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Chapter 9

Brainstem
External features of the brainstem XXX
Dorsal surface of the brainstem XXX
Ventral surface of the brainstem XXX
Internal structure of the brainstem XXX
Caudal medullaXXX
Mid-medulla XXX
Rostral medullaXXX
Midbrain XXX
Reticular formation XXX
The brainstem consists of the medulla oblongata, pons and midbrain. The archaic term bulb is
applied to the brainstem in compound anatomical names gien to nere fibres originating from,
or terminating in, the brainstem !e.g. corticobulbar refers to a"ons that arise in the cerebral
corte" and terminate in the brainstem#. $t is also used clinicall% to denote the medulla in such
terms as bulbar pals% and pseudobulbar pals%, &hich describe s%ndromes associated &ith
medullar% d%sfunction.
The brainstem lies upon the basal portion of the occipital bone !clius# and is connected to,
and largel% coered b%, the cerebellum. Caudall%, the medulla is continuous &ith the spinal cord
'ust belo& the foramen magnum. Rostrall%, the midbrain is continuous &ith the diencephalon of
the forebrain.
The brainstem contains numerous ascending and descending fibre tracts. (ome of these pass
throughout its &hole length, haing their origin in the spinal cord or cerebral hemisphere,
respectiel%) others hae their origin or termination &ithin brainstem nuclei. Certain of these
brainstem nuclei receie fibres from, or send fibres into, cranial neres, *+ pairs of &hich !$$$,
X$$# attach to the surface of the brainstem. These are -no&n as the cranial nerve nuclei. $n
addition, the brainstem contains a comple" and heterogeneous matri" of neurones -no&n as the
reticular formation, &ithin &hich a number of indiiduall% identified nuclei e"ist. The reticular
formation has seeral important functions, including control oer the leel of consciousness, the
perception of pain and regulation of the cardioascular and respirator% s%stems. $t also has
e"tensie connections &ith the cranial nere nuclei, &ith the cerebellum, and &ith brainstem and
spinal motor mechanisms, through &hich it influences moement, posture and muscle tone. The
brainstem contains the cells of origin of monoaminergic !dopamine, noradrenaline, serotonin#
neurones that hae &idespread pro'ections throughout the C.( and are important in sensor%,
motor, autonomic and cognitie functions.
External features of the brainstem
Dorsal surface of the brainstem
The dorsal surface of the brainstem can be ie&ed if the oerl%ing cerebellum is remoed b%
cutting the three pairs of nere fibre bundles, or peduncles, b% &hich it is attached on each side
!/igs 0.*, 0.1#. 2n the dorsal surface of the medulla, the midline is mar-ed b% a dorsal median
sulcus, continuous &ith that of the spinal cord. $n the caudal part of the medulla, the dorsal
columns !fasciculi gracilis and cuneatus, containing first-order sensor% neurones# continue
rostrall% from the spinal cord to their termination in the nuclei gracilis and cuneatus, the
locations of &hich are mar-ed b% t&o small eleations, the gracile and cuneate tubercles.
The caudal t&o-thirds of the medulla contain the rostral continuation of the central canal of the
spinal cord and is, therefore, sometimes referred to as the closed portion of the medulla. $n
passing rostrall%, the central canal moes progressiel% more dorsall% until, in the rostral
medulla, it opens out into the fourth entricle. This portion is sometimes referred to as the open
medulla. The floor of the fourth entricle forms a shallo&, rhomboid depression on the dorsal
surface of the rostral medulla and the pons. The transition from medulla to pons is not clearl%
delineated on the dorsal surface of the brainstem but, appro"imatel%, the caudal third of the floor
of the fourth entricle constitutes the dorsal aspect of the rostral medulla, &hile the rostral t&o-
thirds of the entricular floor is made up of the dorsal aspect of the pons. The fourth entricle is
&idest at the leel of the pontomedullar% 'unction, &here a lateral recess e"tends to the lateral
margin of the brainstem. 3t this point, a small lateral aperture !foramen of Luschka# proides
passage for C(/ &ithin the fourth entricle to reach the subarachnoid space surrounding the
brain. The lateral &alls of the rostral part of the fourth entricle are made up of the superior and
inferior cerebellar peduncles, connecting the brainstem &ith the cerebellum. $n the rostral
pons, the &alls conerge until, at the pontomesencephalic 'unction, the fourth entricle becomes
continuous &ith a small channel, the cerebral aqueduct, &hich passes throughout the length of
the midbrain.
