Male reproductive system
1285.e1
67
RETPAHC
Erectiledysfunctionistheinabilitytoachievetumescencedespiteade-
quatesexualstimulation.Thepathophysiologyoferectiledysfunction
iscomplex,andinvolvesvascular,neurogenicandpsychogeniccompo-
nents.Commonaetiologiesunderlyingadiagnosisoferectiledysfunc-
tioninclude:inabilitytorelaxthecavernoussmoothmuscle;arterial
insufficiencyduetoatheromatousdisease;neurogenicdysfunctiondue
todiabetesorsurgery;andradiationtherapy.
Aprolongederectionthatfailstosubsideafterejaculationiscalled
priapism.Priapismcan,broadly,beclassifiedasbeinghigh-flow(non-
ischaemic)orlow-flow(ischaemic).Low-flowpriapism,whichisdue
toimpairedeffluxofvenousbloodfromthepenis,ischaracterizedby
significantpainandisamedicalemergency.Ifuntreated,priapismleads
toischaemiaofthecorporalsmoothmuscleandirreversibleerectile
dysfunction.Theaetiologyoflow-flowpriapismincludesconditions
suchassickle-celldiseaseandleukaemia,andtheuseofintracavernosal
injectiontherapywithmedicationssuchasprostaglandin.Incontrast,
high-flowpriapismistheconsequenceofexcessivebloodflowintothe
penis,usuallysecondarytoafistulabetweenthecavernousarteriesand
sinusoidalspaces.


MAlE REPRoduCTiVE sysTEM
1286
8
NoiTCEs
Fig. 76.24 The blood vessels
and nerves of the perineal
Perineal artery, posterior scrotal branches
region and external genitalia
in the adult male. The fat
body of the ischio-anal fossa
Bulbospongiosus has been removed and
Posterior scrotal nerves
gluteus maximus has been
incised in order to expose the
Dorsal artery of penis Ischiocavernosus course of the pudendal nerve
and internal pudendal artery.
(With permission from
Artery of bulb of penis
Superficial external Waschke J, Paulsen F (eds),
anal sphincter Sobotta Atlas of Human
Dorsal nerve of penis Anatomy, 15th ed, Elsevier,
Perineal artery Urban and Fischer. Copyright
2013.)
Perineal branches, posterior
cutaneous nerve of thigh
Superficial transverse
perineal muscle
Perineal nerve
Posterior cutaneous
nerve of thigh
Dorsal nerve of penis
Dorsal nerve of penis
Inferior rectal nerves
Internal pudendal
Perineal nerve artery and vein
Sacrospinous ligament
Pudendal nerve Inferior rectal artery
Internal pudendal artery
Posterior cutaneous nerve of thigh
Sacrotuberous ligament
Gluteus maximus Levator ani
Anococcygeal ligament Anococcygeal nerves
Dorsal arteries Perineal arteries layersofthescrotalwallexcepttheskin.Thetestesaresuspendedby
thespermaticcordswithinthesecompartments.
ThedartoslayerofsmoothmusclesiscontinuouswithColles’,
Scarpa’sandthedartosfasciaofthepenis.Theexternalspermatic,cre-
mastericandinternalspermaticlayersofthescrotumarecontinuous
withthecorrespondinglayersinthespermaticcord,andarisefromthe


aponeurosesofexternalobliqueandinternalobliqueandthetransver-
salisfasciaoftheabdominalwall,respectively.Thetestisisalsofixed
tothescrotalwallatitslowerpolebyafibrousbandoftissueknown
asthegubernaculum.
Cavernous artery
Bulbourethral artery Developmental anomalies of the scrotum Congenitalagenesis
ofthescrotumisaveryrareanomalycharacterizedbytheabsenceof
thescrotum;norugaearenotedintheperineum(Yilmazetal2013).
Penoscrotaltransposition,inwhichthescrotumislocatedsuperiorand
Circumflex artery anteriortothepenis,isaconsequenceofabnormalgenitaltubercle
developmentandisassociatedwithhypospadias,renalagenesisand
dysplasia,andimperforateanus.
Vascular supply and lymphatic drainage
Arterialsupplytothescrotumincludesexternalpudendalbranchesof
thefemoralartery,scrotalbranchesoftheinternalpudendalartery,and
Fig. 76.25 The arterial blood supply to the body of the penis. acremastericbranchfromtheinferiorepigastricartery.Ofnote,the
anteriorscrotalwallissuppliedprimarilybybranchesoftheexternal
pudendalartery,whichrunparalleltotherugaeanddonotcrossthe
raphe;thisanatomicaldetailisimportantforplanningappropriate
incisionsduringscrotalsurgery.


1287
67
RETPAHC
Key references
Scrotallymphaticsdonotcrossthemedianraphe.Therefore,lym-
phaticdrainagefromthescrotumisalwaysroutedtotheipsilateral
superficialinguinalnodes.
Innervation
Pelvic
Thescrotumisinnervatedbytheilioinguinalnerve,thegenitalbranch
parasympathetic
roots ofthegenitofemoralnerve,twoposteriorscrotalbranchesoftheperi-
nealnerve,andtheperinealbranchoftheposteriorfemoralcutaneous
nerve(seeFig.76.24).Innervationoftheanteriorthirdofthescrotum
isprimarilyderivedfromthefirstlumbarspinalsegment,bywayofthe
ilioinguinalandgenitofemoralnerves(seeFig.76.2).Innervationofthe
posteriortwo-thirdsofthescrotumisprimarilyderivedfromthethird
sacralspinalsegment,viatheperinealandposteriorfemoralcutaneous
nerves.Inordertoanaesthetizethescrotum,spinalanesthesiamust,
therefore,beinjectedhigherthantheL1level.
Obturator
nerve
Bonus e-book images and table
Hypogastric
nerves
Fig. 76.1 A, Reference curves for mean testicular volume measured
by ultrasound. B, Enlargement of the reference curves for mean
Rectum
testicular volume measured by ultrasound.
Prostate
Left inferior Fig. 76.12 Gangrene of the appendix testis secondary to torsion in
hypogastric an 8-year-old child.
Dorsal sensory plexus
nerve Table 76.1 Volume of the left and right testis, as measured by


Pudendal nerve ultrasound in boys between infancy and adolescence.
Cavernous nerves
Posterior scrotal nerve
Fig. 76.26 The nerve supply to the penis. The corpus cavernosum of
penis receives both a parasympathetic and a sympathetic innervation
from the cavernous nerves. It should be noted that, in life, multiple
cavernous nerves emanate from the prostatic plexus and intertwine with
both dorsal sensory nerves. The afferent fibres from the glans pass via
the dorsal nerves of the penis and via the pudendal nerve. (Adapted from
Drake RL, Vogl AW, Mitchell A (eds), Gray’s Anatomy for Students,
2nd ed, Elsevier, Churchill Livingstone. Copyright 2010.)
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BeckEM,SchlegelPN,GoldsteinM1992Intraoperativevaricoceleanatomy: anastomoses with the testicular blood supply, and clinical implications in
amacroscopicandmicroscopicstudy.JUrol148:1190–4. disease settings.
The first large, detailed, intraoperative study of spermatic cord anatomy in NguyenHT,EtzellJ,TurekPJ1996Normalhumanejaculatoryductanatomy:
men with varicoceles. astudyofcadavericandsurgicalspecimens.JUrol155:1639–42.
BraithwaiteJL1952Thearterialsupplyofthemaleurinarybladder.BrJ Gross and microscopic evaluation of the human ejaculatory duct, correlating
Urol24:64–71. microarchitecture to function, and defining anatomical measurements for
A comprehensive description of the arterial supply of the human bladder, the diagnosis of ejaculatory duct obstruction.
including variations and anomalies. PraderA1966Testicularsize:assessmentandclinicalimportance.Triangle
GoldsteinM2012Surgicalmanagementofmaleinfertility.In:WeinA, 7:240–3.


KavoussiLR,NovickACetal(eds)Campbell-WalshUrology,10thed. A landmark paper introducing the use of a standardized orchidometer for
Philadelphia:Elsevier,Saunders,pp.648–87. measuring testicular volume, and discussing the relationship between
A definitve and unparallelled summary of the indications and operative abnormal testicular size and various underlying medical disorders.
techniques for operations involving the male reproductive tract. SilberSJ1989Roleofepididymisinspermmaturation.Urology33:
HarrisonRG,BarclayAE1948Thedistributionofthetesticularartery(inter- 47–51.
nalspermaticartery)tothehumantestis.BrJUrol20:57–66. An examination of epididymal tubule patency and natural pregnancy rates
A detailed anatomical study that significantly improved knowledge of the following vasoepididymostomy to epididymal tubules in the caput, corpus or
vascular supply of the human testis, with important implications for surgical cauda epididymis, with implications for sperm maturation during
techniques of orchidopexy. epididymal transit.
HofferAP1976Theultrastructureoftheductusdeferensinman.Biol TaguchiK,TsukamotoT,MurakamiG1999Anatomicalstudiesoftheauto-
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MacmillanEW1954Thebloodsupplyoftheepididymisinman.BrJUrol A histological evalaution of the various components of the pelvic plexus,
26:60–71. correlating nerve morphology with function, and demonstrating the presence
An impressive and definitive arteriographical evaluation of the arterial of communication between nerves from the right and left plexuses,
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CHAPTER
77
Female reproductive system
The female reproductive system consists of the lower genital tract (vulva mons becomes prominent with coarse hair and atrophies slightly after
and vagina) and the upper tract (uterus and cervix with associated menopause.
uterine (Fallopian) tubes and ovaries).
Labia majora
LOWER GENITAL TRACT
The labia majora are two prominent, longitudinal folds of skin that
extend back from the mons pubis to the perineum (Fig. 77.1B). They
VULVA
form the lateral boundaries of the vulva. Each labium has an external,
pigmented surface covered with hairs and a smooth, pink internal
The female external genitalia or vulva include the mons pubis, labia surface with large sebaceous follicles. Between these surfaces there is
majora, labia minora, clitoris, vestibule, vestibular bulb and the greater loose connective and adipose tissue, intermixed with smooth muscle
vestibular glands (Fig. 77.1). (resembling the scrotal dartos muscle), vessels, nerves and glands. The
subcutaneous layer consists of a superficial fatty layer similar to Camp-
Mons pubis
er’s fascia, and a deep membranous layer – Colles’ fascia – continuous
with Scarpa’s fascia of the anterior abdominal wall (p. 1069). The
The mons pubis is the rounded, hair-bearing area of skin and adipose uterine round ligament may end in the adipose tissue and skin in the
tissue over the pubic symphysis and adjacent pubic bone. Before anterior part of the labium. A patent processus vaginalis may also reach
puberty, the mons pubis is relatively flat and hairless, and the labia a labium. The labia join anteriorly to form the anterior commissure.
minora are poorly formed. Through adolescence and into adult life, the Posteriorly, they do not join, but instead merge into neighbouring skin,
A B Skin overlying
body of clitoris
Prepuce of clitoris
Glans clitoris
Frenulum Lateral fold Glans clitoris
Urethral opening Medial fold
Vestibule Opening of duct of Labium minus
(between labia minora) para-urethral gland
Hymen Vestibule
Labium minus
Vaginal opening


Opening of duct of Labium majus
greater vestibular gland
Fourchette
Posterior
commissure
(overlies
perineal body)
C D
Prepuce
of clitoris
Frenulum
Glans clitoris of clitoris Area of opening of the
duct of the para-urethral
gland
Labium
minus
Area of opening of the
External duct of the greater
urethral orifice vestibular gland
Vaginal opening
(introitus)
Fig. 77.1 A, The superficial features of the perineum. B, The labia majora and surrounding external genitalia. C, The labia minora. D, An inferior view of
the vestibule with the left labium minus pulled to one side to show the regions of the vestibule into which the greater vestibular and para-urethral glands
open. (A, With permission from Drake RL, Vogl AW, Mitchell A, Tibbitts R, Richardson P (eds), Gray’s Atlas of Anatomy, Elsevier, Churchill Livingstone.
Copyright 2008. B–D, With permission from Drake RL, Vogl AW, Mitchell A (eds), Gray’s Anatomy for Students, 2nd ed, Elsevier, Churchill Livingstone.
1288 Copyright 2010.)


