Anatomyof HumanExtrinsic

CardiacNervesand Ganglia
RONALD D. JANES, MD, J. CHRISTOPHER BRANDYS, MD, DAVID A. HOPKINS, PhD,
DAVID E. JOHNSTONE, MD, DAVID A. MURPHY, MD,
and J. ANDREW ARMOUR, MD, PhD

lhe anatomy of the human extrlnsk cardiac nerves them. These nerves lntm ed with sympathet-
and ganglia was reinvestigated because descriptions k cardiopulmonary nerves anterkr and posterior to
of human cardiac imervatkn vary, detailed analy- the main pulmonary artery to form the ventral and
ses of *man mammal&n cardiac hervatkn dorsal cardkpdmonary plexuses. These plexuses
reveal conskkrable shnilarMes among species and contained relatively large discrete nerves as Wdi as
the anatomic pattern of cardiac innervation ob- smalier krtercomections. Emerging from these plex-
served in subhuman mammals differs si@kantly uses to imervate the ventrkles were 3 disthct rela-
fromthoseckscmed for humans. The presence of a tively large cardiac nerves, the right and left coro-
consistent pattern of cardiac innervation in subhu- nary cardiac nems and the left lateral cardiac
man mammals raised the questkm as to whether a nerve.lnadditkntothese3majornerves,small
similar pattern exists In humans. To Investigate this, cardlacnervesarosefromtheplexusesandthetho-
the cervkal and thorack autonomic nerves and rack vagi. Histologk examinatkn of repmse&We
ganglia were dissected in 13 embalmed and 10 au- dhecWnscoWmedthepresenceofneuraltissue
topsy cadavers. All major sympathetk cardiopulmo- andkhtifiedthelocatkinsof neuronal cell bodies
nary nerves were found to arise from the stellate intheseshctures.CeUbodieswerelocatedinthe
ganglia and the caudal halves of the cervical sym- nodose, superb cervical, mlddle cervical, stellate
pathetic trunks below the level of the crkoid carti- and thorack sympathetk gang&. The middle cervi-
lage. These sympathetic cardkpulmonary nerves cal ganglia varied In size and number. Neuronal cell
usuallyconsistedof3nervesontherlghtsfdeand4 bodies were found In the cervkal and thorack sym-
ontheleft.Incontrasttowidelyacceptedreports, pMWtktlUdCSandiflSmall n-mdkthl gangga lo-
no sympathetic cardiopulmonary nerves were found cated along the coursesofthecardkpuhonaryand
toarisefromthesuperkrcervkalgangliaorthe cardiac nerves. Marked shnlMt& exist between
thoracic sympathetic trunks inferior to the stellate the anatomy of the cardiopulmonary nerves and
ganglia. Parasympathetk cardiopulmonary nerves ganglia of humans and baboons.
were found to arise from the recurrent laryngeal
nerves and the thorack vagi lmmedlately distal to (Am J Cardlol 1988;57:299-309)

S ince the earliestanatomicstudiesof human cardiac

nerves,’ the number and origin of the cardiac nerves,
been variously described.z-loThe most frequent and
perhapsmost widely acceptedanatomic descriptionof
the number and size of cervical sympathetic ganglia, cardiac innervationI consistsof 3 major sympathetic
and the location and size of the cardiac plexuseshave cardiac nerves arising from the superior, middle and
inferior cervical ganglia:the superior, middle and in-
From the Departments of Anatomy, Medicine, Physiology & Bio- ferior sympathetic cardiac nerves.These sympathetic
physics and Surgery, Dalhousie University, Halifax, Nova Sco- nerves,togetherwith a similar number of parasympa-
tia, Canada B3H 4H7. This study was supported by the New thetic cardiac nerves, have been described as inner-
Brunswick and Nova Scotia Heart Foundations and the Medical vating the heart via the cardiac plexuses.2,3,5,7,g-*1
This
Research Council of Canada (PG-18). Dr. lanes was supported classic description of the innervation of the human
by the Patton Fellowship of the Dalhousie Medical Research heart differs markedly from that of subhuman pri-
Foundation and by a Medical Research Council Fellowship. matesI and other mammals.13-~6 In the animal studies,
Manuscript received April 15, 1985; revised manuscript re- in which the cardiac nerves were identified by both
ceived June 20,1985, accepted June Z&1985. anatomic13and physiologic techniques15the sympa-
Address for reprints: J. Andrew Armour, MD, PhD, Depart- thetic cardiac nerveswere found to originate from the
ment of Physiology & Biophysics, Sir Charles Tupper Medical region of the stellate and middle cervical ganglia.No
Building, Dalhousie University, Halifax, Nova Scotia, Canada sympathetic cardiac nerves were found to arise from
B3H 4H7. the superior cervical ganglia or the thoracic sympa-