The dorsal aspect of the midbrain is mar-ed b% four paired eleations, the superior and
inferior colliculi, &hich are parts of the isual and auditor% s%stems, respectiel%. The trochlear
nerve !$V cranial nere# emerges immediatel% caudal to the inferior colliculus.
Ventral surface of the brainstem
2n the entral surface of the medulla, prominent longitudinal columns, the p%ramids, run on
either side of the entral median fissure !/igs 0.4, 0.5#. The p%ramid gies its name to the
underl%ing pramidal or corticospinal tract, &hich consists of descending fibres originating
from the ipsilateral cerebral corte". $n the caudal medulla, 67,0+8 of these fibres cross oer in
the decussation of the pramids !/igs 0.4,0.7#, partl% obscuring the entral median fissure as
the% do so, to form the lateral corticospinal tract of the spinal cord. 9ateral to the p%ramid lies an
elongated eleation, the olive, &ithin &hich lies the inferior olivar nucleus. This has
connections primaril% &ith the cerebellum and is inoled in the control of moement.
The transition from medulla to pons is clearl% delineated on the entral surface of the
brainstem. The entral part of the pons is dominated b% a transerse s%stem of fibres !the
transverse pontine fibres or pontocerebellar fibres# that originate from cells in the entral
pons !pontine nuclei# and pass through the contralateral middle cerebellar peduncle to enter
the cerebellar hemisphere. The pontine nuclei receie corticopontine fibres from the cerebral
corte" !including the motor corte"# and constitute an important connection bet&een cerebral and
cerebellar cortices inoled in the coordination of moement. The massie s%stem of transerse
pontine fibres obscures the underl%ing corticospinal tract.
The entral surface of the midbrain consists, on either side, of a large column of descending
fibres, the crus cerebri or basis pedunculi. $n the midline, the t&o crura cerebri are separated b%
a depression called the interpeduncular fossa. The crus cerebri is continuous rostrall% &ith the
internal capsule of the cerebral hemisphere !/ig. *.16# and consists of corticobulbar and
corticospinal fibres that hae left the cerebral hemisphere ia the internal capsule on their &a% to
the brainstem and spinal cord. The% are primaril% motor in function.
External features of the brainstem
The brainstem consists of the medulla oblongata, pons and midbrain.
2n the dorsal aspect of the brainstem can be seen the dorsal columns, the floor of the fourth
entricle and the superior and inferior colliculi.
The dorsal aspect of the rostral medulla and the pons form the floor of the fourth entricle)
the lateral and median apertures of the fourth entricle permit the passage of C(/ into the
subarachnoid space. The cerebral a:ueduct runs through the midbrain, beneath the colliculi.
2n the entral aspect of the brainstem can be seen the p%ramids, transerse pontine fibres
and the crura cerebri.
The inferior, middle and superior cerebellar peduncles connect the cerebellum to the
medulla, pons and midbrain, respectiel%.
Internal structure of the brainstem
Caudal medulla
3t the transition from spinal cord to medulla, the pattern of gre% and &hite matter undergoes
considerable rearrangement !/ig. 0.7#. The entral horn becomes much attenuated. The dorsal
horn is replaced b% the caudal part of the trigeminal sensor nucleus !nucleus of the spinal
tract of the trigeminal nerve#. The trigeminal sensor% nucleus is regarded as the brainstem
homologue of the dorsal horn, since it receies primar% afferent fibres cone%ing general
sensation from the head, &hich enter the brainstem in the trigeminal nere. $t is a large nucleus
that e"tends the &hole length of the brainstem and into the upper segments of the spinal cord.
This latter, caudal part of the trigeminal nucleus is particularl% associated &ith the modalities of
pain and temperature. The trigeminal nere attaches to the pons and, therefore, fibres that
terminate in the parts of the trigeminal nucleus caudal to this leel descend in a tract !the spinal
tract of the trigeminal#, &hich lies immediatel% superficial to the nucleus.