Lower genital tract
1289
77
RETPAHC
ending near and almost parallel to each other. The connecting skin A
between them posteriorly forms a ridge, the posterior commissure, Ischiocavernosus Suspensory ligament
which overlies the perineal body and is the posterior limit of the vulva.
The distance between the posterior vulva and the anus is 2.5–3 cm and
is termed the ‘gynaecological’ perineum.
Labia minora
The labia minora are two small cutaneous folds, devoid of fat, that lie
between the labia majora. They extend from the clitoris obliquely
down, laterally and back, flanking the vaginal orifice. Anteriorly, each
labium minus bifurcates. The upper layer of each side passes above the
clitoris to form the hood or prepuce, while the lower layer passes below
the clitoris to form the frenulum of the clitoris (Fig. 77.1A–C). Seba-
ceous follicles are numerous on the medial surfaces. Sometimes, an
extra labial fold (labium tertium) is found on one or both sides between Bulbospongiosus Perineal body Superficial transverse perineal muscle
the labia minora and majora. Adhesions between the labia minora are
common in prepubertal girls and may predispose to urinary infections
(Leung and Robson 1992).
B
Vestibule Glans clitoris Skin Body of clitoris
(cross-section)
Corpus cavernosum
Corpora cavernosa
The vestibule is the cavity that lies between the labia minora. It contains
the vaginal and external urethral orifices and the openings of the two
greater vestibular (Bartholin’s) glands and of numerous mucous, lesser
vestibular glands. There is a shallow vestibular fossa between the vaginal
orifice and the frenulum of the labia minora. The posterior part of the
vestibule is a classic site where a fistulous opening of the rectum may
be located in girls born with an imperforate anus (Bill et al 1975).
Urethra
The urethra opens into the vestibule about 2.5 cm below the clitoris


and above the vaginal opening via a short, sagittal cleft with slightly
raised margins: the urethral meatus. The meatus is very distensible and
varies in shape; the aperture may be rounded, slit-like, crescentic or Bulb of vestibule Greater vestibular gland Crus of clitoris
in superficial pouch (attached part of
stellate. The ducts of the para-urethral glands (Skene’s glands) open on
corpus cavernosum)
each side of the lateral margins of the urethra.
C
Bulbs of the vestibule
Body of clitoris (unattached
The bulbs of the vestibule lie on each side of the vestibule (Fig. 77.2B). parts of corpora cavernosa)
They are two elongated masses of erectile tissue, 3 cm long, which flank Mons pubis
the vaginal orifice and unite anterior to it by a narrow commissura
bulborum (pars intermedia). Their posterior ends are expanded and are
in contact with the greater vestibular glands. Their anterior ends taper
and are joined by a commissure, and also to the clitoris by two slender
bands of erectile tissue. Their deep surfaces contact the inferior aspect
of the perineal membrane; superficially, each is covered posteriorly by
bulbospongiosus (Fig. 77.2A) (Delancey 2011).
Greater vestibular glands
(Bartholin’s glands)
The greater vestibular glands are homologues of the male bulbourethral
glands. They consist of two small, round or oval, reddish-yellow bodies
that flank the vaginal orifice, in contact with, and often overlapped by,
the posterior end of the vestibular bulb. Each opens into the vestibule
by a 2 cm duct, situated in the groove between the hymen and the
labium minus at approximately the 5 and 7 o’clock positions (Corton
Crus clitoris (attached part
2012) (see Fig. 77.1D). The glands are composed of tubulo-acinar of corpus cavernosum) Glans clitoris
tissue; the secretory cells are columnar and secrete a clear or whitish
Fig. 77.2 A, The muscles of the vestibule and clitoris. B, The erectile
mucus for lubrication during sexual arousal.


tissues of the clitoris. C, The erectile tissues of the clitoris, vestibule and
Clitoris greater vestibular gland with a surface anatomy overlay. (A–B, Adapted
with permission from Drake RL, Vogl AW, Mitchell A (eds), Gray’s
Anatomy for Students, 2nd ed, Elsevier, Churchill Livingstone. Copyright
The clitoris is an erectile structure, partially enclosed by the anterior 2010. C, With permission from Drake RL, Vogl AW, Mitchell A (eds),
bifurcated ends of the labia minora. It has a root, a body and a glans Gray’s Anatomy for Students, 2nd ed, Elsevier, Churchill Livingstone.
(see Fig. 77.2). The body can be palpated through the skin. It contains Copyright 2010.)
two corpora cavernosa, composed of erectile tissue and enclosed in
dense fibrous tissue, and separated medially by an incomplete fibrous
pectiniform septum. The fibrous tissue forms a suspensory ligament Vascular supply and lymphatic drainage
that is attached superiorly to the pubic symphysis. Each corpus caverno-
of the vulva
sum is attached to its ischiopubic ramus by a crus that extends from
the root of the clitoris. The glans clitoris is a small, round tubercle of
Arteries
spongy erectile tissue at the end of the body, connected to the bulbs of
the vestibule by thin bands of erectile tissue. It is exposed between the The arterial blood supply of the female external genitalia is derived
anterior ends of the labia minora. Its epithelium has high cutaneous from the superficial and deep external pudendal branches of the femoral
sensitivity, important in sexual responses. Congenital absence of the artery superiorly and the internal pudendal artery inferiorly on each
clitoris is very rare. side (Figs 77.3A, 77.4).


FEmALE REPRoduCTivE sysTEm
1290
8
NoiTCEs
A
Common iliac vein Internal iliac vein
Common iliac artery Median sacral artery
Superior vesical artery and vein
Internal iliac artery
Rectum, longitudinal
Ovarian artery
muscular layer
Umbilical artery, patent part Uterine artery
Ureter
Middle rectal artery
Right ovary and vein
Recto-uterine fold
Uterine tube, infundibulum
Uterine tube Left ureter
Round ligament of uterus
External iliac Uterine artery and
artery and vein vein
Vagina
Uterus
Middle rectal
Urinary bladder
artery and vein
Vaginal artery
Left ovary
Inferior vesical
Uterine artery,
artery
tubal branch
Levator ani
Bulb of vestibule
Inferior rectal vein
Round ligament of
uterus
Vaginal venous plexus
Vesical veins Internal pudendal artery and vein
B
Abdominal aorta Common iliac nodes
Iliolumbar artery
Ovarian artery and vein Superior rectal
artery and vein
Internal iliac node
Rectum
External iliac node
Genitofemoral nerve Uterine artery
and vein
Obturator nerve
Superior rectal
External iliac artery and vein node
Obturator artery and vein Vaginal artery
External iliac nodes, Internal pudendal
medial and lateral nodes artery
Inferior epigastric
artery and vein
Cord of umbilical
artery Recto-uterine
pouch
Round ligament of
uterus
Umbilical artery,
patent part
Levator ani
Round ligament of uterus
Uterus, intestinal surface
Superior vesical arteries
Ureter
Urinary bladder Round ligament of uterus
Median umbilical ligament Uterine tube, isthmus
Fig. 77.3 A, The vessels of the female pelvis: sagittal view. B, The nerves and lymphatics of the female pelvis: sagittal view. (With permission from
Waschke J, Paulsen F (eds), Sobotta Atlas of Human Anatomy, 15th ed, Elsevier, Urban & Fischer. Copyright 2013.)


Lower genital tract
1291
77
RETPAHC
Bulbospongiosus Fig. 77.4 The muscles,
External urethral orifice
vessels and nerves of
Internal pudendal artery, the female perineum:
Bulb of vestibule posterior labial branches inferior view. (With
Vein of bulb of Ischiocavernosus permission from
vestibule Waschke J, Paulsen F
Dorsal artery of (eds), Sobotta Atlas of
clitoris Human Anatomy, 15th
ed, Elsevier, Urban &
Perineal artery Urogenital diaphragm Fischer. Copyright
2013.)
Dorsal nerve of
clitoris
Superficial transverse
perineal muscle
Perineal branches,
posterior cutaneous
nerve of thigh
Posterior labial nerves
Inferior rectal nerves
Internal pudendal
artery and vein
Posterior cutaneous
Gluteus maximus
nerve of thigh
Inferior rectal artery
Perineal nerves
Pudendal nerves
Levator ani Anococcygeal nerves
External anal sphincter
Anococcygeal ligament
This blood supply to the labial fat must be carefully preserved during
VAGINA
vaginal surgery, e.g. in creating a Martius fat pad flap to repair a vesico-
vaginal fistula where blood supply has already been compromised by
The vagina is a fibromuscular tube lined by non-keratinized stratified
radiation or fibrosis (Delancey 2011).
epithelium. It extends from the vestibule (the opening between the
Veins labia minora) to the uterus. The upper end of the vagina surrounds the
Venous drainage of the vulval skin is via external pudendal veins to the vaginal projection of the uterine cervix. The anular recess between the
long saphenous vein. Venous drainage of the clitoris is via deep dorsal cervix and vagina is the fornix; the different parts of the fornix are given
veins to the internal pudendal vein, and via superficial dorsal veins to separate names, i.e. anterior, posterior and right and left lateral, but they
the external pudendal and long saphenous veins. are continuous (Fig. 77.6).
In the standing position, the vagina ascends posteriorly and superi-