29*
900 HUMAN EXTRINSIC CARDIAC NERVES

thetic chains in these animals, which contrastswith
previous descriptions of human cardiac nerves.
Becausethe descriptionsof the anatomy of the hu-
man cardiac nerves are inconsistentand differ mark-
edly from those of subhuman primates and other
mammals, and as accurate anatomic information
about the innervation of the human heart is clinically
relevant,*e-20
we reexamined the grossand microscop
ic anatomy of the extrinsic cardiac nerves and ganglia
of man.
Methods
The grossand microscopic anatomyof the innerva-
tion of the human heart was investigatedin 23 human
cadavers.In 13embalmed* cadavers(6men, 7 women,
mean age 66 years, range 38 to 891,we completely
dissectedthe autonomic ganglia and nerves between
the base of the skull and the ninth thoracic vertebra,
tracing all nerves arising from the vagi, sympathetic
trunks and gangliato the organsthey innervated in the
neck or thorax. In 4 representative dissections, all
autonomic nerves and ganglia were removed en bloc
for histologicstudy. Dissectionsfocusingon the origins
* The embalming fluid contained 25% glycerine, 25% methyl-
hydrate, 6% phenol, 6% formaldehyde and 38% water.
of the cardiopulmonary nerves and their courses
through the mediastinum onto the heart were then
performed during autopsiesof 9 males and 1 female
(mean age 58 years,range 6 to 81). Neural tissue was
obtainedfrom 6 of theseautopsiesfor histologicexam-
ination. Drawings were made of all dissectionsand
photographswere taken of dissectionsin 4 embalmed
cadavers.
Histologic procedures were similar for all speci-
mens, except that tissues obtained from embalmed
cadaverswere first washed overnight in running tap
water. The tissueswere placed into increasing con-
centrationsof ethanol,cleared in benzene(1 hour for
embalmed tissue;20 minutes for autopsy tissue)and
then embedded in paraffin, Serial 10.rm-thick sec-
tions, cut on an MSE rotary microtome, were stained
either with cresyl violet or with cresyl violet and 1~x01
fast blue and were examined using bright-field
microscopy.
Becausea variety of nomenclatureshavebeenused
to describe human cardiac innervation, anatomic
terms similar to thoseusedin subhumanprimateshave
been adopted, where possible, to standardize termi-
nology. To facilitate interspecies comparisons, we
used “anterior” and “posterior” interchangeablywith
“ventral” and “dorsal,” respectively.The major crani-

,,\
\

FIGURE 1. Course ot the right sympathetic trunks, vagus nerve and their major branches In the neck and thorax ot embalmed cadaver 11.
Five major rlgM cardiopulmonary nervea (CPN) are Illustrated lateral to the trachea a8 they coume to the base ot the heart. The right tho-
raclc sympathetic trunk, a tlat band ot tissue. termlnatea superiorly ln a relatlvety IndIstInct right dellate gangtlon. The nerves arldng from
the medlal aspect of the right sympathetic trunk do not course to the heart.
al thoracic nerves arising from the sympathetic trunks
have been called “sympathetic” and those arising from
the vagi “parasympathetic.” The nerves and plexuses
have been identified as cardiopulmonary, as they pre-
sumably innervate pulmonary tissue as well.
Results
Cardiopulmonary nerves in man arise from the
stellate ganglia and the caudal halves of the cervical
sympathetic trunks and, together with nerves which
arise from the recurrent laryngeal nerves or the vagi
immediately distal to them, form 2 cardiopulmonary
plexuses. These plexuses contain distinct nerves that
unite to form 3 major cardiac nerves at the base of the
heart which project onto the heart. In addition to these
major nerves, other small cardiac nerves arise from
the plexuses as well as from the thoracic vagi posterior
to the pulmonary hila. Many macroscopic and micro-
scopic ganglia are found in the cardiopulmonary
nerves.
Gross anatomy: Figures 1 and 2 show anatomic
overviews of the right and left sympathetic trunks, vagi
and some of the major cardiopulmonary nerves. The
superior cervical ganglion on either side is situated at
the base of the skull posterior to the carotid sheath,
which, at this level, contains the internal carotid ar-
tery, the internal jugular vein and the cervical vagus
nerve. The superior cervical ganglia give rise to rami
which are connected to the upper cervical nerves (Cl
to CA) and to the adjacent vagi. The pharyngeal and
superior laryngeal nerves arise from the vagi at or
immediately inferior to the nodose ganglion. The cer-
vical sympathetic trunks, which extend inferiorly from
the superior cervical ganglia anterior to the preverte-
bra1 fascia, give rise to fine branches that connect with
the cervical nerves (C2 to CS].
One to 4 visible ganglionic swellings are located in
each cervical sympathetic trunk immediately cranial
to its junction with the ventral and dorsal subclavian
ansae. One of these ganglia is consistently located at
the junction of the ventral and dorsal ansae. The visi-
ble ganglionic swellings and microscopically identi-
fied ganglia along the inferior half of the cervical sym-
pathetic trunk were named, collectively, “middle
cervical ganglia.” Each dorsal subclavian ansa arises
as a craniomedial extension of the stellate ganglion
[Fig. 3 and 4) and is shorter and thicker than the ventral
ansa, which loops anteriorly around the subclavian
artery as a thin nerve. In a few cadavers, on one or
both sides, there was no distinct dorsal ansa because
the stellate and inferiormost middle cervical ganglia
formed a large swelling, with a narrowing in the mid-

