimately, to the sensory cortex in the
postcentral gyrus.

90
Olfactory System: Receptors NEUROPHYSIOLOGY

Olfactory A. Distribution Cribriform plate

bulb of olfactory of ethmoid bone
epithelium
(blue area)

Lateral nasal wall

Septum

B. Schema of
section through
olfactory mucosa
Cribriform plate

Schwann cell

Olfactory gland

Unmyelinated
olfactory axons
Basement
membrane
Sustentacular
cells
Endoplasmic
reticulum
Nucleus

Olfactory cells

Dendrites

Terminal bars
(desmosomes)
Olfactory rod
(vesicle)

Villi

Cilia

Mucus
©

FIGURE 2.40 OLFACTORY RECEPTORS•

The sensory cells that make up the olfactory epithelium respond to qualities that can be sensed: floral, ethereal (e.g., pears), musky,
odorants by depolarizing. Like taste buds, an olfactory cell can camphor (e.g., eucalyptus), putrid, and pungent (e.g., vinegar, pep-
respond to more than one odorant. There are six general odor permint).

91
NEUROPHYSIOLOGY Olfactory System: Pathway

Efferent fibers

Afferent fibers
Granule cell (excited by Fibers from contralateral olfactory bulb
and inhibiting to mitral
and tufted cells) Fibers to contralateral olfactory bulb

Mitral cell
Anterior commissure
Recurrent process
Tufted cell Medial olfactory stria

Periglom- Olfactory trigone and

erular cell olfactory tubercle
Glomerulus
Anterior perforated
Olfactory substance
nerve fibers
Lateral olfactory
stria

Lateral olfactory
tract nucleus

Piriform lobe

Uncus

Amygdala
(in phantom)

Entorhinal area

Olfactory epithelium Olfactory tract

Olfactory nerves Anterior olfactory nucleus

Olfactory bulb Cribriform plate of ethmoid bone ©

FIGURE 2.41 OLFACTORY PATHWAY•

Olfactory stimuli are detected by the nerve fibers of the olfactory sure (some efferent projections course to the contralateral olfactory
epithelium and conveyed to the olfactory bulb (detailed local cir- bulb, blue lines) or terminate in the ipsilateral olfactory trigone
cuitry shown in upper left panel). Integrated signals pass along the (olfactory tubercle). Axons then project to the primary olfactory
olfactory tract and centrally diverge to pass to the anterior commis- cortex (piriform cortex), entorhinal cortex, and amygdala.

92
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