Lymphatic drainage
orly, forming an angle of 60–70° with the horizontal plane (Corton
A meshwork of connecting vessels from the skin of the labia, clitoris 2012). The vagina forms an angle of over 90° to the uterine axis (see
and perineum join to form three or four collecting trunks that drain to Fig. 77.12); this angle varies with the contents of the bladder and
superficial inguinal nodes lying on the cribriform fascia covering the rectum. The apex of the vagina is directed posteriorly towards the ischial
femoral artery and vein; these nodes drain through the cribriform fascia spines. The width of the vagina increases as it ascends. The inner sur-
to the deep inguinal nodes lying medial to the femoral vein (Corton faces of the anterior and posterior vaginal walls are in contact with each
2012). The deep inguinal nodes drain via the femoral canal to the pelvic other, forming a transverse slit. The vaginal mucosa is attached to the
nodes (Table 77.1). The last of the deep inguinal nodes lies under the uterine cervix higher on the posterior cervical wall than on the anterior;
inguinal ligament within the femoral canal and is often called Cloquet’s the anterior wall is approximately 7.5 cm long and the posterior wall
node. Lymph vessels from the clitoris drain directly to the deep inguinal is approximately 9 cm long. The fibromuscular anterior wall of the
nodes, and direct clitoral lymphatics may pass to the internal iliac nodes vagina supports the base of the bladder in its middle and upper por-
(Fig. 77.3B). Lymph vessels in the perineum and lower part of the labia tions, and the urethra (which is embedded in it) inferiorly. The fibro-
majora drain to the rectal lymphatic plexus. muscular posterior wall of the vagina supports the rectum. The upper
quarter of the posterior vagina is separated from the rectum by the
Innervation
peritoneum of the recto-uterine pouch, and by moderately dense fibro-
muscular tissue (Denonvilliers’s fascia) in its middle half (Delancey


The main nerve supply of the vulva is the pudendal nerve (S2, 3 and 2011). In its lower quarter, it is separated from the anal canal by the
4) through its inferior rectal and perineal branches and the dorsal nerve fibromuscular perineal body. The upper part of the vagina is supported
of the clitoris (Table 77.2; Figs 77.4, 77.5). Shoja et al 2013). The laterally by levator ani, together with the transverse cervical, pubocervi-
sensory innervation of the anterior and posterior parts of the labium cal and uterosacral ligaments. Pubovaginalis provides a U-shaped mus-
majus differs. The anterior third of the labium majus is supplied by the cular sling around the mid-vagina. The lower vagina is surrounded by
ilioinguinal nerve (L1), the posterior two-thirds are supplied by the the skeletal muscle fibres of bulbospongiosus (see Fig. 77.4). As the
pudendal nerve through the posterior labial branches of the perineal ureters pass anteromedially to reach the fundus of the bladder, they
nerve (S3), and the lateral aspect is also innervated by the perineal pass close to the lateral fornices, where care must be taken to avoid
branch of the posterior cutaneous nerve of the thigh (S2). Vulvar nerves damage during hysterectomy (Fig. 77.7). As they enter the bladder, the
are susceptible to trauma and inflammation, leading to vulvar pain ureters are usually anterior to the vagina; at this point, each ureter is
syndromes or vulvodynia (Shoja et al 2013). crossed transversely by a uterine artery (Fig. 77.8).


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Table 77.1 Lymphatic drainage of the female genitalia
Structure Nodes
Vulva
Labia, clitoris, and perineum Superficial inguinal nodes
Perineum and lower labia majora Rectal lymphatic plexus
Clitoris Superficial inguinal nodes
Direct to deep inguinal nodes
Direct to internal iliac nodes
Vagina
Upper vagina Internal and external iliac nodes
Mid-vagina Internal iliac nodes
Lower vagina Superficial inguinal nodes
Uterus
Fundus Para-aortic nodes
Isthmus of tube and round ligament Superficial inguinal nodes
Body of uterus, including cervix External iliac nodes
Internal iliac nodes
Obturator nodes
Uterine (Fallopian) tubes Para-aortic nodes
Internal iliac nodes
Superficial inguinal nodes
Ovaries Para-aortic nodes


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T12 Fig. 77.5 The autonomic innervation of the
female reproductive system. (Based on Barucha
Paravertebral sympathetic chain AE 2006 Pelvic floor: anatomy and function.
L1 Neurogastroenterol Motil 18:507–19.)
L2
Aortic plexus
L3
L4
Lumbar splanchnic nerves
L5
Superior hypogastric plexus
Left hypogastric nerve
Right hypogastric nerve S1
S2
S3
Pelvic splanchnic nerves
S4
Uterovaginal plexus
S5
Pudendal nerve
Vesical plexus Inferior hypogastric plexus
Middle rectal plexus
Inferior rectal nerve
Perineal nerve
Anterior fornix Anterior lip Fig. 77.6 The cervix after it has been ruptured. It has no established function. The hymen
Cervical os of vagina of cervix Left vaginal wall and vaginal fornices, as may be imperforate; this is usually detected in adolescence.
seen via a Cusco’s Remnants of the duct of Gartner (embryologically, the caudal end
bivalve speculum. of the mesonephric duct) (see Figs 72.13, 72.19) are occasionally seen
protruding through the lateral fornices or lateral parts of the vagina and
may cause cysts (Gartner’s cysts). Prepubertal distal longitudinal folds
are common within the vagina and they disappear during puberty
(Altchek et al 2008).
Vascular supply and lymphatic drainage
Arteries
The arterial supply of the vagina is derived from the internal iliac arter-
ies by two median longitudinal vessels, the azygos arteries of the vagina,
which descend anterior and posterior to the vagina; they also supply
the vestibular bulb, fundus of the bladder, and adjacent part of the
rectum (see Fig. 77.3A). The uterine, internal pudendal and middle
rectal branches of the internal iliac artery may contribute to the blood
Right Posterior fornix Posterior lip of Cusco’s speculum
supply of the vagina.
vaginal wall of vagina cervix
Veins


The vagina opens externally via a sagittal introitus positioned below The vaginal veins, one on each side, arise from lateral plexuses that
the urethral meatus (see Fig. 77.1C). The size of the introitus varies; it connect with uterine, vesical and rectal plexuses and drain to the inter-
is capable of great distension during childbirth and, to a lesser degree, nal iliac veins. The uterine and vaginal plexuses may provide collateral
during sexual intercourse. The hymen is a thin fold of mucous mem- venous drainage to the lower limb.
brane situated just within the vaginal orifice. The internal surfaces of
Lymphatic drainage
the folds are normally in contact with each other and the vaginal
opening appears as a cleft between them. The hymen varies greatly in Vaginal lymphatic vessels link with those of the cervix, rectum and
shape and dimensions. When stretched, it is anular and widest poste- vulva. They form three groups but the regions drained are not sharply
riorly; it may also be semilunar and concave towards the mons pubis, demarcated. Upper vessels accompany the uterine artery to the internal
cribriform, fringed, absent or complete and imperforate. The hymenal and external iliac nodes; intermediate vessels accompany the vaginal
ring normally ruptures after first sexual intercourse, but can rupture artery to the internal iliac nodes; and vessels draining the vagina below
earlier during non-sexual physical activity. Small round carunculae the hymen, and from the vulva and perineal skin, pass to the superficial
hymenales (also known as carunculae myrtiformis) are its remnants inguinal nodes (see Table 77.1).


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Innervation
The lower vagina is supplied by the pudendal nerve (S2, S3 and S4).
The upper vagina is supplied by the pelvic splanchnic nerves (S2, S3
and, sometimes, S4) (see Fig. 77.5 and Table 77.2).
Abdominal aorta
Developmental anomalies of the vagina
Ovarian artery
Congenital anomalies of the vagina are vaginal agenesis, absent hymen,
transverse vaginal septum and persistent cloaca. Vaginal agenesis, in
the presence of other Müllerian duct anomalies and renal agenesis,
is termed Mayer–Rokistansky–Kuster–Hauser syndrome. An absent
hymen in patients with vaginal agenesis is associated with renal agen-
Uterine tube esis (Kimberley et al 2012). A congenital transverse septum may be
present within the vagina and manifests clinically in adolescence with
primary amenorrhoea and haematocolpos. Children with a persistent
cloaca have a congenital defect characterized by fusion of the rectum,
2 vagina and urethra into a single common channel that varies in length
from 1 to 7 cm.
Microstructure
Ovary
Uterus The vagina has an inner mucosal and an external muscular layer. The
1 Ureter
mucosa adheres firmly to the muscular layer. There are two median
Uterine artery longitudinal ridges on its epithelial surface: one anterior and the other
posterior. Numerous transverse bilateral rugae extend from these
3 vaginal columns, divided by sulci of variable depth, giving an appear-
ance of conical papillae. These transverse rugosities are most numerous
Bladder on the posterior wall and near the orifice; they increase under the influ-
ence of oestrogen during puberty and pregnancy, are especially well
developed before parturition, and decrease after the menopause (Corton
2012). The epithelium is non-keratinized, stratified, squamous similar
Vagina to, and continuous with, that of the ectocervix. After puberty, it thickens
and its superficial cells accumulate glycogen, which gives them a clear
appearance in histological preparations.
Fig. 77.7 The most common sites of ureteric injury during hysterectomy.


The vaginal epithelium does not change markedly during the men-
Key: 1, distal ureter at the level of the uterine artery; 2, dorsal to the
strual cycle, but its glycogen content increases after ovulation and then
infundibulopelvic ligament, near the pelvic bone; 3, intramural portion of
diminishes towards the end of the cycle. Natural vaginal bacteria, par-
ureter at the angle of the vaginal cuff.
ticularly Lactobacillus acidophilus, break down glycogen in the desqua-
mated cellular debris to lactic acid. This produces a highly acidic (pH
3) environment that inhibits the growth of most other microorganisms.
The amount of glycogen is less before puberty and after the menopause,
when vaginal infections are more common. There are no mucous
glands, but a fluid transudate from the lamina propria and mucus from
the cervical glands lubricate the vagina (Fig. 77.9). The muscular layers
are composed of smooth muscle and consist of a thick outer longitu-
dinal and an inner circular layer; the two layers are not distinct but are
Median sacral artery
Ovarian artery
Ovarian
vessels
E
Ureter
Ureter
Broad
ligament
Vaginal Uterine
artery artery
Branches of
anterior trunk
of internal
iliac artery
Fig. 77.9 Stratified squamous non-keratinizing epithelium (E) covering the
Fig. 77.8 The relationship of the ureter to the uterine and vaginal arteries. ectocervix and vagina. The cells of the middle and upper layers appear
(With permission from Drake RL, Vogl AW, Mitchell A (eds), Gray’s clear due to their glycogen content. (Courtesy of Mr Peter Helliwell and
Anatomy for Students, 2nd ed, Elsevier, Churchill Livingstone. Copyright the late Dr Joseph Mathew, Department of Histopathology, Royal
2010.) Cornwall Hospitals Trust, UK.)


