Secti on IC

EMBRYOLOGY
Topic 1 SOME PRELIMINARY CONSIDERATIONS

SHORT NOTE
Q.1. Discuss about chromosome.

A typical chromosome is made up of two rod-shaped structures or

chromatids placed more or less parallel to each other (Fig. 1C.1.1).

The chromatids are united to each other at a light staining area called
the centromere or kinetochore.
Each chromatid has two arms, one on either side of the centromere.
Each chromosome has a very large number of Satellite
structures called genes on it, which guide the
performance of particular cellular functions and Secondary
constriction
in turn lead to the development of Short arm of
particular features of a species or an chromatid

individual.
Chromosomes are predominantly made up of a
nucleic acid called deoxyribonucleic acid (DNA). Centromere

Chromosomes control the development and functioning of

cells by determining what types of proteins will
Long arm of
be synthesized in them. Fig. 1C.1.1 Structureofa
chromatid
chromosome.
Topic 2 SPERMATOGENESIS AND OOGENESIS

SHORT NOTES
Q.1. Oogenesis.Q.2. Graafian follicles.

In females, maturation from primitive germ cell to mature At maturity,

the follicle of 10 mm or more in diameter is known gamete
is called oogenesis, which begins before birth. as the tertiary follicle, vesicular follicle
or Graafian follicle.
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Fig. 1C.2.1 Graafianfollicleshowingdifferentstagesofdevelopment.

Q.1. Formation, functions and fate of notochord. whose cells become intercalated
in the hypoblast so
1. Prenotochordal cells invaginating in that for a short time
the primitive pit move forwards the midline of the embryo
until they reach the prechordal consists of two cell layers
plate, that form the notochordal
plate (Fig 1C.4.1).
The Graafian follicle is surrounded by (Fig. 1C.2.1): 2. The theca externa, which
gradually merges with the 1. The theca interna, which is
composed of cells having ovarian stroma. characteristics of steroid secretion,
is rich in blood vessels.

Topic FORMATION OF GERM

3 LAYERS
SHORT NOTE
Q.1. Effects of
fertilization
. The phases of fertilization include the following:
Phase 1: Penetration of the corona radiata
The process by which male and female gametes fuse is Phase 2: Penetration
of the zona pellucida known as fertilization. It occurs in the
ampullary region of the uterine tube. Phase 3: Fusion of the oocyte and
sperm cell membranes.

Topic FURTHER DEVELOPMENT OF

4 EMBRYONIC DISC
SHORT ESSAY
As the hypoblast is replacedby endoderm cells Primitive pit and moving in
at the streak, cells of the
Connecting stalk
notochordal plate neurenteric canal proliferate and Amnion
detach from the endoderm. They then
Allantois
form a solid cord of cells—the Wall of
yolk sac Notochord Cloacal plate
Prechordal plate (membrane)

Notochordal plate
 Intraembryonic mesoderm

Extraembryonic
mesoderm Embryology
Endoderm 121
Notochord

definitive notochord that underlies the neural tube; serves as the

basis for the axial skeleton.
Elongation of the notochord is a dynamic process; the cranial end
forms first and caudal regions are added as the primitive streak
assumes a more caudal position.
The notochordand prenotochordal cells extend cranially to the prechordal plate
and caudallyto the primitive pit. At the point where the pit
forms an indentation in the epiblast,the neurenteric canal temporarily
connects the amniotic and yolk sac cavities.
Notochord underlies the neural tube and serves as the basis
for the axial skeleton. Fig. 1C.4.1 Schematicviewsandscanningelectronmicrographs illustrating
formationofthenotochord.

SHORT NOTE
Q.1. Formation and fate of notochord. During further development, the plate detaches
from the endoderm and a solid cord known as notochord
is formed.
Prenotochordal cells invaginate—in the primitive pit and The definitive
notochord underlies the neural tube and move forwards cephalad
until they reach the prechordal serves as the basis for the axial
skeleton. plate. They intercalate in the endoderm as notochordal plate.
Topic 5 PHARYNGEAL ARCHES

SHORT ESSAYS
Q.1. Development of mandible. 1C.5.1A) between the tuberculum
impar and the copula at a
The ventral portion of viscerocranium point known as the foramen
gives rise to the mandibular cecum.
process, which containsMeckel’s
cartilage.
Mesenchyme around Meckel’s cartilage
condenses and ossifies by
membranous ossification to
give rise to the mandible.
Meckel’s cartilage disappears
except in the sphenomandibular
ligament.
The dorsal tip of the mandibular
process, along with that of the
second pharyngeal arch, gives rise
to the incus, the malleus and
the stapes.
The mandible is also formed by
membranous ossification of
mesenchymal tissue surrounding
Meckel’s cartilage. Q.2.
Development of thyroid. Or
Development of thyroid gland and its
anomalies.