Right Coronary Cardiac Nerve

-.
*%\I -L ..

dcal

Sympathetic Trunk

Scalenus Medius

Thoracic Sympathetic Trunk

‘Thwacic V-8

FIGURE 2. The left ventral cardiopulmonary nerve of embalmed cadaver 11, Whkh can be seen as lt passes ante&r to the arch of the aorta,
Is the main contributor to the right coronary cardiac nerve at the base of the heart. Cardiopulmonary branches from the left thorack vagus
arise at the level of the pulmonary hllum. The @fl thoraclc sympathetic trunk ls a relatively flat band of tissue, whkh gradually enlarges to
terminate In a relatively dlstlnct lefl stellate gangllon. The small nerves seen arlslng from the left thoraclc sympathetic trunk course to the
aorta, esophagus and adjacent muscle.
302 HUMAN EXTRINSIC CARDIAC NERVES
dle (Fig. 4D).The stellateganglionsometimesconnects
to the inferiormost middle cervical ganglionby a small
ansa passingaround the vertebral artery (Fig. 3B).
The stellate ganglia are consistently located over
the headsof the first and secondribs asenlargements
at the superiorends of the thoracicsympathetictrunks.
Macroscopically,the stellategangliavary in shapeand
sometimes merge indistinctly with the most inferior
middle cervical ganglionor inferiorly with the thoracic
sympathetictrunk, or both. Sometimesthe stellategan-
glion is a distinct enlargement composed of one or
more swellings (Fig.3A). In no cadaverdid we identify
macroscopically separate inferior cervical and first
thoracic ganglia.Rami from the stellate ganglion con-
nect with the first and second intercostal nerves, as
well as the eighth cervical and first thoracic divisions
of the brachial plexus (Fig. 3).Small nerves originate
from the stellate ganglion and courseto the vertebral
and subclavian arteries and medially to adjacentves-
sels and muscles. The thoracic sympathetic trunks
coursing inferiorly from the stellate ganglia are flat,
band-like structures with indistinct enlargements
along their lengths [Fig. 1 and 2).
All the sympathetic cardiopulmonary nerves we
identified arosefrom the stellate ganglia and the cau-
da1halvesof the cervical sympathetictrunks below the
level of the cricoid cartilage;none originated from the
superior cervical ganglia or directly from the cranial
cervical sympathetictrunks. No nerve arising from the
thoracic sympathetic trunks inferior to the stellate
ganglia interconnectedwith cardiopulmonary nerves
or passedto the heart directly. The small nerves that
arise from the’ superior thoracic sympathetic ganglia
passmedially to the adjacentvessels,muscle, aorta or
esophagus,where they become too fine to dissect
further.
i ,.‘”
f -.
A Cervical !$nipdi~tii Trunk
-4
Scalenus Anterior
Nerve 10 Subclavian Artery
Thoracic Sympathetic Trunk
Dorsal Lateral CPN
FIGURE 3. Drawlng of the connectkns of the stellate @gfia of embalmed cadaver 11. A, the right stellate oardkpulmonary nerve (CPN)
arlses from the rlght stellate gaiigfh&ih$ @maI Irtqral CPN from a right mldqtle cervical gangllon and the dorsal medlal CPN from a mere
superior kcatlon on the Mrvlcal symp&hetk trunk. B, ihe IefMtellbte &N arises from the I& stelfate gangllon, the left dorsal lateral CPN
from a left mlddle cervkal gangllq, the IefidorSal &al
ventral CPN from the left ventral ansa.
The right sympathetic cardiopulmonary nerves
usually comprise 3 major nerves which arise from the
region of the right stellate and middle cervical ganglia
[Fig. 3AJ. The right stellate cardiopulmonary nerve
arisesfrom the right stellate ganglion,the right dorsal
lateral cardiopulmonary nerve from the dorsal ansaor
the most inferior right middle cervical ganglion, and
the right dorsal medial cardiopulmonary nerve from
the right cervical sympathetictrunk in associationwith
a more superiorly locatedright middle cervical gangli-
on. Although this was the most consistentpattern ob-
served,the origin and number of the right sympathetic
cardiopulmonary nervesvaried. For instance,in some
cadavers a cardiopulmonary nerve arose from the
ventral ansa(Fig.4A),while in otherssmall sympathet-
ic cardiopulmonary nerves interconnected before
forming the dorsal medial and dorsal lateral cardio-
pulmonary nerves [Fig. 4B). No cardiopulmonary
nerve arosedirectly from the right cervical vagus.Va-
gal branchesfrom the recurrent laryngeal nerves,the
recurrent cardiopulmonarynerves,join the dorsalme-
dial and dorsal lateral cardiopulmonary nerves (Fig.
4A). Cranial and caudal vagal cardiopulmonary
nervesarise from the right thoracicvagusbetween the
origin of the right recurrent laryngeal nerve and the
level of the ostium of the azygousvein and course
toward the heart anterior to the trachea (Fig. 1 and 5).
The right thoracic vagusgives off small branchespos-
terior to the pulmonary hilum, which coursemedially
to the heart.Largebranchesfrom both the right cranial
and caudal vagal cardiopulmonary nerves course to
the pulmonary hilum, where they pass to the lung,
posterior to the secondarybronchi.
The left sympatheticcardiopulmonary nervesarise
from the region of the left stellate and middle cervical
ganglia, the left stellate cardiopulmonary nerve from
:, .’
B
Dorsal Medial C P N
Middle Cervical Ga
Dorsal Lateral C P
Ventral Ansa C P N
Thoracic Sympathetic Trunk
&N Tom a r$re superior posltkn on the cervical sympathetic trunk and the left
 Februatv 1. 1986 THE AMERICAN JOURMAL OF CA!WlO!-CC;Y Volume 57 303