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O
C O
B
R
S V
R
A
A B
B S
U
C V
I I
A
R
C D
Fig. 77.10 T2-weighted magnetic resonance images through the pelvis, demonstrating the normal female anatomy. A, Sagittal view. Abbreviations: A,
anus; B, bladder; C, cervix; R, rectum; S, pubic symphysis; V, vagina; *, endometrium; **, inner myometrium of uterus (also known as the junctional
zone); ***, outer myometrium of uterus. B–D, Axial views. B, Abbreviations: O, ovaries; R, rectum; *, endometrium, **, inner myometrium of uterus
(junctional zone); ***, outer myometrium of uterus. C, Abbreviations: B, bladder; C, cervix (which, from external to internal, has several layers, as seen on
T2-weighted images: a high-signal-intensity outer cervical stroma (contiguous with the outer myometrium); a low-signal-intensity inner cervical stroma
(contiguous with the inner myometrium); high-signal-intensity endocervical glands (contiguous with endometrium); and a very high-signal-intensity
endocervical canal (contiguous with the endometrial canal)); R, rectum. D, Abbreviations: A, anus; I, ischio-anal fossa; S, pubic symphysis; U, urethra; V,
vagina.
connected by oblique interlacing fibres. The longitudinal fibres are In the adult nulliparous state, the cervix usually tilts forwards relative
continuous with the superficial muscle fibres of the uterus. A layer of to the axis of the vagina (anteversion), and the body of the uterus tilts
loose connective tissue, containing extensive vascular plexuses, sur- forwards relative to the cervix (anteflexion) (see Fig. 77.12). In 10–15%
rounds the muscle layers. of women, the whole uterus leans backwards at an angle to the vagina
and is said to be retroverted. A uterus that angles backwards on the


cervix is described as retroflexed.
UPPER GENITAL TRACT
Body
UTERUS
The body of the uterus is pear-shaped and extends from the fundus
The uterus is a thick-walled, muscular organ situated in the pelvis superiorly to the cervix inferiorly. The uterine tubes enter the uterus on
between the urinary bladder and the rectum (Figs 77.10–77.12). It lies both sides at the uterine cornua. The round and ovarian ligaments are
posterior to the bladder and uterovesical space, and anterior to the inferoanterior and inferoposterior, respectively, to each cornu. The
rectum and recto-uterine pouch; it is mobile, which means that its posi- dome-like fundus is superior to the entry points of the uterine tubes
tion varies with distension of the bladder and rectum. The broad liga- and covered by peritoneum that is continuous with that of neighbour-
ments are lateral. ing surfaces. The fundus is in contact with coils of small intestine and,
The uterus is divided structurally and functionally into two main occasionally, by distended sigmoid colon. The lateral margins of the
regions: the muscular body of the uterus (corpus uteri) forms the upper body are convex; on each side, their peritoneum is reflected laterally to
two-thirds, and the fibrous cervix (cervix uteri) forms the lower third. form the broad ligament, which extends as a flat sheet to the pelvic wall


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Table 77.2 Innervation of the female genitalia
Origin Proximal course Distal course Organ Function
Parasympathetic S2–S4 Pelvic splanchnic nerves Pelvic ganglia Uterine tube Vasodilation
Uterus
Cavernous (deep, cavernosal) nerves of clitoris Vagina Transudation
Clitoris Erection
Sympathetic T12, L1–L2 Superior mesenteric and renal plexus Ovarian plexus Ovary Vasoconstriction
Superior hypogastric plexus Hypogastric nerve
↓
Inferior hypogastric plexus
↓
Uterovaginal plexus (Frankenhäuser’s ganglion) Uterine tube, uterus, upper vagina Contraction
Somatic S2, 3, 4 Pudendal nerve Dorsal nerve of clitoris Clitoris
Pudendal nerve Posterior labial nerves Lower vagina Contraction
Labia majora
Ischiocavernosus
Bulbospongiosus
(With permission from Shoja MM, Sharma A, Mirzaya N, Groat C, Watanabe K, Loukas M and Tubbs RS. Neuroanatomy of the Female Abdominoplevic Region: A Review with Application to Pelvic Pain Syndromes. Clinical
Anatomy 26:66–76, 2013.)


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A
Axis of
uterine body Axis of vagina
Ovary and
uterine tube
Round ligament Recto-
of uterus uterine pouch
Uterus
Rectum
Vesico-uterine
pouch Vaginal
vault
Bladder
Vagina Axis of cervix
Pubis
Greater Angle of
Urethra vestibular gland anteflexion
Clitoris Bulb of vestibule
Angle of anteversion
B
Fig. 77.12 Angles of anteflexion and anteversion. (With permission from
Drake RL, Vogl AW, Mitchell A (eds), Gray’s Anatomy for Students, 2nd
ed, Elsevier, Churchill Livingstone. Copyright 2010.)
size of the uterus until approximately 7 years of age, when there is
Ureter Recto-uterine greater enlargement of the body of the uterus than the cervix.
fold
Broad ligament Developmental anomalies of the uterus There may be failure
Round ligament in fusion of the paramesonephric (Müllerian) ducts, which results in a
of uterus uterus that is not pear-shaped. There may only be a septum (septate
Inferior uterus) or partial clefting of the uterus (bicornuate uterus); the most
epigastric artery extreme example is a septate vagina, two cervices and two discrete uteri,
each with one uterine tube (uterus didelphys) (Minto et al 2001) (see
Fig. 72.15).
Lateral umbilical fold
Cervix
Medial umbilical fold Recto-uterine
The adult, non-pregnant cervix is narrower and more cylindrical than the
pouch
body of the uterus and is typically 2.5 cm long. The upper end commu-
nicates with the uterine body via the internal os, and the lower end opens
Median umbilical fold
into the vagina at the external os (see Fig. 77.6). In nulliparous women,
Vesico-uterine pouch
the external os is usually a circular aperture, whereas, after childbirth, it
is a transverse slit. Two longitudinal ridges, one each on its anterior and


Fig. 77.11 A, Anatomical relations of the female genital tract, bladder and posterior walls, give off small, oblique, palmate folds that ascend later-
rectum. B, Pelvic peritoneal reflections, showing the broad ligament and
ally like the branches of a tree (arbor vitae uteri); the folds on opposing
its contents. (A, With permission from Drake RL, Vogl AW, Mitchell A,
walls interdigitate to close the canal. The narrower isthmus forms the
Tibbitts R, Richardson P (eds), Gray’s Atlas of Anatomy, Elsevier,
upper third of the cervix. Although unaffected in the first month of
Churchill Livingstone. Copyright 2008. B, With permission from Drake RL,
pregnancy, it is gradually taken up into the uterine body during the
Vogl AW, Mitchell A (eds), Gray’s Anatomy for Students, 2nd ed, Elsevier,
second month to form the ‘lower uterine segment’ (see below). In non-
Churchill Livingstone. Copyright 2010.)
pregnant women, the isthmus undergoes menstrual changes, although
these are less pronounced than those occurring in the uterine body. The
external end of the cervix enters the upper end of the vagina, thereby
(see Figs 77.11B, 77.15). The anterior surface of the uterine body is dividing the cervix into supravaginal and vaginal parts. The supravaginal
covered by peritoneum reflected on to the bladder at the uterovesical part is separated anteriorly from the bladder by cellular connective tissue:
fold (see Fig. 77.14). This normally occurs at the level of the internal the parametrium, which also passes to the sides of the cervix and laterally
os, the most inferior margin of the body of the uterus. The vesico- between the two layers of the broad ligaments.
uterine pouch, between the bladder and uterus, is obliterated when the
bladder is distended, but may be occupied by small intestine when the Peritoneal folds and ligaments of the pelvis
bladder is empty. The posterior surface of the uterus is convex trans-
versely. Its peritoneal covering continues down to the cervix and upper The uterus is connected to a number of ‘ligaments’. Some are true liga-


vagina, and is then reflected back to the rectum along the surface of the ments, in that they have a fibrous composition and provide support to
recto-uterine pouch, which lies posterior to the uterus (see Fig. 77.16). the uterus; some provide no support to the uterus; and others are simply
The sigmoid colon, and occasionally the terminal ileum, lie posterior folds of peritoneum.
to the uterus.
The cavity of the uterine body usually measures 6 cm from the Peritoneal folds
external os of the cervix to the wall of the fundus and is flat in its The parietal peritoneum is reflected over the upper genital tract to
anteroposterior plane (Salardi et al 1985). In coronal section, it is tri- produce anterior (uterovesical), posterior (rectovaginal) and lateral
angular, broad above where the two uterine tubes join the uterus, and peritoneal folds. The lateral folds are commonly called the broad liga-
narrow below at the internal os of the cervix. There is no change in the ments (Fig. 77.13).


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Round ligament Ovary Uterus Uterus Recto-uterine pouch Round ligament External iliac artery
Broad ligament Rectum Uterine tube
Rectum Uterine tube Broad ligament
Fig. 77.13 A laparoscopic view of the uterus. The patient is tilted head
Fig. 77.15 A laparoscopic view of the broad ligament.
down so that the small bowel is moved away from the uterus to give this
view.
Catheter balloon in bladder Uterovesical fold Ovary Uterus Recto-uterine pouch Uterosacral ligament
Uterine tube Uterus Rectum Uterine tube Sigmoid colon descending Ovarian vessels
Fig. 77.14 A laparoscopic view of the anterior part of the pelvis, showing to rectum
the uterovesical fold anterior to the uterus. Fig. 77.16 A laparoscopic view of the posterior pelvis, showing the
recto-uterine pouch (pouch of Douglas) with the sigmoid colon
descending towards the rectum.
uterovesical and rectovaginal folds
The anterior, or uterovesical, fold consists of peritoneum reflected on
to the bladder from the uterus at the junction of its cervix and body Mesosalpinx
(Fig. 77.14). The posterior or rectovaginal fold extends lower than the The mesosalpinx is attached above to the uterine tube and posteroin-
anterior fold and consists of peritoneum reflected from the posterior feriorly to the mesovarium (see Fig. 77.17). Superior and laterally, it is
vaginal fornix on to the front of the rectum, thereby creating the deep attached to the suspensory ligament of the ovary; medially, it is attached
recto-uterine pouch (pouch of Douglas). The recto-uterine pouch is to the ovarian ligament. The fimbria of the tubal infundibulum projects
bounded anteriorly by the uterus, supravaginal cervix and posterior from its free lateral end. Between the ovary and uterine tube, the mes-
vaginal fornix; posteriorly, by the rectum; and laterally, by the uterosac- osalpinx contains vascular anastomoses between the uterine and


ral ligaments. ovarian vessels, the epoophoron and the paroophoron. The meso-
varium projects from the posterior aspect of the broad ligament, of
Broad ligament which it is the smaller part. It is attached to the hilum of the ovary and
The lateral folds, or broad ligaments, extend on each side from the carries vessels and nerves to the ovary.
uterus to the lateral pelvic walls, where they become continuous with
the peritoneum covering those walls (Figs 77.15, 77.17–77.18). The Mesometrium
upper border is free and the lower border is continuous with the peri- The mesometrium is the largest part of the broad ligament, and extends
toneum over the bladder, rectum and side wall of the pelvis. The borders from the pelvic floor to the ovarian ligament and uterine body. The
are continuous with each other at the free edge via the uterine fundus, uterine artery passes between its two peritoneal layers typically 1.5 cm
and diverge below near the superior surfaces of levatores ani. A uterine lateral to the cervix; it crosses the ureter shortly after its origin from the
tube lies in the upper free border on either side. The broad ligament is internal iliac artery and gives off a branch that passes superiorly to the
divided into an upper mesosalpinx, a posterior mesovarium and an uterine tube, where it anastomoses with the ovarian artery (Fig. 77.19).
inferior mesometrium. Between the pyramid formed by the infundibulum of the tube, the