The thyroid gland appears as an

epithelial proliferation in the
floor of the pharynx (Fig.
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Fig. 1C.5.1 (A)Thyroidglandinthefloorofpharynx.(B)Descendingpathwayofthyroidgland.

As the development progresses, the Foramen cecum
thyroid descends in front of Thyroglossal cyst
the pharyngeal gut (Fig. Epiglottis
1C.5.1B), hyoid bone and the
laryngeal cartilages as a Hyoid bone

bilobed diverticulum. Thyroglossal cysts

During this migration, the thyroid Thyroid cartilage
remains connected to the
Cricoid cartilage
tongue by a narrow canal—the
thyroglossal duct, which disappears Thyroid gland
later.
It acquires a small median isthmus and Fig. 1C.5.2 Locationofthyroglossalcyst.
two lateral lobes by the A thyroglossal fistula usually arises
time it reaches its final secondarily after rupture of a
position in front of the thyroglossal cyst and is
trachea in the seventh week. connected to the outside by
By the end of the third a fistulous canal, but
month, the thyroid begins to may be present at birth.
function, at the time at Q.3. Cartilage, arch, artery and muscular
which the first follicles containing derivatives of first pharyngeal arch.
colloid become visible.
Follicular cells produce the colloid that First pharyngeal arch(Fig.1C.5.3)
serves as a source of The first pharyngeal arch consists of
thyroxinand triiodothyronine. a dorsal portion known as the
Parafollicular or C-cells derived maxillary process, which extends
from the ultimobranchial body forwards beneath the region of
serve as a source of the eye, and a ventral
calcitonin. portion— the mandibular process—
Anomalies which containsMeckel’s cartilage.
Thyroglossal cyst is a cystic During further development Meckel’s
remnant of the thyroglossal cartilage disappears except for
duct. It lies at any two small portions at its
point along the midline of the dorsal end that persist and
neck, starting from base of the form the incus and malleus.
tongue up to the thyroid Mesenchyme of the maxillary
isthmus (Fig. 1C.5.2). process gives rise to the
Body of tongue premaxilla, maxilla, zygomatic
bone and part of the
temporal bone through
membranous ossification. The
 Embryology 121

mandible is also formed by

membranous ossification of
mesenchymal tissue surrounding
Meckel’s cartilage. The
first arch also contributes to
formation of the bones of
the middle ear.
Musculature of the first pharyngeal Man Mandi
arch includesthe muscles of d b
mastication (i.e. the temporalis, i u
masseter and medial and b l
lateral pterygoids), anterior belly of a
r
the digastric, mylohyoid,
d
tensor tympani and tensor palatine. iv
The nerve supply to the muscles of is
the first arch is provided i
by the mandibular branch of o
the trigeminal nerve. Since n
o
mesenchyme from the first f
arch also contributes to the tr
dermis of the face, sensory i
supply to the skin of the g
face is provided by e
ophthalmic, maxillary and m
i
mandibular branches of the n
trigeminal nerve. a
Tuberculumimpar l
Laterallingualswelling
Terminalsulcus Body of tongue n
e
r
v
e

Foramen Palatine tonsil

cecum
Copula
(hypobranchial Laryngealorific e Root of tongue
eminence)
Arytenoidswellings Epiglottis
Epiglottal
swelling

5 weeks 5 months
Fig. 1C.5.3 Pharyngealarches(ventralportion).

SHORT NOTES
Q.1. Derivatives of first pharyngeal arch.