the left stellate ganglion,the left dorsal lateral cardio-
pulmonary nerve from the most inferior left middle
cervical ganglionand the left dorsal medial cardiopul-
monary nerve from the left cervical sympathetictrunk
in associationwith a more superiorly located left mid-
dle cervical ganglion (Fig. 3B, 4C and D). The ventral
cardiopulmonary nerve arises from the region of the
left middle cervical ganglia(Fig. 4D) or the left ventral
ansa or both (Fig. 3B) and coursestoward the heart
anteriorto the aortic arch (Fig.21.The left vagal cardio-

A Right Side Left Side

Cerwcal Sympalhellc Trunkd

MIddIeCarwcal Gangka

Sutlawan Artery
lb Cerwal Sympalhetlc Trunk

Recurrent Laryngeal Nerve

Stellate Ganglion

Ventral Ansa Nerve

Dorsal Medial C P N

rsa, Lateral C P N

‘Craw Vagal C.P.N

CervCal SympalhellC

&al &opulmcnary Nerves

MiddleCerwcal Gangka Cerwal Sympathetic Trunk

MtddleCerwcal Ganglia

Aecurrenl LaryngealNerve
Ventral Ansa Nerve

CardiopulmonaryNer
Dorsal Medial C P N

Dorsal Lateral C P N.
Darsal Lateral C P N

bB, CramalVagal C P N

FIGURE 4. Variations In the slzo and shape of the stellate and mlddle cervical ganglia, a8 well as the orlgln and number of the
cardiopulmonary nerves (CPN). Sf/pp/ed areas Indicate locatlons of hlstologlcally Identlfled neural cell bodies. As there were no
connections between the left vague and the left sympathetic CPNs near their orlgln, the vagus has been omltled from panels C and D for
clarity. A, In autopsy cadaver 10, the right recurrent CPNs from the rlght recurrent laryngeal nerve unlted wlth both the right dorsal medlal
and dorsal lateral CPNs. There was no rlght stellate CPN but a nerve dld arks from the ventral ansa, the right ventral ansa CPN. B, In autop-
sy cadaver 5 there were 3 relatively large rlgM mlddle cervkal ganglia. Several CPNs arose from these ganglia and 1 CPN arose from the
rlght stellate gangllon. These nerves lnterconnected wlth CPNs arlslng from the rlght vagus before forming the right dorsal medial and
doraal lateral CPNs. C, In embalmed cadaver 2 there were many neural lnfercomWWns on the Ien Me, which uHlmately formed 5 dfscrefe
CPNs, 4 dorsal CPNs (left stellate CPN, left dorsal lateral CPN, left dorsal IntermedIate CPN and left dorsal medtal CPN) and 1 ventral CPN
(left ventral CPN). There are several mlddle cervkal and medlastlnal ganglia. A photomlcrograph of these ganglia ls shown ln Figure ‘IA. D,
In autopsy cadaver 10 there were only 3 left CPNs, and no left stellate CPN was found. The stellate ganglkn was connected directly to the
most Inferior lefl mlddte cervkal gang&n as a continuous swelling.
so4 HUMAN EXTRINSIC CARDIAC NERVES

Left Dorsal Medial CPN

Right Dorsal Lateral CPN Left Dorsal Lateral CPN

Lett Pulmonary

Right
Lett Lateral
Cardiac Nerve

Left Ventral Cardiopulmonary Nerve

Right Dorsal Medial
Cardiopulmonary Nerve Left Dorsal Medial CPN

Right Dorsal Lateral CPN Left Dorsal Lateral CPN

Left Stellate CPN

Right Stellate CPN-

/T& Vagus

Vagus

-Left Recurrent Laryngeal Nerve

Right Cranial Vagal CPN
- Left Coronary Cardiac Nerve

i - Lett Cranial Vagal CPN

kwk Left Lateral Cardiac Nerve

Right Caudal Vagal CPN”

Left Caudal Vagal CPN
Right Coronary Cardiac Nerve

FIGURE 5. Right and left parasympathetic and eympathetlc cardlopuhnonary nerve8 (CPN) In the medlastlnum superior to the heart. A, the
rlght dorsal Iat$ral CPN passes anterior to the right pulmonary artery and postorlor to the main pulmonary artery. The Interconnections of
the dorsal plexus posterior to the pulmonary artery are shown In broken /he& B, the pulmonary artery (PA) Is removed to Illwtrate the con-
nectlonr of the dorsal plexus. The right coronary card& nerve courses along the right coronary artery, the lefl coronary cardiac nerve
along the left coronary artery and the left lateral cardiac nerve onto the left lateral myocardlum. Fine InterconnectIons between nerves, as
well as from nerves to ad)acent vascular structures, are omltted for clarity. LAD = left anterior desiendlng coronary artery; SVC = ruperlor
vena cava.
 ?%Bruary 1, 1986 T’REZWERICAN JO~JRNA~- OF CARCIOLOGY Volume 57 305