$n the entral medulla, the ma'orit% of fibres of the p%ramid undergo decussation and then pass
laterall%, dorsall% and caudall% to form the lateral corticospinal tract.
!id"medulla
2n the entral surface of the mid-medulla the p%ramids are prominent, aboe their decussation.
2n the dorsal surface, the ascending fibres of the dorsal columns reach their termination in the
gracile and cuneate nuclei, &hich appear beneath their respectie tracts !/ig. 0.;#. The dorsal
columns consist of first-order sensor% neurones) the cell bodies of these neurones lie in the dorsal
root ganglia of spinal neres and hae central processes that ascend ipsilaterall% through the cord
and into the medulla. The% terminate in the nuclei gracilis and cuneatus, upon the cell bodies of
second-order neurones. The a"ons of the second-order neurones course entrall% and mediall% as
internal arcuate fibres, decussating in the midline. Thereafter, the% turn rostrall% forming a
distinct tract, the medial lemniscus, &hich runs through the rostral medulla, the pons and
midbrain, to terminate upon third-order neurones in the entral posterior nucleus of the thalamus.
#ostral medulla
<assing into the rostral medulla, a number of ne& features appear, mostl% related to the
entricular s%stem and cerebellar connections. 2n the entral surface of the medulla, the
descending fibres of the p%ramids remain conspicuous. $mmediatel% dorsal to the medial aspect
of the p%ramid lie the ascending fibres of the medial lemniscus, on either side of the midline
!/ig. 0.6#. $n the midline is located the medullar% part of the brainstem raph= nuclear comple",
represented at this leel b% the nucleus raph= magnus, a ma'or origin of serotonergic neurones.
Dorsolateral to the p%ramid and lateral to the medial lemniscus is the inferior olivar nucleus,
l%ing &ithin the prominence of the olie. The inferior oliar% nucleus has roughl% the form of a
crenated bag &ith an opening, or hilum, facing mediall%, through &hich afferent and efferent
fibres pass. The nucleus is concerned &ith the control of moement and receies afferents from
the motor and sensor% cortices of the cerebral hemisphere and from the red nucleus of the
midbrain. $ts main efferent connection is to the cerebellum ia the inferior cerebellar peduncle.
>ithin the cerebellum, a"ons originating from the inferior oliar% nucleus, -no&n as climbing
fibres, end in e"citator% s%napses in the dentate nucleus and upon <ur-in'e cells of the cerebellar
corte".
Dorsal to the inferior oliar% nucleus and lateral to the medial lemniscus lie second-order
sensor% fibres ascending to the entral posterior thalamus from the trigeminal nucleus !the
trigeminothalamic tract or trigeminal lemniscus# and from the spinal cord !spinothalamic
fibres, referred to in the brainstem as the spinal lemniscus#.
The dorsal surface of the rostral medulla forms part of the floor of the fourth entricle. ?oth
immediatel% and deep beneath the floor of the entricle lie a number of cranial nere nuclei,
some of &hich can be clearl% identified in simpl% stained sections, others of &hich cannot.
$mmediatel% beneath the entricular floor, 'ust lateral to the midline, lies the hpoglossal
nucleus, &hich contains motor neurones innerating the muscles of the tongue ia the
h%poglossal nere. 9ateral to the h%poglossal nucleus lies the dorsal motor nucleus of the
vagus, containing preganglionic paras%mpathetic neurones that run in the agus nere. The most
caudal aspect of the entricular floor is -no&n as the area postrema. 3t this point, the blood,
brain barrier, &hich limits the passage of certain chemicals from the blood to the brain, is absent.
This region is the central site of action of substances that cause omiting !emetics#. $n the lateral
part of the floor of the fourth entricle are located the vestibular nuclei, &hich receie primar%
afferent fibres from the estibular nere. Ventromedial to the h%poglossal nucleus, close to the
midline, is located the medial longitudinal fasciculus. This consists of both ascending and
descending fibres and can be identified also in the pons and midbrain. >ithin the brainstem, it
lin-s the estibular nuclei &ith the nuclei suppl%ing the e"traocular muscles !abducens, trochlear
and oculomotor nuclei# and subseres the coordination of head and e%e moements.