The derivatives of first pharyngeal

arch, i.e. mandibular arch are
listed in Table 1C.5.1 and
shown in Fig. 1C.5.4.
Table 1C.5.1 Derivativesoffirstpharyngealarch
Pha Nerve Muscl Skelet
r e o
y s n
n
g
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u f rtil o
l m ag r
a a e b
r st in el
ic cu ly
a a s; o
r ti M f
c o ec di
h n ke g
(t l’s a
e ca st
m rtil ri
p ag c;
o e, t
r m e
al all n
is eu s
, s, o
m an r
a ter palati
s ior n
s lig e
e a ,
t m t
e en e
r; t n
m of s
e m o
di all r
al eu ty
a s, m
n sp p
d he a
la no ni
t m Malleus Incus
e an
r di Stapes
al bu Meckel’s cartilage
Styloid process
p lar
t lig I Stylohyoid ligament
e a II Lesser horn of hyoid bone
r m Greater horn of hyoid bone
y en III
g t, Body of hyoid bone IV
oi po
Thyroid cartilage VI
d rti
s on Cricoid cartilage
); of Tracheal rings
m m
Fig. 1C.5.4 Thefirstpharyngealarchderivatives. Q.2.
yl an
o di Derivatives of second pharyngeal arch.
h bl Or
y e Derivatives of second pharyngeal arch
oi (muscles). Or
d
Derivative of mesoderm of second
;
a pharyngeal arch.
n
t The derivatives of second pharyngeal
e arch, i.e. hyoid arch are
ri listed in Table 1C.5.2.
 Embryology 121

Table 1C.5.2 Derivativesofsecondpharyngealarch ric; n

Phar sty of
y loh b
n yoi o
g d; d
e sta y
a pe of
l diu h
a s y
r Skele oi
c t d
h Ner Muscle o b
s n o
n
Hyoid VII Facial Stape
e
a mu s,
r scl st Q.3. Structures developed from third
c es yl pharyngeal arch
h of oi Structures developed from third
ex d pharyngeal arch are listed in
pr p
es r
Table 1C.5.3.
sio o Table 1C.5.3 Derivativesofthirdpharyngealarch
n c Pha
(b e r
uc s y
cin s, n
ato st g
r, yl e
au o a
ric h l
ula y
ris; oi a
fro d r
nta li c
lis, g h Muscle Sk
pla a Nerve s
tys m
Thir Glossop Styloph Gr
ma e
d har ary
, nt
yng ng
or , a eal eu
bic le r s
ula s c
ris s h
ori e
s, r
or h Q.4. Derivatives of third endodermal pouch
bic o pharynx.
ula r
ris n The third and fourth pouches are
oc a
characterized at their distal
uli) n
; d extremity by a dorsal and
po u a ventral wing (Fig. 1C.5.5).
ste p gland, while the ventral wing
rio p forms the thymus, as shown in
r e the previous figure.
bel r
Q.5. Formation and fate of first pharyngeal
ly p
of o pouch.
dig rti
ast o
 1. The first pharyngeal pouch forms a stalk-like
I diverticulum—the tubotympanic recess—which comes
in contact with the epithelial lining of the first pharyn-
120 Quick Review Series: BDS
II
1st Year
Palatine geal cleft—the future external auditory meatus.
tonsil III 2. The distal portion of the diverticulum widens into
a
Parathyroid saclike structure—the primitive tympanic or middle ear
IV
gland (inferior)
Thymus cavity—and the proximal part remains narrow, forming
Parathyroid the auditory tube. gland (superior)
Q.6. Give persistent structures of fibrous envelope of Meckel’s cartilage.
Fig. 1C.5.5
Derivativesof During further development Meckel’s cartilage disappears
third
except for two small portions at its dorsal end that
pharyngeal
pouch. persist and form the incus and malleus.
In the fifth
week,
epithelium
of the
dorsal
wing of
the
third
pouch

differentiates
into the
inferior
parathyroid

Topic
FACE, NOSE AND PALATE
6
SHORT ESSAYS
Q.1 Development of face. Maxillary prominences can be
Or Frontonasal process of embryo. distinguished lateral to the
stomodeum, and mandibular
1. At the end of the prominences can be
fourth week, facial prominences distinguished caudal to this
consisting primarily of neural structure.
crest-derived mesenchyme appear (Fig. The frontonasal prominence, formed by
1C.6.1). proliferation of mesenchyme
ventral to the brain vesicles,
constitutes the upper border of
the stomodeum.
Under inductive influence of
the ventral portion of the
forebrain on both sides of
the frontonasal prominence,
local thickenings of the
surface ectoderm—the nasal or
Frontonasal olfactory placodes—originate.
prominence
5. During the fifth week, the nasal
placodes invaginate to form
Nasal pit Eye Nasal pit nasal pits. They create a ridge
Maxillary Lateral nasal
prominence
of tissue that surrounds each
prominence
Embryology 121
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Lateral nasal a palatal component, which forms the