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Fig. 77.17 Pelvic peritoneal
reflections, demonstrating the
broad ligament and its contents.
Ovarian vessels (With permission from Drake RL,
Vogl AW, Mitchell A (eds),
Ureter
Gray’s Anatomy for Students,
2nd ed, Elsevier, Churchill
Livingstone. Copyright 2010.)
Broad ligament
Suspensory ligament
of ovary
Mesovarium
Deep inguinal ring
Round ligament
Sagittal section of broad ligament
of the uterus
Mesosalpinx
Uterine
Inguinal canal Ovary
tube
Ligament of ovary Mesovarium
Superficial inguinal ring Broad
ligament
Ureter
Labium majus
Uterine artery
Rectum
Fundus of uterus
Ureter
Uterine tube, fimbriae
Vermiform appendix
Ovarian artery and vein
Caecum
Suspensory ligament of
ovary (infundibulopelvic
ligament)
Uterine tube, ampulla Uterine tube, infundibulum
Uterine tube, ampulla
Ovary, medial surface
Mesosalpinx
Mesovarian border
Uterine tube, isthmus
Ligament of ovary
Broad ligament of uterus
Round ligament of uterus
Vesico-uterine pouch
Medial umbilical fold
Urinary bladder
Uterus, vesical surface Median umbilical fold
Fig. 77.18 The ovaries and broad ligament: superior view with the uterus lifted away from the bladder. (With permission from Waschke J, Paulsen F
(eds), Sobotta Atlas of Human Anatomy, 15th ed, Elsevier, Urban & Fischer. Copyright 2013.)
upper pole of the ovary, and the lateral pelvic wall, the mesometrium neither attachment nor extension to the caudal labium (the homologue
contains the ovarian vessels and nerves lying within the fibrous suspen- of the hemiscrotum) (Attah and Hutson 1991). Near the uterus, the
sory ligament of the ovary (infundibulopelvic ligament). This ligament round ligament contains a considerable amount of smooth muscle but


continues laterally over the external iliac vessels as a distinct fold. The this gradually diminishes and the terminal portion is purely fibrous.
mesometrium also encloses the proximal part of the round ligament of The round ligament also contains striated muscle, blood vessels, nerves
the uterus, as well as smooth muscle and loose connective tissue. and lymphatics. The latter drain the uterine region around the entry of
the uterine tube to the superficial inguinal lymph nodes.
Ligaments of the pelvis
In the fetus, a projection of peritoneum (processus vaginalis) is
The ligaments of the pelvis consist of the round, uterosacral, transverse carried with the round ligament for a short distance into the inguinal
cervical and pubocervical ligaments. canal. This is generally obliterated in adults, although it is sometimes
patent even in old age. A patent processus vaginalis in the inguinal canal
Round ligament in females is often referred to as the canal of Nuck; it may be asymp-
Each round ligament is a narrow smooth muscle band 10–12 cm long, tomatic or it may give rise to an inguinal hernia or hydrocele of the
which extends from the lateral cornu of the uterus through the broad canal of Nuck. In the canal, the ligament receives the same coverings as
ligament to enter the deep inguinal ring lateral to the inferior epigastric the spermatic cord, although they are thinner and blend with the liga-
artery (see Figs 77.13, 77.15, 77.18). Although conventionally described ment itself, which may not reach the mons pubis. The round and
as ending in the labium majus, a cadaveric dissection study found that, ovarian ligaments both develop from the gubernaculum and are
in girls, the round ligament ended just outside the external ring, with continuous.


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uterosacral, transverse cervical and horizontal axis of the upper vagina. The uterus and vagina are supported
pubocervical ligaments by the close interaction of the uterosacral and transverse cervical liga-
ments with the muscles of the pelvic floor, including the levatores ani
The uterosacral, transverse cervical and pubocervical ligaments are con-
and coccygei, the perineal membrane and the perineal body. The
densations of the visceral or endopelvic connective tissue that connect
support of the pelvic floor has been reviewed in detail by Delancey
the pelvic viscera to the side wall of the pelvis; they radiate like the
(2011) (see also Ch. 73).
spokes of a wheel around the hub of the cervix, providing it with con-
siderable support (see Figs 77.13–16) (Delancey 2011). The connective
Vascular supply and lymphatic drainage
tissue lateral to the uterus and the cervix – the parametrium – continues
down along the vagina as the paracolpium. The uterosacral ligaments
contain fibrous tissue and smooth muscle. They pass back from the Arteries
cervix and uterine body on both sides of the rectum, and are attached The arterial supply to the uterus comes from the uterine artery (see Fig.
to the anterior aspect of the sacrum. They can be palpated laterally on 77.19; Fig. 77.21), which arises as a branch of the anterior division of
rectal examination and can be felt as thick bands of tissue passing the internal iliac artery. From its origin, the uterine artery crosses the
downwards on both sides of the posterior fornix on vaginal examina- ureter anteriorly in the broad ligament before branching as it reaches
tion. The transverse cervical ligaments (cardinal ligaments, ligaments of the uterus at the level of the cervico-uterine junction (see Fig. 74.21B).
Mackenrodt) (Fig. 77.20) extend from the side of the cervix and lateral One major branch ascends the uterus tortuously within the broad liga-


fornix of the vagina, and are attached extensively on the pelvic wall. ment until it reaches the region of the ovarian hilum, where it anasto-
The lower parts of the ureters and pelvic blood vessels traverse the moses with branches of the ovarian artery. Another branch descends to
transverse cervical ligaments. Fibres of the pubocervical ligament pass supply the cervix and anastomoses with branches of the vaginal artery
forwards from the anterior aspect of the cervix and upper vagina to to form two median longitudinal vessels: the azygos arteries of the
diverge around the urethra, and are attached to the posterior aspect of vagina, which descend anterior and posterior to the vagina. Although
the pubic bones. there are anastomoses with the ovarian and vaginal arteries, the domi-
The transverse cervical and uterosacral ligaments are almost verti- nance of the uterine artery is indicated by its marked hypertrophy
cally orientated in the standing position and maintain the near- during pregnancy.
The tortuosity of the vessels as they ascend in the broad ligaments
is repeated in their branches within the uterine wall. Each uterine artery
Fundus of uterus Ovarian artery and vein
gives off numerous branches. These enter the uterine wall, divide and
Ovary and tube Anastomosis of ovarian Mesosalpinx run circumferentially as groups of anterior and posterior arcuate arter-
and uterine vessels
ies. They ramify and narrow as they approach the anterior and posterior
midline so that no large vessels are present in these regions. However,
the left and right arterial trees anastomose across the midline and uni-
lateral ligation can be performed without serious effects. Terminal
branches in the uterine muscle are tortuous and are called helicine
arterioles. They provide a series of dense capillary plexuses in the myo-
metrium and endometrium. From the arcuate arteries, many helical
arteriolar rami pass into the endometrium. Their detailed appearance
Broad
ligament changes during the menstrual cycle. In the proliferative phase, helical
arterioles are less prominent, whereas they grow in length and calibre,
Uterine artery becoming even more tortuous in the secretory phase.
and vein
Veins
Ureter
The uterine veins extend laterally in the broad ligaments, running adja-


cent to the arteries and passing over the ureters. They drain into the
Vagina internal iliac veins (see Fig. 77.3). The uterine venous plexus anasto-
moses with the vaginal and ovarian venous plexuses.
Lymphatic drainage
Fig. 77.19 The broad ligament (left), and blood supply to the uterus and Uterine lymphatics exist in the superficial (subperitoneal) and deep
ovaries (right). (With permission from Drake RL, Vogl AW, Mitchell A, parts of the uterine wall. Collecting vessels from the body of the uterus
Tibbitts R, Richardson P (eds), Gray’s Atlas of Anatomy, Elsevier, and cervix pass laterally in the parametrium to three main groups of
Churchill Livingstone. Copyright 2008.) lymph nodes: the external and internal iliac and the obturator nodes
Fig. 77.20 The supporting ligaments of the
pelvis, showing the transverse cervical
ligaments. (With permission from Waschke
J, Paulsen F (eds), Sobotta Atlas of Human
Anatomy, 15th ed, Elsevier, Urban &
Sacroiliac joint
Fischer. Copyright 2013.)
Pararectal space
Retrorectal space
Recto-uterine ligament
Rectum
Rectouterine pouch
Transverse cervical ligament
Cervix uteri
(cardinal ligament)
Uterovesicular pouch
Paravesical space
Ureter
Urinary bladder
Ureteric orifice
Retropubic space
Internal urethral orifice
Pubic symphysis


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Suspensory ligament of ovary
(infundibulopelvic ligament)
Uterine tube
Ovarian artery
Tubal branch
Fundus of uterus
Ovary
Branch to fundus
Ovarian branch
of uterus
Ligament of ovary
Helicine arterioles
B 56% C 40%
Uterine artery >90%
Vaginal branch
A
D 4% E 90% F 10%
Fig. 77.21 A, The normal arterial supply to the uterus and ovary. B–D, Variations in arterial blood supply to the ovary. E–F, Variations in arterial blood
supply to the fundus of the uterus. (A, With permission from Waschke J, Paulsen F (eds), Sobotta Atlas of Human Anatomy, 15th ed, Elsevier, Urban &
Fischer. Copyright 2013.)
(see Table 77.1). The external and internal iliac nodes surround their Endometrium
corresponding arteries. The obturator nodes lie in the obturator fossa The endometrium is formed by a layer of connective tissue, the endome-
between the external and internal iliac vessels; the obturator nerve trial stroma, which supports a single-layered columnar epithelium.
passes through the lower part of this group of lymph nodes. Before puberty, the epithelium is ciliated and cuboidal. It contains
Lymph vessels from the fundus of the uterus and the uterine glands that are composed largely of columnar cells secreting glycopro-
tubes may accompany the lymph drainage of the ovaries to para-aortic teins and glycogen. After puberty, the structure of the endometrium
nodes (see Fig. 77.3B). The region surrounding the isthmus of the varies with the stage of the menstrual cycle (see below). The glands are
uterine tube may drain along the round ligament to the superficial tubular, run perpendicular to the luminal surface and penetrate up to
inguinal nodes. the myometrial layer. The stroma consists of a highly cellular connective
tissue between the endometrial glands, and contains blood and lym-
Innervation phatic vessels.
myometrium
The nerve supply to the uterus is predominantly from the inferior hypogas-