The dorsolateral part of the rostral medulla is dominated b% the inferior cerebellar peduncle, or
restiform bod. This consists of fibres passing bet&een the medulla and the cerebellum.
<rominent among these are oliocerebellar fibres, connections bet&een the estibular nuclei and
the cerebellum, and the fibres of the dorsal spinocerebellar tract, cone%ing proprioceptie
information from the lo&er limb. 2n the dorsal and lateral aspects of the inferior cerebellar
peduncle lie the dorsal and entral cochlear nuclei, &hich receie afferent fibres from the
cochlear nere. Medial to the inferior cerebellar peduncle and entral to the estibular nuclei lies
the nucleus solitarius, surrounding a small, dense fascicle of fibres, the solitar% tract or tractus
solitarius. The nucleus solitarius receies isceral afferent fibres entering the brainstem in the
facial, glossophar%ngeal and agus neres. Deep beneath the entricular floor, entral to the
nucleus solitarius and 'ust dorsal to the inferior oliar% nucleus, is located the nucleus
ambiguus. This sends motor fibres into the glossophar%ngeal and agus neres and cranial roots
of the accessor% nere and, thence, to the muscles of the phar%n" and lar%n".
$ons
The pons ma% be diided into a entral, or basal, portion and a dorsal portion, also -no&n as the
tegmentum. The entral portion is mar-ed b% numerous transersel% oriented fascicles of
pontocerebellar fibres that originate from scattered cell groups, the pontine nuclei, and pass into
the contralateral side of the cerebellum through the massie middle cerebellar peduncle
!brachium pontis# !/igs 0.@,0.*+#. Corticospinal fibres !&hich continue into the medullar%
p%ramid# appear as small, separate bundles running longitudinall% bet&een the fascicles of
transerse pontine fibres.
The ascending fibres of the medial lemniscus become separated from the p%ramid and
displaced dorsall%, together &ith the spinal lemniscus and trigeminothalamic tract !trigeminal
lemniscus#, b% interening transerse pontocerebellar fibres. The medial lemniscus also rotates
through 0+A so that it lies almost horiBontall%, mar-ing the boundar% bet&een entral and
tegmental portions of the pons. $n the caudal to mid-pons !/ig. 0.0#, an additional group of
transersel% running fibres is located close to the ascending lemniscal fibres but dorsal to the
pontocerebellar fibres. This is the trape%oid bod, &hich consists of acoustic fibres crossing the
brainstem from the cochlear nuclei. The% ascend into the midbrain as the lateral lemniscus !/ig.
0.*+# and terminate in the inferior colliculus.
?eneath the floor of the fourth entricle, in the pontine tegmentum, lie a number of cranial
nere nuclei. These include the abducens nucleus !innerating the lateral rectus muscle#, the
facial motor nucleus !innerating the muscles of facial e"pression# and the trigeminal motor
nucleus !innerating the muscles of mastication#, &hich each suppl% motor a"ons to their
respectie cranial neres. 3lso the trigeminal sensor% nucleus, alread% encountered in the
medulla, reaches its ma"imum e"tent in the pons, ad'acent to the origin of the trigeminal nere.
$n the rostral half of the pons, the superior cerebellar peduncles form the lateral &alls of the
fourth entricle, the thin superior medullar% elum spanning bet&een them to form its roof. The
superior peduncle contains some cerebellar afferent fibres, such as the entral spinocerebellar
tract, &hich cone%s proprioceptie information from the lo&er limb. $t consists mainl%,
ho&eer, of ascending cerebellar efferents concerned &ith the coordination of moement, that
are destined for the red nucleus of the midbrain and the entral lateral nucleus of the thalamus.
The superior cerebellar peduncles conerge to&ards the midline as the% pass into the midbrain.