prominence
triangular primary palate.
Medial nasal
prominence The intermaxillary segmentis continuous
Maxillary with the rostral portion of the
prominence Eye
nasal septum, which is formed by
Mandibular
prominence Nasolacrimal the frontal prominence. Structures
Philtrum
A groove B contributing to the formation
Fig. 1C.6.2 Developmentofface. of face are explained in
The upper lip is formed by the Table 1C. 6.1.
Table 1C.6.1 Structurescontributingtoformationofthe
two medial nasal prominences and
face
the two maxillary
Name of the Structures formed
prominences. prominence
The lower lip and jaw form Frontonasal Forehead,bridgeofnose,
from the mandibular medialandlateralnasal
prominences that merge across the prominences
midline. Maxillary Cheeks,lateralportionof
Initially, the maxillary and upperlip
lateral nasal prominences are Medialnasal Philtrumofupperlip,crest
separated by a deep andtip ofnose
furrow—the nasolacrimal groove. Lateralnasal Alareofnose
The maxillary prominences Mandibular Lowerlip
enlarge to form the cheeks and Q.2. Give the development of palate and
maxillae. associated anomalies.
The nose is formed from five
facial prominences; the frontal The primary palate is derived from the
prominence gives rise to the intermaxillary segment.
bridge; the merged medial nasal The main part of the definitive
prominences provide the crest and palate is formed by two
tip; and the lateral nasal shelves-like outgrowths—the palatine
prominences form the sides shelves—from the maxillary
(alare) (Fig. 1C.6.3). prominences.
Maxilla with
During the sixth week of
Philtrum of lip four incisor teeth
development outgrowths appear and
Intermaxillary segment are directed obliquely downwards
Primary on each side of the
palate tongue.
Fused In the seventh week, the palatine
Maxillary
palatal shelves ascend to attain a
process
plates horizontal position above the
Fig. 1C.6.3 Formationofmaxillaryprocess(5-week-old tongue and fuse, forming the
embryo).
secondary palate (Fig. 1C.6.3).
As a result of medial growth of the Anteriorly, the palatine shelves fuse
maxillary prominences, the two with the triangular primary
medial nasal prominences merge not palate, and the incisive foramen is
only at the surface but the midline landmark between the
also at a deeper level primary and secondary palate.
forming the intermaxillary segment,
At the same time as the
which is composed of:
palatine shelves fuse the nasal
a labial component, which forms the septum grows down to join
philtrum of the upper lip; with the cephalic aspect of the
an upper jaw component, which newly formed palate.
carries the four incisor teeth and
 Embryology 121

The anomaliesassociated with which may be due to

development of palate are as smallness of the fusion
follows: process itself or failure of the
Cleft lip and cleft palate are tongue to drop from between the
common defects that result in shelves because of micrognathia.
abnormal facial appearance and Cleft lip and cleft palate are
defective speech. common defects that result in
Cleft palate results from a lack of abnormal facial appearance and
fusion of the palatine shelves, defective speech.
which may be due to The frequency of isolated cleft palate is
smallness of the fusion much lower than that of
process itself or failure of the cleft lip (1/2500 births), occurs
tongue to drop from between the more often in females (67%)
shelves because of micrognathia than males and is not
(Fig. 1C.6.3). related to maternal age.
The frequency of isolated cleft palate is Anticonvulsant drugs, such as
much lower than that of phenobarbital and diphenylhydantoin,
cleft lip around 1 in given during pregnancy increase the
2500 births, occurs more often in risk of cleft palate.
females (67%) than males and is Palatine aponeurosis.
not related to maternal
age. The posterior border of the hard
Anticonvulsant drugs, such as palate provides attachment to
phenobarbital and diphenylhydantoin, the palatine aponeurosis.
given during pregnancy increase the Development of palate.
risk of cleft palate.
SHORT
NOTES
Q.1. Hare lip or cleft lip. The primary palate is derived from the
intermaxillary segment.
Cleft lip and cleft palate are The main part of the definitive
common defects that result in palate is formed by two
abnormal facial appearance and shelf-like outgrowths from the
defective speech. maxillary prominences known as
Median cleft lip is a palatine shelves, which appear in the
rare abnormality caused by sixth week of development and
incomplete merging of the two are directed obliquely downwards
medial nasal prominences in the on each side of the
midline. tongue.
Most cases of cleft lip and During seventh week the palatine
cleft palate are multifactorial. shelves ascend to attain a
Cleft lip, approximately 1 in horizontal position above the
1000 births, occurs more frequently tongue and fuse, forming the
in males (80%) than in secondary palate.
females;its incidence increases In the Ventral view of palate, lip
slightly with maternal age and and nose (Fig. 1C.6.4)
varies among populations. Anteriorly the shelves fuse with the
Q.2. Cleft palate. triangular primary palate, and the
incisive foramen is the midline
Cleft palate results from a lack of landmark between the primary and
fusion of the palatine shelves, secondary palates.
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At the same time, as the indicates that tissue of the

