C H A P T E R 11 | A l i me n tar y S y ste m 219
Lesser omentum
Hepatoduodenal ligament
Diaphragm
Bare area of liver
Falciform ligament
Umbilical vein
(carries oxygenated
blood to embryo)
Extraembryonic coelom
Midgut loop
Peritoneal cavity (former
intraembryonic coelom)
F I G U R E 1 1 – 9   Median section of caudal half of an embryo at the end of the fifth week, showing the liver and associated liga-
ments. The arrow indicates the communication of the peritoneal cavity with the extraembryonic coelom.
Development of Pancreas
10 The pancreas develops between the layers of the mesen-
tery from dorsal and ventral pancreatic buds of endoder-
mal cells, which arise from the caudal part of the foregut
(Fig. 11-10A and B, and see also Fig. 11-9). Most of the
pancreas is derived from the larger dorsal pancreatic bud,
which appears first and develops at a slight distance
cranial to the ventral bud.
The smaller ventral pancreatic bud develops near the
entry of the bile duct into the duodenum and grows
between the layers of the ventral mesentery. As the duo-
denum rotates to the right and becomes C shaped, the
bud is carried dorsally with the bile duct (see Fig. 11-10C
to G). It soon lies posterior to the dorsal pancreatic
bud and later fuses with it. The ventral pancreatic bud
forms the uncinate process and part of the head of the
pancreas.
As the stomach, duodenum, and ventral mesentery
rotate, the pancreas comes to lie along the dorsal abdomi-
nal wall (in a retroperitoneal position). As the pancreatic
buds fuse, their ducts anastomose, or open into one
another (see Fig. 11-10C). The pancreatic duct forms
from the duct of the ventral bud and the distal part of the
duct of the dorsal bud (see Fig. 11-10G). The proximal
part of the duct of the dorsal bud often persists as an
accessory pancreatic duct that opens into the minor duo-
denal papilla, located approximately 2 cm cranial to the
main duct (see Fig. 11-10G). The two ducts often com-
municate with each other. In approximately 9% of people,
the pancreatic ducts fail to fuse, resulting in two ducts.
Molecular studies show that the ventral pancreas
develops from a bipotential cell population in the ventral
region of the duodenum where the transcription factor
Hepatogastric ligament Dorsal mesogastrium
Dorsal pancreatic bud
Celiac artery
Gallbladder
Dorsal aorta
Superior mesenteric artery
Free edge of ventral mesogastrium
Inferior mesenteric artery
PDX1 is expressed. A default mechanism involving
FGF-2, which is secreted by the developing heart, appears
to play a role. Formation of the dorsal pancreatic bud
depends on the notochord secreting activin and FGF-2,
which block the expression of Shh in the associated
endoderm.
Histogenesis of Pancreas
The parenchyma (basic cellular tissue) of the pancreas is
derived from the endoderm of the pancreatic buds, which
forms a network of tubules. Early in the fetal period,
pancreatic acini (secretory portions of an acinous gland)
begin to develop from cell clusters around the ends of
these tubules (primordial pancreatic ducts). The pancre-
atic islets develop from groups of cells that separate from
the tubules and lie between the acini.
Recent studies show that the chemokine, stromal-
cell derived factor 1 (SDF-1), expressed in the mesen-
chyme, controls the formation and branching of the
tubules. Expression of transcription factor neurogenin-
3 is required for differentiation of pancreatic islet endo-
crine cells.
Insulin secretion begins during the early fetal period
(at 10 weeks). The cells containing glucagon and soma-
tostatin develop before differentiation of the beta cells
that secrete insulin. Glucagon has been detected in fetal
plasma at 15 weeks.
The connective tissue sheath and interlobular septa of
the pancreas develop from the surrounding splanchnic
mesenchyme. When there is maternal diabetes mellitus,
the beta cells that secrete insulin in the fetal pancreas are
chronically exposed to high levels of glucose. As a result,
these cells undergo hypertrophy to increase the rate of
insulin secretion.