$n the most rostral part of the pons, close to the pontomesencephalic 'unction !a region called the
isthmus#, the fourth entricle becomes greatl% attenuated as it merges into the cerebral a:ueduct
!/ig. 0.*+#. Central gre% matter begins to encompass the lumen !much li-e the gre% matter
surrounding the central canal of the spinal cord# and at this leel lies a group of pigmented
neurones, the locus coeruleus, &hich is a ma'or site of noradrenergic neurones.
!idbrain
The midbrain !/igs 0.** , 0.*4# is formall% diided into dorsal and entral portions at the leel
of the cerebral a:ueduct. The dorsal portion is -no&n as the tectum, &hich consists largel% of
the inferior and superior colliculi !corpora quadrigemina#. The entral portion of the
midbrain is -no&n as the tegmentum. $t is bounded entrall% b% the massie fibre s%stem of the
crus cerebri. The term cerebral peduncle is sometimes used as a s%non%m for crus cerebri but,
strictl% spea-ing, the cerebral peduncle refers to the &hole of the entral midbrain, on either side,
e"cluding the tectum.
$n the caudal part of the midbrain, the inferior colliculus constitutes part of the ascending
acoustic !auditor# pro&ection. 3scending auditor% fibres run in the lateral lemniscus, &hich
terminates in the inferior colliculus. Cfferent fibres from the colliculus terminate in the medial
geniculate nucleus of the thalamus, &hich in turn pro'ects to the auditor% corte" of the temporal
lobe.
The superior colliculus of the rostral midbrain is part of the isual s%stem. $ts main afferents
are corticotectal fibres originating from the isual corte" of the occipital lobe and from the
frontal e%e field of the frontal lobe. These inputs are concerned &ith controlling moements of
the e%es, such as those occurring &hen a moing ob'ect is follo&ed !smooth pursuit# or &hen the
direction of the gaBe is altered !saccadic e%e moements#. $n addition, corticotectal fibres from
the isual corte" are inoled in the accommodation reflex !Ch. *+#.
3 small number of isual fibres running in the optic tract terminate 'ust rostral to the superior
colliculus, in the pretectal nucleus. This nucleus has connections &ith nearb% midbrain
preganglionic paras%mpathetic neurones 'Edinger()estphal nucleus* controlling the smooth
muscle of the e%e and is part of the circuit mediating the pupillar light reflex !Ch. *+#.
Ventral to the colliculi, the cerebral a:ueduct runs the length of the midbrain. (urrounding the
a:ueduct is a pear-shaped arrangement of gre% matter, the periaqueductal !or central# gre. $n
the entral part of the peria:ueductal gre%, at the leels of the inferior and superior colliculi,
respectiel%, lie the trochlear and oculomotor nuclei, &hich innerate the e"traocular muscles
controlling e%e moements. .ear the oculomotor nucleus lies the Cdinger->estphal nucleus.
Close to these nuclei runs the medial longitudinal fasciculus, &hich lin-s them to the abducens
nucleus in the pons and the estibular nuclei in the medulla and is important in the control of
gaBe.
3t the leel of the inferior colliculus !/igs 0.**, 0.*1#, the central portion of the tegmentum is
dominated b% the superior cerebellar peduncles !brachium con&unctivum#. These fibres
originate in the cerebellum, coursing entromediall% as the% run into the midbrain. ?eneath the
inferior colliculus, the superior cerebellar peduncles decussate in the midline. Rostral to the
decussation, at the leel of the superior colliculus !/ig. 0.*4#, the central portion of the
tegmentum is occupied b% the red nucleus, in &hich some of the fibres of the superior cerebellar
peduncle terminate. The red nucleus is inoled in motor control. $ts other ma'or source of
afferents is the motor corte" of the frontal lobe. Cfferent fibres from the red nucleus cross in the
entral tegmental decussation and descend to the spinal cord in the rubrospinal tract. $n
addition, the red nucleus pro'ects to the inferior oliar% nucleus of the medulla, ia the central
tegmental tract.