palatine shelves fuse the nasal third arch overgrows that of

Topic ALIMENTARY SYSTEM

7 SHORT ESSAY
Q.1.
Developme
nt of
tongue. swelling—the tuberculum impar. These three swellings originate
from the first pharyngeal arch (Fig. 1C.7.1).
septum grows down and joins the second.
with the cephalic aspect of the The extreme posterior part of the
newly formed palate. tongue is innervated by the
Primary Incisive superior laryngeal nerve, reflecting
palate foramen
its development from the
fourth arch.
Some of the tongue muscles probably
differentiate in situ; but
most are derived from myoblasts
Uvula originating in occipital
2. A second median swelling somites. Thus, tongue musculature is
called the copula or innervated by the hypoglossal
1. At approximately 4 weeks, the nerve.
tongue appears in embryo in The general, sensory innervation of the
hypobranchial eminence is tongue is easy to understand:
formed by mesoderm of the The body is suppliedby the
the form of two lateral trigeminal nerve—the nerve of
lingual swellings and one the first arch.
median second, third and part of The root is suppliedby the
the fourth arch. glossopharyngeal and vagus
Fig. 1C.6.4 Ventralviewofpalate,lipandnose. nerves—the nerves of the
A third median swelling formed by the third and fourth arches,
posterior part of the fourth respectively.
arch.
As the lateral swellings increase in
size, they overgrow the
tuberculum impar and merge,
forming the anterior two-thirds or
body of the tongue; sensory
innervation to this area is
by the mandibular branch of
the trigeminal nerve.
The posterior part, or root, of the
tongue originates from the
second, third and part of the
fourth pharyngeal arch. The
sensory innervation to this
part of the tongue suppliedby
the glossopharyngeal nerve
 Embryology 121

Q.1. Give a detailed account of development of Dental lamina

milk
Special sensory innervation (taste) to the
the appearance of a bell (bell C. Bell stage D. Advanced Bell stage
stage). Fig. 1C.7.2 Developmentalstagesofatoothbud.
4. Mesenchymal cells of the
papilla adjacentto the inner dental
layer differentiate into odontoblasts,
which later
produce dentine; the remaining cells 5. Epithelial cells of the
of the dental papilla form the outer dental epithelium differentiate
pulp of the tooth. into ameloblasts,which produce long
enamel prisms
body of the tongue is
provided by the chorda
tympani branch of the facial
nerve.
Tuberculumimpar
Laterallingualswelling Bodyoftongue
Terminalsulcus

Foramen Palatinetonsil
cecum
Copula
(hypobranchial Laryngealorifice Rootoftongue
eminence)
Arytenoidswellings Epiglottis
Epiglottal
swelling
Fig. 1C.7.1 Pharyngealarchesinvolvedindevelopmentoftongue.