tric plexus (see Fig. 77.5) (Shoja et al 2013). Some branches ascend with, The myometrium is composed of smooth muscle and loose connective
or near, the uterine arteries in the broad ligament. They supply the uterine tissue, and contains blood vessels, lymphatic vessels and nerves. It is
body and tubes, and connect with tubal nerves from the inferior hypogas- dense and thick at the uterine midlevel and fundus but thin at the tubal
tric plexus and with the ovarian plexus. The uterine nerves terminate in orifices. The body of the uterus has four muscular layers. The submu-
the myometrium and endometrium, and usually accompany the vessels cosal (innermost) layer is composed of longitudinal and some oblique
(see Table 77.2). Nerves to the cervix form a plexus that contains small smooth muscle fibres. Where the lumen of the uterine tube passes
paracervical ganglia. Sometimes, one ganglion is larger and is termed the through the uterine wall, this layer forms a circular muscle coat. The
uterine cervical ganglion. Branches may pass directly to the cervix uteri or vascular layer is external to the submucosal layer and is rich in blood
may be distributed along the vaginal arteries. vessels, as well as longitudinal muscle; it is succeeded by a layer of
Efferent preganglionic sympathetic fibres are derived from neurones predominantly circular muscle, the supravascular layer. The outer, thin,
in the last thoracic and first lumbar spinal segments; the sites where longitudinal muscle layer, the subserosal layer, lies adjacent to the
they synapse on their postganglionic neurones are unknown but are serosa.
presumably in the superior and/or inferior hypogastric plexuses (Lee The muscular fibres of the outer two layers converge at the lateral
et al 1973). Preganglionic parasympathetic fibres arise from neurones angles of the uterus and continue into the uterine tubes. Some fibres
in the second to fourth sacral spinal segments and relay in the paracervi- enter the broad ligaments as the round and ovarian ligaments; others
cal ganglia. Sympathetic activity may produce uterine contraction and turn back into the uterosacral ligaments. At the junction between the


vasoconstriction, and parasympathetic activity may produce uterine body and the cervix, the smooth muscle merges with dense, irregular
inhibition and vasodilation, but these activities are complicated by connective tissue containing both collagen and elastin, and forms the
hormonal control of uterine functions. majority of the cervical wall. Bilateral longitudinal fibres extend in the
lateral submucosal layer from the fundal angle to the cervix. Their
Microstructure muscle fibres differ structurally from those of typical myometrium, and
they may provide fast-conducting pathways that coordinate the contrac-
tile activities of the uterine wall.
Body of the uterus
The uterus is composed of three main layers. From its lumen outwards, serosa
these are the endometrium (mucosa), myometrium (smooth muscle The uterine body is covered by peritoneal serosa, which continues
layer) and serosa (or adventitia) (Fig. 77.22). downwards posteriorly to cover the supravaginal cervix. The anterior


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Ligament of ovary
Ovarian stroma
Uterine tube, isthmus Mesosalpinx
Fundus of uterus
Uterine tube, ampulla
Uterine tube Transverse ductules
Uterine tube, folds
Round ligament of uterus
Ligament of ovary
Uterine tube,
infundibulum;
fimbriae
Uterine cavity, endometrium
Myometrium Corpus albicans
Ovarian artery and veins
Cervical canal, palmate folds
Vesicular appendix (stalked hydatid)
Cervix of uterus, vaginal part
Vesicular ovarian follicles
External os of uterus Corpus luteum
Broad ligament of uterus
Uterine ostium,
Vagina Uterine tube, uterine part
intramural part
Cervix of uterus, supravaginal part Body of uterus
Isthmus of uterus
Fig. 77.22 The uterus, uterine tubes and ovaries. (With permission from Waschke J, Paulsen F (eds), Sobotta Atlas of Human Anatomy, 15th ed, Elsevier,
Urban & Fischer. Copyright 2013.)
EEGG
BV
L
LP
E
EC
EC
Fig. 77.23 The endocervical glands. These are are deep infoldings of the
lining of the endocervical canal (EC). The epithelium (E) is simple
columnar and mucus-secreting. The underlying lamina propria (LP) is
richly supplied with blood vessels (BV) and lymphatics (L). (Courtesy of Mr
Peter Helliwell and the late Dr Joseph Mathew, Department of
Histopathology, Royal Cornwall Hospitals Trust, UK.)
Fig. 77.24 The transformation zone of the uterine cervix. The single-
cervix and the lateral surfaces of the uterine body and cervix are not layered columnar epithelium lining the endocervical canal (EC) and its
covered by peritoneum. endocervical glands (EG) changes abruptly (arrow) to the stratified
squamous non-keratinizing epithelium of the external os and ectocervix
Cervix uteri
(below arrow).


The cervix consists of fibroelastic connective tissue and contains rela-
tively little (10%) smooth muscle. The elastin component of the cervical
stroma is essential to the stretching capacity of the cervix during child- The squamocolumnar junction, where the columnar secretory epi-
birth. The cervical canal is lined by a deeply folded mucosa with a thelium of the endocervical canal meets the stratified squamous cover-
surface epithelium of columnar mucous cells (Fig. 77.23). There are ing of the ectocervix, is located at the external os before puberty. As
branched tubular glands present within the mucosa, which are lined by oestrogen levels rise during puberty, the cervical os opens, exposing the
a similar secretory epithelium. The glands extend obliquely upwards endocervical columnar epithelium on to the ectocervix. This area of
and outwards from the canal. They secrete clear, alkaline mucus, which columnar cells on the ectocervix forms an area that is red and raw in
is relatively viscous except at the midpoint of the menstrual cycle, when appearance, called an ectropion (cervical erosion). It is then exposed to
it becomes more copious and less viscous to encourage the passage of the acidic environment of the vagina and, through a process of squa-
sperm. At the vaginal end of the cervix, the aperture of a gland may mous metaplasia (p. 40), transforms into stratified squamous epithe-
block and it then fills with mucus to form a Nabothian follicle (or cyst). lium. This area is thus known as the ‘transformation zone’ (Fig. 77.24).
None of the mucosa is shed during menstruation and so, unlike the Other hyperoestrogenic states, such as pregnancy and the use of oral
body of the uterus, it is not divided into functional and basal layers, contraceptive pills, can also result in an ectropion. This area is the most
and lacks spiral arteries. The surface of the intravaginal part of the cervix usual site of cervical intraepithelial neoplasia (CIN), which may


(ectocervix) is covered by non-keratinizing stratified squamous epithe- progress to malignancy. In postmenopausal women, the squamocolum-
lium, which contains glycogen. nar junction recedes into the endocervical canal.


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Magnetic resonance imaging of the uterus In prepubertal females, the uterus is smaller (only 4 cm in length),
and vagina and on T2-weighted images, the endometrium is minimal or absent,
and the junctional zone is indistinct. In postmenopausal women, the
corpus decreases in size and the zonal anatomy is indistinct.
On T2-weighted magnetic resonance imaging (MRI), the uterus displays
On T2-weighted MRI, the cervix has an inner, low-signal stroma
a zonal anatomy, with three distinct zones: the endometrium, junc-
continuous with the junctional zone of the uterus. Often, this is sur-
tional zone and myometrium (Fig. 77.25) (Minto et al 2011). The
rounded by an outer zone of intermediate signal intensity, which is
endometrium and uterine cavity appear as a high-signal stripe; the
continuous with the outer myometrium. The appearances do not
thickness varies with the stage of the menstrual cycle. In the early pro-
change with the menstrual cycle or oral contraceptive pill use. The
liferative phase, it measures up to 5 mm, and widens to up to 1 cm in
central stripe is very high-signal and is a consequence of the secretions
the mid-secretory phase. A band of low signal, the junctional zone,
produced by the endocervical glands.
borders the endometrium. It represents the inner myometrium and is
The vagina is best seen on T2-weighted MRI as a thin layer of high-
of constant thickness and signal throughout the menstrual cycle; it
signal intensity of the mucosa and an outer, low-signal layer of the
usually measures 5 mm. The outer myometrium is of medium-signal
submucosa and muscularis (see Fig. 77.25). MRI accurately demon-
intensity in the proliferative phase, and of high-signal intensity in the
strates congenital and acquired Müllerian anomalies, including uterine
mid-secretory phase as a result of the increased vascularity and promi-
and vaginal aplasia, duplication and septa (Grant et al 2010). Vaginal
nence of the arcuate vessels.


and perivaginal cysts are well visualized with T2-weighted MRI.
High-signal stroma Left ovary
at hilum of right ovary UTERINE (FALLOPIAN) TUBES
The uterine tubes are attached to the upper part of the body of the
uterus, and their ostia open into the uterine cavity (Figs 77.26, 77.27,
77.28). The medial opening of the tube (the uterine os) is located at
the superior angle of the uterine cavity. The tube passes laterally and
superiorly, and consists of four main parts: intramural, isthmus, ampulla
and fimbria (see Fig. 77.22). The intramural part is 0.7 mm wide and
1 cm long, and lies within the myometrium. It is continuous laterally
with the isthmus, which is 1–5 mm wide and 3 cm long; it is rounded,
muscular and firm. The isthmus is continuous laterally with the
ampulla, the widest portion of the tube with a maximum luminal
diameter of 1 cm. The ampulla is 5 cm long and has a thin wall and a
tortuously folded luminal surface. Typically, fertilization takes place in
its lumen. The ampulla opens into the trumpet-shaped infundibulum
at the abdominal os. Fimbriae, numerous mucosal finger-like folds
1 mm wide, are attached to the ends of the infundibulum and extend
from its inner circumference beyond the muscular wall of the tube. One
of these, the ovarian fimbria, is longer and more deeply grooved than
the others, and is typically applied to the tubal pole of the ovary. At the
time of ovulation, the fimbriae swell and extend as a result of engorge-
ment of the vessels in the lamina propria, which aids capture of the
released oocyte. All fimbriae are covered, like the mucosal lining
A
throughout the tube, by a ciliated epithelium whose cilia beat towards
the ampulla.
Small follicles in Urinary Uterus
low-signal outer stroma bladder Vascular supply and lymphatic drainage
Fluid in recto-uterine pouch
Myometrium Junctional zone (pouch of Douglas) Arteries


The blood supply to the uterine tubes is derived from ovarian and
uterine arteries (see Fig. 77.21A). The lateral third of the tube is supplied
by the ovarian artery, which continues in the mesosalpinx to anasto-
mose with branches from the uterine artery. The medial two-thirds of
the tube are supplied by the uterine artery.
Ostium of left uterine tube Fundus Ostium of right uterine tube
B
Bladder Endometrial cavity Vagina Cervix
Fig. 77.25 T2-weighted MRI scans of the uterus and ovaries. A, A coronal
T2-weighted MRI through a female pelvis showing the uterus and both
ovaries with the high-signal central stroma and lower-signal outer stroma.
B, A sagittal T2-weighted MRI through a female pelvis, showing the zonal
anatomy of the uterus. Fig. 77.26 A hysteroscopic view of the endometrial cavity of the uterus.