The most entral part of the midbrain tegmentum is occupied b% the substantia nigra. This
consists of t&o subnucleiD the pars compacta and the pars reticulata. The pars compacta
consists of pigmented, melanin-containing neurones that s%nthesise dopamine as their
transmitter. These neurones pro'ect to the caudate nucleus and putamen !striatum# of the basal
ganglia in the forebrain and constitute the nigrostriatal path&a% !Ch. *5#. This connection is
important for the control of oluntar% moement, posture and muscle tone and degeneration of
the pars compacta is associated &ith <ar-inson's disease. Dopamine-containing neurones e"tend
dorsomediall% from the entromedial part of the pars compacta into a region -no&n as the
entral tegmental area !VT3) /ig. 0.*4#. These cells of the VT3 are the origin of the ascending
mesolimbic dopaminergic path&a% and the% innerate a number of forebrain structures including
the nucleus accumbens, am%gdala and septum and the cingulate, orbital and prefrontal cortices
!/ig. 0.*5#. The non-pigmented subdiision of the substantia nigra is called the pars reticulata. $t
is considered to be a functional homologue of the internal segment of the globus pallidus, &hich
is also part of the basal ganglia, and the t&o share er% similar connections.
Ventral to the substantia nigra lies the massie crus cerebri. This consists entirel% of
descending cortical efferent fibres that hae left the cerebral hemisphere b% traersing the
internal capsule. 3ppro"imatel% the middle 7+8 of the crus consists of corticobulbar and
corticospinal fibres. The corticobulbar fibres end predominantl% in or near the motor cranial
nere nuclei of the brainstem. The corticospinal !p%ramidal# fibres traerse the pons to enter the
medullar% p%ramid and, thence, the corticospinal tracts.
2n either side of the corticobulbar and corticospinal fibres, the crus cerebri contains
corticopontine fibres that originate from &idespread regions of the cerebral corte" and terminate
in the pontine nuclei of the entral pons. /rom the pontine nuclei connections are established
&ith the cerebellum, ia the middle cerebellar peduncle, that are inoled in the coordination of
moement.
#eticular formation
The reticular formation consists of a comple" matri" of neurones that e"tends throughout the
length of the brainstem. This is, in ph%logenetic terms, a relatiel% old part of the brainstem and
its neurones fulfil a number of important functions, some of &hich are necessar% for surial.
The reticular formation has &idespread afferent and efferent connections &ith other parts of the
C.(, &hich reflect its comple" and multimodal functions. (ome reticular neurones hae long
a"ons that ascend and descend for considerable distances &ithin the brainstem, allo&ing profuse
interactions throughout the neura"is. >ithin the reticular formation a number of more-or-less
&ell-defined nuclei can be identified &hile some functions are subsered b% more dispersed
net&or-s that do not correspond e"actl% to anatomicall% identified nuclei. The latter applies to
the so-called respirator% and cardioascular centres. These consist of diffuse neuronal net&or-s
located &ithin the lateral medullar% and caudal pontine reticular formation that control
respirator% moements and cardioascular function.
Descending reticulospinal tracts originate from the medullar% and pontine reticular formation
!Ch. @#. These predominantl% influence muscle tone and posture. >ithin the reticular formation
of the medulla lies a group of particularl% large cell bodies, the nucleus reticularis
gigantocellularis, that contributes descending fibres to the lateral reticulospinal tract.
(ome of the ascending fibres of the reticular formation constitute the so-called reticular
activating sstem. These neurones, man% of &hich are cholinergic, receie input, either directl%
or indirectl%, from multiple sensor% sources. Through the intermediar% of thalamic nuclei
!mostl% the intralaminar nuclei#, the% cause actiation of the cerebral corte" and heightened
arousal.
The raph+ nuclei are a series of midline nuclei that e"tend throughout the length of the
brainstem !/igs 0.6, 0.*1#. Man% of the neurones are serotonergic, utilising serotonin !7-
h%dro"%tr%ptamine, 7-ET# as their transmitter. Their a"ons are &idel% distributed throughout the
C.(, including ascending pro'ections to the thalamus, h%pothalamus, striatum, am%gdala,
hippocampus and &idespread regions of the cerebral corte" !/ig. 0.*7#. The functional
significance of such dierse connections is not full% understood but ascending fibres to forebrain
structures are inoled in mood and cognitie function and in the neural mechanisms of sleep.