pulmonary nerves consist of the recurrent cardiopul- atria1 appendage and anterior to the left pulmonary
monary nerves, which arise from the left recurrent veins, before it courses onto the left lateral wall of the
laryngeal nerve, and the cranial and caudal vagal car- ventricle. The smaller ventral cardiopulm~onary plex-
diopulmonary nerves, which arise from the left vagus us is located anterior to the aorta and pulmonary artery
distal to the origin of the left recurrent laryngeal nerve and is connected to the dorsal cardiopulmonary plexus
[Fig. 5). The left caudal vagal cardiopulmonary nerves by nerves passing around both sides of the pulmonary
arise from the left vagus posterior to the pulmonary artery. The left ventral cardiopulmonary nerve unites
hilum. On the left side, as on the right, nerves connect with small nerves from the ventral plexus to form the
the vagus and the left sympathetic cardiopulmonary right coronary cardiac nerve, the latter projects onto
nerves only at or below the level of the recurrent la- the heart along the right coronary artery.
ryngeal nerve (Fig. 5B). The left parasympathetic car- Microscopic anatomy: Histologic examination re-
diopulmonary nerves, before interconnecting with vealed that the nerves and ganglia have relatively
other nerves in the dorsal and ventral plexuses, give thick connective tissue sheaths (Fig. 6). Neuronal cell
rise to large pulmonary branches that enter the left bodies are in the nodose and superior cervical ganglia.
lung posterior to its major bronchi. A number of esoph- The sensory neurons of the nodose ganglia are larger
ageal branches arise from the left recurrent laryngeal than the neurons of the superior cervical ganglia. Cell
nerve as it passes superiorly along the left lateral side bodies are located throughout the cervical vagi and
of the esophagus, sympathetic trunks, although visible ganglionic swell-
The left and right cardiopulmonary nerves have ings are only found in the inferior halves of the cervi-
relatively consistent courses through the superior me- cal sympathetic trunks. The sympathetic ganglia along
diastinum. The right stellate cardiopulmonary nerve the cervical sympathetic trunks vary in size and shape
passes along the posteromedial surface of the superior (Fig. 4) and have been called collectively the middle
vena cava, giving rise to a small branch, which projects cervical ganglia. These ganglia include macroscopi-
to the junction of the superior vena cava and right tally visible ganglia as well as ganglia that could only
atrium, and then joins with other cardiopulmonary be identified microscopically.
nerves posterior to the great vessels. The right dorsal Microscopic examination of the stellate ganglia re-
medial and dorsal lateral cardiopulmonary nerves fre- vealed that they are relatively long structures that vary
quently unite to become 1 large nerve at the level of in shape, with medial enlargements at their superior
the tracheal bifurcation. The larger of these 2 nerves, ends (Fig. 6 and 7B). Neuronal cell bodies in the stel-
or a single nerve if they unite, courses lateral to the late ganglia and the larger of the middle cervical gan-
trachea, anterior to the right main bronchus and right glia are organized in clusters separated by bundles of
pulmonary artery, and posterior to the root of the aorta. axons (Fig. 7D). The dorsal ansa, which arises from the
From this region, this relatively large nerve courses cranial end of the stellate ganglion, frequently con-
onto the heart along the left main coronary artery as tains 1 or more ganglia, and in some instances is com-
the left coronary cardiac nerve (Fig. 5). The left coro- posed of a continuous band of neuronal cell bodies.
nary cardiac nerve then separates into branches that Where the dorsal and ventral ansae unite superome-