SHORT NOTES
teeth. Permanent
tooth bud

Inner } Dental
By the sixth week of development
the basal layer of the Outer
epithelium epithelial lining of the oral
cavity forms a C-shaped A. Bud
stage B. Cap stage structure—the dental lamina—
along the length of the upper
and lower jaws.
This lamina subsequently gives rise to
10 dental buds in each jaw,
which form the primordia of the ectodermal components of the
teeth.
The deep surface of the buds invaginates, resulting in the cap stage
of tooth development. As the dental cap grows and the indentation
deepens, the tooth takes on
120 Quick Review Series: BDS 1st Year

that are deposited over the dentine. The

contact layer between the enamel and
dentine layers is known as the
enamel–dentine junction.
Formation of the root of the
tooth begins when the dental
epithelial layers penetrate
into the underlying
mesenchyme and form the
epithelial root sheath.
Cells of the dental papilla lay
down a layer of dentine
continuous with that of the
crown. Mesenchymal cells on the
outside of the tooth and in
contact with dentine of the
root differentiate into
cementoblasts and produce a
thin layer of specialized
bone—the cementum.
Outside of the cemental
layer, mesenchyme gives rise to
the periodontal ligament,
which holds the tooth firmly in
position and functions as a
shock absorber.
With further lengthening of the
root, the crown is gradually
pushed through the overlying
tissue layers into the oral
cavity. The eruption of deciduous
or milk teeth occurs 6–24
months after birth (Fig. 1C.7.2).
Q.2. Describe the structure of tooth (labelled
diagram only).

A B
 Embryology 121

Fig. 1C.7.3 Thestructureoftooth:(A)beforebirth;(B)after

eruption.
Q.1. Development of pituitary gland. membrane known as Rathke’s
pouch and
The hypophysis or pituitarygland A downward extension of the
develops from two completely diencephalon—the infundibulum.
different parts (Fig. 1C.9.1): Rathke’s pouch appears as an
An ectodermal outpocketing of the evagination of the oral

Topic ELEMENTARY GENETICS

8
SHORT NOTE
Q.1. Karyotype each chromosome individually. Classification of chromosomes in
and any two this way is called karyotyping.
clinical Example: Turner syndrome: It is characterized by the
features of absence of ovaries (gonadal dysgenesis) and short
Turner stature. Other commonly associated abnormalities are
syndrome. webbed neck, lymphoedema of the extremities, skeletal
deformities and a board chest.
Individual
chromosomes
differ from
one
another in
total
length, in
the
relative length
of the
two arms
and in
various other

characteristics;
these
differences
enable us
to
identify

Topic
MISCELLANEOUS
9
SHORT NOTES
stomodeum immediately in cavity when the embryo is
front of the buccopharyngeal approximately 3 weeks old, and
subsequently grows dorsally towards the
120 Quick Review Series: BDS 1st Year

infundibulum. During further two lateral lobes by the

development, cells in the time it reaches its final
anterior wall of Rathke’s position in front of the
pouch increase rapidly in number and trachea in the seventh week. By
form the anterior lobe of the the end of the third
hypophysis or adenohypophysis. month the thyroid begins to
A small extension of this lobe—the function at the time at
pars tuberalis–grows along the which the first follicles containing
stalk of the infundibulum and colloid become visible (Fig. 1C.9.2).
eventually surrounds it. Follicular cells produce the
The posterior wall of colloid that serves as a
Rathke’s pouch develops source of thyroxinand
into the pars intermedia. triiodothyronine. Parafollicular or C-
The infundibulum gives rise to the cells derived from the
stalk and the pars nervosa or ultimobranchial body serve as a
posterior lobe of the source of calcitonin.
hypophysis (neurohypophysis). It Development of parathyroid gland.
is composed of neuroglial
cells. In addition, it In the fifth week of embryonic
containsa number of nerve development, epithelium of the
fibres from the hypothalamic dorsal wing of the third
area. pharyngeal pouch differentiates
Development of thyroid gland. into the inferior parathyroid
gland.
The thyroid gland appears as an Epithelium of the dorsal wing of
epithelial proliferation in the the fourth pharyngeal pouch
floor of the pharynx between the forms the superior parathyroid
tuberculum impar and the gland.
copula at a point known as Whenthe parathyroid gland loses
the foramen cecum. As a contact with the wall of the
bilobed diverticulum, the thyroid pharynx, it attaches itself to
descends in front of the the dorsal surface of the
pharyngeal gut. During this caudallymigrating thyroid as the
migration the thyroid remains superior parathyroid gland.
connected to the tongue by
a narrow canal called the
thyroglossal duct, which disappears
later. (Fig. 1C.5.1).
As the development progresses the
thyroid gland descends in
front of the hyoid bone and
the laryngeal cartilages. It
acquiresa small median isthmus and
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