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Intramural
Isthmus of part of Uterine
uterine tube uterine tube cavity Uterine tube
LP SM
Fig. 77.29 The uterine tube (ampulla) with the mucosal lining thrown into
extensive folds that fill the lumen. The superficial epithelium includes both
secretory and ciliated cells, covering a lamina propria (LP). Smooth
Ampulla of uterine tube Vaginal speculum Spill of muscle (SM) occupies the wall of the tube. (Courtesy of Mr Peter Helliwell
contrast into and the late Dr Joseph Mathew, Department of Histopathology, Royal
peritoneal Cornwall Hospitals Trust, UK.)
cavity
Fig. 77.27 A digitally subtracted hysterosalpingogram. Radiopaque
contrast is introduced via a catheter inserted through the cervical os; the Innervation
catheter is introduced using a vaginal speculum. The contrast fills the
triangular uterine cavity. The lumina of the narrow intramural and isthmic The uterine tube is innervated by autonomic fibres that are distributed
parts of the uterine tubes may be traced inferolaterally from the superior
mainly with the ovarian and uterine arteries. Most of the tube has a dual
angles as they expand into the wider ampullae. Some contrast media has
sympathetic and parasympathetic supply (see Table 77.2). Preganglionic
escaped into the pelvic cavity from the abdominal ostia. (Courtesy of Dr
parasympathetic fibres are derived from the vagus for the lateral half of
Julia Hillier, Chelsea and Westminster Hospital, London.)
the tube, and pelvic splanchnic nerves for the medial half. Preganglionic
sympathetic supply is derived from neurones in the intermediolateral
column of the tenth thoracic to the second lumbar spinal segments;
Broad ligament Round ligament Origin of left uterine Uterus
postganglionic sympathetic fibres are most likely derived from the supe-
tube from uterus
rior hypogastric plexus, via the superior hypogastric and hypogastric
nerves (see Fig 77.5) (Chen et al 2014). Visceral afferent fibres travel
with the sympathetic nerves and enter the cord through corresponding


dorsal roots; they may also travel with parasympathetic fibres. The am-
pullary submucosa contains modified Pacinian corpuscles.
Microstructure
The walls of the uterine tubes show typical visceral mucosal, muscular
and serosal layers (Fig. 77.29). The mucosa is thrown into longitudinal
folds, which are most pronounced distally at the infundibulum and
decrease to shallow bulges in the intrauterine (intramural) portion. The
mucosa is lined by a single-layered, tall, columnar epithelium, which
contains mainly ciliated cells and secretory (peg) cells (so called because
they project into the lumen further than their ciliated neighbours), and
occasional intraepithelial lymphocytes. In the tube, ciliated cells pre-
dominate distally and secretory cells proximally. Their activities vary
with the stage in the menstrual cycle and with age. Secretory cells are
most active around the time of ovulation. Their secretions include
nutrients for the gametes and aid capacitation of the spermatozoa. Cili-
ated cells increase in height and develop more cilia in the oestrogenic
first half of the menstrual cycle. Their cilia waft the oocyte from the
Uterine tube Ovary Fimbriated end of uterine tube
open-ended infundibulum towards the uterus in fluids secreted by the
Fig. 77.28 A laparoscopic view of the origin of the uterine tube from the peg cells. The epithelium regresses in height towards the end of the cycle
uterus.
and postmenopausally, when ciliated cells are reduced in number.
The lamina propria provides vascular connective tissue support and
abundant lymphatic drainage vessels. The smooth muscle of the mus-
Veins cularis is arranged as an inner circular, or spiral, layer and an outer,
Venous drainage is similar to the arterial supply. The venous drainage thinner, longitudinal layer. Together, their contractile activity produces
of the lateral two-thirds of the uterine tube is via the pampiniform peristaltic movements of the tube, which assist propulsion of the
plexus to the ovarian veins, which open into the inferior vena cava on gametes and the fertilized ovum. The uterine tubes are covered exter-


the right side and the renal vein on the left side. The medial two-thirds nally by a highly vascular serosa.
of the tube drain via the uterine plexus to the internal iliac vein.
Lymphatic drainage OVARIES
Lymph drainage is via ovarian vessels to the para-aortic nodes and
uterine vessels to the internal iliac chain. It is possible for lymph to In the adult, non-pregnant state, the ovaries lie on each side of the
reach the inguinal nodes via the round ligament (see Table 77.1). uterus close to the lateral pelvic wall, suspended in the pelvic cavity by


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a double fold of peritoneum, the mesovarium, which is attached to the inferior extremity points downwards towards the pelvic floor. The ante-
upper limit of the posterior aspect of the broad uterine ligament. They rior border faces the posterior leaf of the broad ligament and contains
are dull white in colour and consist of dense fibrous tissue, in which the mesovarium. The posterior border is free and faces the peritoneum,
ova are embedded. Before regular ovulation begins, they have a smooth which overlies the upper part of the internal iliac artery and vein, and
surface but, thereafter, their surfaces are distorted by scarring that the ureter. On the right side, superior and lateral to the ovary, are the
follows the degeneration of successive corpora lutea (Speroff and Fritz ileocaecal junction, caecum and appendix. On the left side, the sigmoid
2004) (Fig. 77.30). Their average dimensions are 4 × 2 × 3 cm in repro- colon passes over the superior pole of the ovary and joins the rectum,
ductively mature women; they more than double their size during which lies between the medial surfaces of both ovaries. The meso-
pregnancy. In the neonate, their dimensions are 1.3 × 0.6 × 0.4 cm. Prior salpinx lies below the uterine tube. The ovarian ligament is inferior and
to the first menstrual period (menarche), the ovaries are about one- medial. The mesovarium and ovary lie inferiorly at its fimbrial end. The
third of the normal reproductive adult size; they gradually increase in round ligament is anterior to the tube. The superior and posterior sur-
size with body growth (Sforza et al 1993, Badouraki et al 2008) and faces of the tube lie free in the peritoneal cavity.
decrease significantly after the menopause. In embryonic and early fetal life, the ovaries are situated in the
The lateral surface of the ovary contacts parietal peritoneum in the lumbar region near the kidneys. They gradually descend along the
ovarian fossa. Behind the ovarian fossa are retroperitoneal structures, gubernaculum, stopping at the lesser pelvis. The relationship between


including the ureter, internal iliac vessels, obturator vessels and nerve, the location of the ovaries and surface landmarks is important in order
and the origin of the uterine artery (Fig. 77.31). The medial surface to enable shielding of the ovaries from exposure to radiation during
faces the uterus and uterine vessels in the broad ligament, and the radiological procedures. In girls, the ovaries lie at or below the iliac
peritoneal recess here is termed the ovarian bursa. Above the superior crest, just medial to the anterior superior iliac spine, and above the
extremity are the fimbria and distal section of the uterine tube. The pubic symphysis at all ages (Bardo et al 2009). Accessory ovarian tissue
may occur in the mesovarium or along the course of the gubernacula;
rarely, the ovaries may descend along the whole course of the guber-
Recto-uterine Uterus Origin of Ovary held up by laparoscopic nacula and are found in the labia majora. During pregnancy, the ovaries
pouch uterine tube instrument to demonstrate the
are lifted high in the pelvis; by 14 weeks of gestation, they become
irregular surface of the ovary
partly abdominal structures, and by the third trimester, they are totally
abdominal structures and lie vertically behind and lateral to the parous
uterus.
Peritoneal and ligamentous supports
of the ovary
The peritoneal and ligamentous supports of the ovary consist of the
infundibulopelvic and ovarian ligaments and the mesovarium.
Infundibulopelvic (suspensory) ligament
The suspensory or infundibulopelvic ligament of the ovary is a perito-
neal fold attached to the upper part of the lateral surface of the ovary,
which contain the ovarian vessels and nerves (see Fig. 77.18). On the
right side, the infundibulopelvic ligament is attached to a fold of peri-
toneum that is posterior and inferior to the caecum and appendix. On
the left side, the peritoneal attachment is higher than on the right, and
is lateral to the junction of the descending and sigmoid colons. As it
joins the peritoneum covering psoas major, the suspensory ligament of


the ovary passes superiorly over the external iliac vessels, genitofemoral
nerve and ureter.
Sigmoid loop in Small bowel in Fimbrial end of
Ovarian ligament (ligament of the ovary)
posterior pelvis posterior pelvis uterine tube
The ovarian ligament (ligament of the ovary) attaches the uterine (infer-
Fig. 77.30 A laparoscopic view of the right ovary, showing its distorted
omedial) extremity of the ovary to the lateral angle of the uterus, pos-
surface.
teroinferior to the uterine tube (see Fig. 77.22). It lies in the posterior
leaf of the broad ligament and contains some smooth muscle cells. It
is continuous with the medial border of the round ligament. Both liga-
Obturator Obturator External External
artery nerve iliac vein iliac artery ments are remnants of the gubernaculum.
Mesovarium
The mesovarium is a short peritoneal fold that attaches the ovary to the
back of the broad ligament. It carries blood vessels and nerves to the
ovarian hilum. The uterine tube arches over the ovary and ascends in
relation to its mesovarian border, then curves over its tubal end and
passes down on its posterior, free, border and medial surface.
Vascular supply and lymphatic drainage
Arteries
The ovarian arteries are branches of the abdominal aorta and originate
below the renal arteries. Each descends behind the peritoneum and, at
the brim of the pelvis, crosses the external iliac artery and vein to enter
the true pelvic cavity. Here, the artery turns medially in the ovarian
suspensory ligament and splits into a branch to the mesovarium that
supplies the ovary, and a branch that continues into the uterine broad
ligament, below the uterine tube, and supplies the tube. There are vari-
ations in the site of the division to the ovary and tube (see Fig. 77.21).
On each side, a branch passes lateral to the uterus to unite with the
uterine artery. Other branches accompany the round ligaments through
Sigmoid colon Obliterated internal Internal iliac Bifurcation of the inguinal canal to the skin of the labium majus and the inguinal


iliac artery artery common iliac artery region. Early in intrauterine life, the ovaries flank the vertebral column
Fig. 77.31 The broad ligament dissected and pelvic lymph nodes inferior to the kidneys, and so the ovarian arteries are relatively short;
removed to reveal the structures of the right-hand pelvic side wall. they gradually lengthen as the ovaries descend into the pelvis.