course along the circumflex and left anterior descend- dially there is a ganglion, a middle cervical ganglion,
ing coronary arteries and onto the adjacent epicardi- which varies in size and shape. Histologic examination
um. The left stellate, left dorsal medial and left dorsal of the thoracic sympathetic trunks revealed that they
‘lateral cardiopulmonary nerves course posterior to the contain a number of relatively small ganglia located in
subclavian artery, lateral to the trachea and posterior the narrow neural trunks connecting the larger seg
to the aortic arch, to connect with nerves from the right mental ganglia (Fig. 6). Connective tissue makes up the
side posterior to the main pulmonary artery. The left remaining part of the trunks to give the gross anatomic
ventral cardiopulmonary nerve is the only nerve that appearance of large band-like structures (Fig. 1 and 2).
consistently courses anterior to the aortic arch. Mediastinal ganglia are present along the courses
The right and left cardiopulmonary nerves inter- of the cardiopulmonary nerves, where there are visi-
connect in the mediastinum at the base of the heart as ble swellings, where branches arise and along nerves
dorsal and ventral cardiopulmonary plexuses. The where there are no visible swellings (Fig. 7C). Many of
dorsal plexus is located dorsal to the aortic arch and these ganglia are found in the sympathetic cardiopul-
pulmonary artery along the anterior surface of the tra- monary nerves near the middle cervical and stellate
chea, near its bifurcation (Fig. 5B). All right and left ganglia (Fig. 4 and 7A). In some instances, apparent
cardiopulmonary nerves, except the left ventral car- swellings along the courses of cardiopulmonary
diopulmonary nerve, join to form the dorsal plexus. nerves did not contain neurons on histologic examina-
Many small nerves arise from this plexus to innervate tion. Mediastinal ganglia are also located along the
the adjacent atria, aorta, pulmonary artery, pulmonary course of the cardiopulmonary nerves adjacent to the
veins and trachea. The right dorsal lateral cardiopul- heart or in the cardiac nerves on the heart (Fig. 7F). No
monary nerve courses through the dorsal plexus as a large, consistently located ganglion is found in either
relatively distinct major nerve. Left cardiopulmonary the ventral or dorsal cardiopulmonary plexuses.
nerves and nerves arising from the dorsal plexus inter-
connect with the right dorsal lateral cardiopulmonary Discussion
nerve to form the left coronary cardiac nerve (Fig. 5). Recent anatomic and physiologic studies of cardiac
The left lateral cardiac nerve, which originates from innervation in dogs,13-*5J-25 cats,*4 monkeys*4 and ba-
the dorsal plexus, passes laterally, posterior to the left boons12 show that there are specific cardiac nerves,
333 HUMAN EXTRINSIC CARDIAC NERVES
each of which, when stimulated, producesrelatively
specific cardiovascularresponses.One of the striking
features of the anatomy described in thesestudies is
the relative similarity in the anatomic organizationof
the cardiac nerves and ganglia among species.These
descriptions of cardiac nerves and ganglia in mam-
mals differ markedly from previous,somewhatincon-
sistent,descriptionsof human cardiac innervation.l-I1
The present study shows that the anatomy of the
innervation of the human heart is more similar to that
of baboon@ than to that of dogs,13-15 cats14or mon-
keys,‘4although similarities are also presentwith re-
spectto theseother species.The nomenclatureusedto
describe the cardiac innervation of the baboon was
used,with minor modifications, to describethe anato-
my of the human cardiac nervesand ganglia. In both
humans and baboons,12the superior cervical ganglia
arelarge swellings at the baseof the skull (Fig. 1 and 2)
that are connected with the cervical sympathetic