There are also serotonergic pro'ections to the cerebellum and descending fibres to the spinal
cord. The latter originate from the nucleus raph= magnus of the medulla and are inoled in the
modulation of nociceptie mechanisms in the dorsal horn !/ig. @.**#. (ome of these descending
neurones contain en-ephalin.
The locus coeruleus is a group of pigmented neurones that lies in the rostral pontine
tegmentum !/ig. 0.*+#. $t is the principal noradrenergic cell group of the brain, of &hich there are
a number in the brainstem. $t pro'ects to the cerebellum and man% areas of the forebrain,
including the diencephalon, limbic structures and cerebral corte" !/ig. 0.*;#. The locus
coeruleus, li-e the raph= nuclei, has been implicated in the neural mechanisms regulating sleep,
particularl% RCM !rapid e%e moement# sleep. The noradrenergic inneration of the forebrain
also appears to be important in higher mental functions such as mood and cognition. 2ther
noradrenergic fibres pro'ect throughout the brainstem and spinal cord.
Internal structure of the brainstem
Cranial neres $$$,X$$ attach to the brainstem, their fibres either originating from, or
terminating in, the cranial nere nuclei.
The reticular formation controls the leel of consciousness, the cardioascular s%stem and
the respirator% s%stem.
3scending sensor% s%stems pass through the brainstem en route to the thalamus. /irst-order
proprioceptie fibres in the dorsal columns rela% in the dorsal column nuclei. (econd-order fibres
decussate to form the medial lemniscus. (pinothalamic fibres form the spinal lemniscus. (econd-
order fibres originating in the trigeminal sensor% nucleus constitute the trigeminothalamic tract
!trigeminal lemniscus#.
Descending fibre s%stems end in the brainstem, pass through it and originate &ithin it.
Corticobulbar fibres terminate in the midbrain, pons and medulla. The corticospinal tract
runs through the crus cerebri, the basal part of the pons and the medullar% p%ramid) 67,0+8 of
fibres cross in the p%ramidal decussation to form the lateral corticospinal tract.
The reticular formation, red nucleus and estibular nuclei gie rise to descending fibres that
pass to the spinal cord.
Brainstem lesions
3 unilateral brainstem lesion caused b% stro-e, tumour or multiple sclerosis causes ipsilateral
cranial nere d%sfunction, contralateral spastic hemiparesis, h%perrefle"ia and an e"tensor
plantar response !upper motor neurone lesion#, contralateral hemisensor% loss and ipsilateral
incoordination !/ig. 0.*6#. 3 bilateral lesion destro%s the ital centres that control breathing
and the circulation, leading to coma and death.
,igure 9-. Dorsal aspect of the brainstem-
,igure 9-/ Lateral aspect of the brainstem-
,igure 9-0 Ventral aspect of the brainstem-
,igure 9-1 Ventral aspect of the brainstem sho2ing the decussation of the pramids-
,igure 9-3 4ransverse section through the caudal medulla at the level of the decussation of
the pramids- The sections sho&n in /igures 0.7,0.*4 hae been stained b% the >eigert,<al
method. 3reas rich in nere fibres stain dar-l%, &hile areas rich in cell bodies are relatiel% pale.
,igure 9-5 4ransverse section through the mid"medulla at the level of the great sensor
decussation-
,igure 9-6 4ransverse section through the rostral medulla at the level of the inferior
olivar nucleus-
,igure 9-7 4ransverse section through the caudal pons-
,igure 9-9 4ransverse section through the mid"pons at the level of the trigeminal nerve-
,igure 9-.8 4ransverse section through the rostral pons-
,igure 9-.. 4ransverse section through the brainstem at the level of the pontine(
mesencephalic &unction-
,igure 9-./ 4ransverse section through the caudal midbrain at the level of the inferior
colliculus-
,igure 9-.0 4ransverse section through the rostral midbrain at the level of the superior
colliculus-
,igure 9-.1 Dopaminergic neurones and their principal pro&ections-
,igure 9-.6 Brainstem lesion-
,igure 9-.3 9erotonergic neurones and their principal pro&ections-
,igure 9-.5 :oradrenergic neurones and their principal pro&ections-

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