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Early primary Late primary Secondary Tertiary (Graafian follicle) Fig. 77.32 The microstructure
follicle follicle (antral) follicle of the ovary and follicles at
Ovarian surface Primordial Atretic Tunica Ruptured follicle various stages in their cyclical
(germinal) epithelium follicles follicle albuginea development and the formation
of corpora lutea and albicantes.
Note that, in the human ovary,
developing follicles are rarely
Oocyte at ovulation
seen.
Active corpus luteum
Peritoneum Hilum Cortex Medulla
Regressing corpus luteum
Ovarian vessels Corpus albicans
Veins hilum. It contains the ovarian follicles at various stages of development,
and corpora lutea and their degenerative remnants, depending on age
The ovarian veins emerge from the ovary as a plexus (pampiniform
or stage of the menstrual cycle. The follicles and the structures derived
plexus) in the mesovarium and suspensory ligament (see Fig. 77.3A).
from them are embedded in a dense stroma composed of a meshwork
Two veins emerge from the plexus and ascend with the ovarian artery;
of thin collagen fibres and fusiform fibroblast-like cells, arranged in
they usually merge into a single vessel before entering either the inferior
characteristic whorls. Stromal cells differ from fibroblasts in general
vena cava on the right side, or the renal vein on the left side.
connective tissue in that they contain lipid droplets, which accumulate
Lymphatic drainage in pregnancy. Stromal cells give rise to the thecal layers of maturing
The main lymphatic drainage of the ovaries is along vessels that follow ovarian follicles.
the ovarian veins to para-aortic nodes situated near the origin of the
renal arteries (see Table 77.1). Drainage may also occur via pelvic nodes ovarian follicles
into lower para-aortic nodes and, rarely, may follow the round ligament Primordial follicles
to the inguinal nodes. The development of ovarian follicles can be divided into different stages
(Fig. 77.32). At birth, the ovarian cortex contains a superficial zone of
Innervation primordial follicles. These consist of primary oocytes 25 µm in diam-


eter, each surrounded by a single layer of flat follicular cells. The oocyte
The ovarian innervation is derived from autonomic plexuses (see Table nuclei are slightly eccentric and have a characteristically prominent
77.2). The upper part of the ovarian plexus is formed from branches of nucleolus. They contain the diploid number of chromosomes (dupli-
the renal and aortic plexuses, and the lower part is reinforced from the cated as sister chromatids), arrested at the diplotene stage of meiotic
superior and inferior hypogastric plexuses. These plexuses consist of prophase since 16–20 weeks of fetal life (Speroff and Fritz 2004) (Fig.
postganglionic sympathetic fibres, preganglionic parasympathetic fibres 77.33). Many follicles degenerate either during prepubertal (including
from the sacral outflow, and visceral afferent fibres (Lee et al 1973). The prenatal) life, or through atresia at some stage after beginning the
efferent preganglionic sympathetic fibres are derived from the tenth and process of cyclical maturation during the child-bearing period. Their
eleventh thoracic spinal segments. remnants are visible as atretic follicles, the remains of which accumulate
throughout the period of reproductive life. After puberty, cohorts of up
Microstructure to 20 primordial follicles become activated in each menstrual cycle
(fewer are activated with advancing age). Their development takes a
number of cycles. Of the follicles activated in each cohort, usually only
In young females, the surface of the ovary is covered by a single layer
one follicle from one or other ovary becomes dominant, reaches matu-
of cuboidal epithelium, which contains some flatter cells. It appears
rity and releases its oocyte at ovulation.
dull white, in contrast to the shiny, smooth peritoneal mesothelial
Abnormal, non-growing follicles with indistinct germinal vesicle
covering of the mesovarium, with which it is continuous. A white line
membranes and absent nucleoli are sometimes found in prepubertal
around the anterior mesovarian border usually marks the transition
ovaries and, occasionally, in pubertal ovaries, but are not found in adult
between peritoneum and ovarian epithelium. The surface epithelium is


ovaries (Anderson et al 2014).
also termed the germinal epithelium, but this is a misnomer because it
is not the source of germ cells. Immediately beneath the epithelium,
there is a tough collagenous coat, the tunica albuginea. The ovarian Primary follicles
tissue it surrounds is divisible into a cortex, containing the ovarian fol- Primary follicles develop from primordial follicles. The first sign is a
licles, and a medulla, which receives the ovarian vessels and nerves at change in the follicle cells from flattened to cuboidal, followed by their
the hilum. proliferation to give rise to a multilayered follicle consisting of granu-
losa cells surrounded by a thick basal lamina (Fig. 77.34). Stromal cells
Ovarian cortex
immediately surrounding the follicle begin to differentiate into spindle-
Before puberty, the cortex forms 35%, the medulla 20% and interstitial shaped cells, which constitute the theca folliculi that will become the
cells up to 45% of the volume of the ovary. After puberty, the cortex theca interna. They are later surrounded by a more fibrous layer, the
forms the major part of the ovary, enclosing the medulla, except at the theca externa. At the same time, the oocyte increases in size and secretes


upper genital tract
1305
77
RETPAHC
TTEE
PP
pp AA
SS GGCC TTII
Fig. 77.33 Ovarian cortex with primordial (p) and early primary (P) follicles Fig. 77.35 An antral (secondary) follicle, showing the layers of the
within the stroma (S); stimulated primary follicles have cuboidal cells developing follicle wall. Granulosa cells (GC) enclose the antrum (A), filled
forming their wall. A large oocyte nucleus is visible in the plane of section with liquor folliculi, and surround the oocyte (not seen in the plane of
of most of the follicles (arrow). (Courtesy of Mr Peter Helliwell and the late section) within an eccentric cellular mass (right). The outer theca interna
Dr Joseph Mathew, Department of Histopathology, Royal Cornwall (TI) is a highly vascular layer that develops steroid secretory activity and
Hospitals Trust, UK.) is enclosed by the outermost theca externa (TE). (Courtesy of Mr Peter
Helliwell and the late Dr Joseph Mathew, Department of Histopathology,
Royal Cornwall Hospitals Trust, UK.)
Fig. 77.36 A low-power
view of a section of a
human ovary,
containing a corpus
luteum (C). (Courtesy of
Mr Peter Helliwell and
the late Dr Joseph
NN Mathew, Department of
Histopathology, Royal
Cornwall Hospitals
Trust, UK.)
FF
C
Fig. 77.34 A high-magnification micrograph of a primary follicle in a
human ovarian cortex. The pale oocyte, with its eccentric nucleus (N), is
separated from the follicle (F) by the zona pellucida (arrow). Cells of the cells synthesize oestrogens (primarily oestradiol). Follicular develop-
follicle wall are in the early stages of proliferation to form a multilayered,
ment is stimulated by follicle-stimulating hormone (FSH).
late primary follicle. (Courtesy of Mr Peter Helliwell and the late Dr Joseph
Mathew, Department of Histopathology, Royal Cornwall Hospitals Trust,
Tertiary (Graafian) follicle
UK.)
Although a number of follicles may progress to the secondary stage by


about the first week of a menstrual cycle, usually only one tertiary fol-
licle develops; the remainder become atretic. The surviving follicle
a thick layer of extracellular proteoglycan-rich material – the zona pel- increases considerably in size as the antrum takes up fluid from the
lucida – between its plasma membrane and the surrounding granulosa surrounding tissues and expands to a diameter of 2 cm. The term Graa-
cells of the early follicle; this is important for the process of fertilization. fian follicle is often used to describe this mature follicular stage. The
The granulosa cells in contact with the zona pellucida send cytoplasmic oocyte and a surrounding ring of tightly adherent cells, the corona
processes radially inwards; these contact and communicate with oocyte radiata, breaks away from the follicle wall and floats freely in the fol-
microvilli at gap junctions (Motta et al 2003). The follicular cells – in licular fluid. The primary oocyte, which has remained in the first meiotic
particular, the granulosa cells – continue to proliferate and so the thick- prophase (see Fig. 1.16) since fetal life, completes its first meiotic divi-
ness of the late primary follicle wall increases. sion to produce the almost equally large secondary oocyte and a small
first polar body with very little cytoplasm. The secondary haploid
Secondary (antral) follicles oocyte immediately begins its second meiotic division, but when it
Secondary (antral) follicles develop from primary follicles. The number reaches metaphase, the process is arrested until fertilization has
of granulosa cells continues to increase. Cavities begin to form between occurred. The follicle moves to the superficial cortex, causing the surface
them and are filled with a clear fluid (liquor folliculi) containing of the ovary to bulge. The tissues at the point of contact (the stigma)
hyaluronate, growth factors, and steroid hormones secreted by the with the tough tunica albuginea and ovarian surface epithelium are
granulosa cells. The follicle is now about 200 µm in diameter and eroded until the follicle ruptures and its contents are released into the


usually lies deep in the cortex. The cavities coalesce to form one large, peritoneal cavity for capture by the fimbria of the uterine tube. The
fluid-filled space – the antrum – which is surrounded by a thin, uniform oocyte at ovulation is still surrounded by its zona pellucida and corona
layer of granulosa cells, except at one pole of the follicle where a thick- radiata of granulosa cells. If fertilization does not occur, it begins to
ened granulosa layer envelops the eccentrically placed oocyte, to form degenerate after 24–48 hours.
the cumulus oophorus. The oocyte has now reached its maximum size
of about 80 µm and the inner and outer thecae are clearly differentiated Corpus luteum
(Fig. 77.35). As follicles mature, the theca interna becomes more prom- After ovulation, the remainder of the follicle becomes the corpus
inent and its cells more rounded and typical of steroid-secreting endo- luteum (Fig. 77.36). The walls of the empty follicle collapse and fold.
crine cells. They produce androstenedione, from which the granulosa The granulosa cells increase in size and synthesize a cytoplasmic


FEmALE REPRoduCTivE sysTEm
1306
8
NoiTCEs
carotenoid pigment (lutein), giving them a yellowish colour (hence, levels of oestrogen, which is produced by the ovary and which acts
corpus luteum). These large (30–50 µm) granulosa lutein cells form through receptors present on both the stromal and the epithelial cells.
most of the corpus luteum. The basal lamina surrounding the follicle The glands become tortuous and their lining epithelial cells become tall
breaks down, and numerous smaller theca lutein cells infiltrate the columnar in nature (Speroff and Fritz 2004).
folds of the cellular mass, accompanied by capillaries and connective
tissue. Extravasated blood from thecal capillaries accumulates in the
centre as a small clot, but this rapidly resolves and is replaced by con- SECRETORY PHASE
nective tissue. All lutein cells have a cytoplasm filled with abundant
smooth endoplasmic reticulum, characteristic of steroid-synthesizing The secretory phase coincides with the luteal phase of the ovarian cycle.
endocrine cells. Granulosa lutein cells secrete progesterone and oestra- The endometrial changes are driven by progesterone and oestrogen,
diol (from aromatization of androstenedione synthesized by theca secreted by the corpus luteum. Steroid receptors in the endometrium
lutein cells). The two cell types also respond differently to circulating activate a programme of new gene expression that produces, in the fol-
gonadotropins: theca lutein cells respond to luteinizing hormone (LH) lowing 7 days, a highly regulated sequence of differentiative events,
and granulose lutein cells respond to FSH. Theca lutein cells express presumably required to prepare the tissue for blastocyst implantation.
receptors for human chorionic gonadotrophin (hCG). If the oocyte is Part of the response is direct, but there is evidence that some of the
not fertilized, the corpus luteum (of menstruation) functions for effects may be mediated through growth factors (Speroff and Fritz
12–14 days after ovulation, then atrophies. The lutein cells undergo 2004). The first morphological effects of progesterone are evident