trunks, the lower halves of which contain a variable
number of ganglionic swellings. Also, the thoracic
sympathetictrunks are wide, flat neural structuresthat
Cervical Sympathetic Trunk n ’ , n Vscebral
FIGURE 8. CompoeM drawing derived from eevefal hktokgio eecllona of the left middle cervkal and etellato ganglia ot embalmed cadaver
11 (Fig.- 35). Arowe Indicate the kcatkne ot cluelers of neuron8 and Borden /hea the extent ol the connecttve tlssue sheathe. Note the
thickness ot the connective ttssue sheathe. CPN = cardiopulmonary nerve.
end superiorly as relatively indistinct stellate ganglia.
In both humans and baboons,sympathetic cardiopul-
monary nerves arise from the stellate ganglia and the
inferior halves of the cervical sympathetic trunks to
join with vagal cardiopulmonary nerves arising from
the vagi and recurrent laryngeal nerves. Together,
thesenervesinterconnectto form distinct ventral and
dorsalcardiopulmonary plexuses,which, in turn, proj-
ectmajor cardiacnervesontothe heart aswell assmall
nerves to the adjacent atria. No cardiopulmonary
nerves arise from the superior cervical ganglia, the
superior half of the cervical sympathetictrunks or the
thoracic sympathetic trunks inferior to the stellate
ganglia in either man or baboons.In humans, in con-
trast to baboons,12no direct connectionsbetween the
middle cervical or stellate ganglia and the thoracic
vagi are found. Vagal connectionswith the sympathet-
ic cardiopulmonarynervesoccuralmost exclusively at
or below the level of the recurrent laryngeal nerves.
Small cardiopulmonary nerves,which arise from the
vagi posterior to the pulmonary hila, coursemedially
to the heart in humans. In baboons,similar nerves
FIWRE 7. photomkrographs ot human Wrack and cervical sympathetk ganglla and cardiopulmonary nerves (CPNs). Cresyl vkkt stain.
A, lett middle cervkal and small medlaetlnal ganglia (arrows) associated with sympathetic CPNs (embalmed cadaver 2). Callbratkn bar =
2 mm. S, kngltudlnel sectkn through the mlddle of a lett stellate gangllon (embalmed cadaver 9). Callbratlon bar = 2 mm. C, left
medtastlnal ganglia (arrows) kcated along the course of a CPN (embalmed cadaver 9). Callbratlon bar = 1 mm. D, hlgh magnltkatkn ot
the boxed area ot the lefl stellate gangllon shown In panel 6. Callbratkn bar = 0.5 mm. E, hlgh magnltlcatlon ot the boxed area of the medl-
astlnal ganglkn shown In panel C. Note blnucleate neuron. Callbratlon bar = 50 pm, F, small gangllon (arrow) located adjacent to the aorta
at the orlgln ot the lett maln coronary artery (embalmed cadaver 10). Callbratlon bar = 0.5 mm.
308 HUMAN EXTRINSIC CARDIAC NERVES
have been shown to have significant cardiac effectsI
In monkeyd4 there are more interconnections be-
tween the vagi and the middle cervical ganglia and
cervical sympathetictrunks than in either humans or
baboons, even though the cardiopulmonary nerves
originate from the sameregionsof the vagi and sympa-
thetic trunks.
The stellate and secondthoracic ganglia are sepa-
rate ganglia joined by relatively thin neural trunks,
which frequently contain groupsof neurons[Fig. 61,as
in monkeys.14Thus, the band-like grossanatomic ap-
pearanceof the thoracic sympathetictrunks in humans
is, to a large extent, a result of connective tissue
sheathsthat obscurethe ganglionicswellings.The stel-
late and middle cervical ganglia contain fewer neu-
rons than expectedfor their size becauseof their con-
nective tissue sheaths and the fact that there are
regionswithin the gangliathat arecomposedprimarily
of axons (Fig. 7D).
A variable number of ganglia of different sizesare
locatedin the caudal halves of the cervical sympathet-
ic trunks in man and are homologousto the multiple,
small middle cervical gangliafound in the sameregion
of the monkey.14Previousreportsin humanshave giv-
en different names to the visible ganglionic swellings
in this region, such as intermediate cervical gangli-
on,4,26accessorymiddle cervical ganglion,5vertebral
ganglion2~5~6*10and low middle cervical ganglion.26Be-
causeno special featuresappearto justify giving these
larger visible ganglia separatenames,and especially
as their sizes and positions vary, we have named all
ganglia in this region, including the microscopically
identified ganglia,middle cervical ganglia,in keeping
with terminology describing the anatomy of the mon-
key.l4 Neurons are present in the large superior cer-
vical sympathetic ganglia, the cranial halves of the
cervical sympathetic trunks, and throughoutthe sym-
pathetic trunks. Consideringthe widespread distribu-
tion of neuronswithin the cervical sympathetictrunks
between the superior cervical and stellate ganglia, it
may be appropriateto considerthesealso asrepresent-
ing middle cervical ganglia.
Many small ganglia exist along the coursesof the
cardiopulmonary nerves in humans, similar to the
anatomy of mediastinal ganglia of dogs21-23 and mon-
keys.14Although reportsin man have described sym-
patheticgangliain cervical rami communicantesz7and
in cardiopulmonary nerves adjacent to the heart,s
small mediastinal ganglia have not been describedas
occurring so extensively along cardiopulmonary
nerves. The “cardiac ganglion” (Wrisberg’sganglion)
is usually described as being located along the inner
curvature of the arch of the aortaagAs there are many
ganglia in the mediastinum immediately superior to
the heart,none of which are consistentlylocatedalong
the inner curvature of the aortic arch, identification of
any one of theseasWrisberg’sganglion appearsnot to
be warranted.The locationsof the cardiac sympathetic
efferent postganglionicneuronal cell bodiesinnervat-
ing the human heart are unknown, but when one takes
into accountthe large number of mediastinal and mid-
dle cervical ganglia and the location of sympathetic
postganglionic neuronal cell bodies in other spe-
cies,lQQ3 it seems likely that many of them will be
found outside the stellate ganglia. Other investiga-
tors’-4@*7~s~lO
have described the cardiac plexuses as
containing numerous small interconnections, and
found that the ventricles are innervated by extensions
of theseplexuses,not by relatively large, discretecar-
diac nerves. Our findings may differ from those of
previous reports because combined anatomic and
physiological studies of cardiac nerves in subhuman
mammald2-15921-25 were not available as a frame of
reference,nonembalmed autopsymaterial was rarely
used,and little, if any, histologic examinationwas car-
ried out in theseprevious human anatomic studies.
Electrical stimulations of individual cardiopulmo-
nary nervesand ganglia in dogs,13-1s.2s monkeysi and
baboons12confirm that these structureshave specific
effectson cardiac chronotropism,dromotropism and/
or a regional inotropism. Few studiesin humans have
attemptedto investigatethe effects of stimulating car-
diac nerves or ganglia.26-28 Recently, however, it was
shown that electrical stimulation of individual cardiac
nerves in patients undergoingcoronary artery bypass
surgery can produce specific cardiac effects.zgThe
complexity of the anatomy of human cardiac nerves
and ganglia must be taken into account in order to
study their physiology.
Acknowledgment: We gratefully acknowledgethe
help and encouragementof Dr. D.G. Gwyn and Dr.
J.H. Cooper. We thank the pathologistsand morgue
attendants of the MacKenzie Diagnostic Center for
their cooperation.We also thank Kathleen McCleave
and Camille Haddad for their technical assistanceand
Joan Livingston and Cheryl Forbes for their typing
assistance.The artwork is by Bev Ransomand by the
Audio-Visual Department of the Faculty of Medicine,
Dalhousie University.
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