25 DEVELOPMENTAL ANATOMY AND
PHYSIOLOGY OF THE STOMACH
Steven J. Czinn  •  Samra S. Blanchard
The gastrointestinal tract begins as a primitive tubular system
and is one of the first organs to polarize the embryo by form-
ing an entry and exit with an anterior and posterior axis, also
known as the craniocaudal axis, extending from the mouth to
the cloaca (Figure 25-1). The nonneural elements of the gut
are derived from endodermal and mesodermal cells. Bilateral
folding of these layers forms the intestinal lumen, which is sur-
rounded by concentric endodermal and splanchnic epithelia,
creating a tubular gut. Cells from the outer epithelium migrate
outward and form a loose mesenchyme, which later forms the
muscle and connective tissue, while neural elements migrate
from the neural crest at vagal and sacral levels to form the
enteric nervous system.
The tubular gut has three distinct sections: the foregut,
midgut, and hindgut. One of the first gross morphologic dis-
tinctions is the rotation and distention of the posterior foregut
to begin differentiating the stomach just distal to esophagus.
The stomach is also separated from the esophagus by the newly
formed diaphragm from the developing abdominal cavity. At
the end of the fourth and beginning of the fifth week, the
stomach can be recognized as a fusiform dilation, which is ini-
tially oriented in the median plane. This primordial stomach
soon enlarges and broadens ventrodorsally. During the next
2 weeks, the dorsal border of the primordial stomach grows
faster than its ventral border (lesser curvature), demarcating
the greater curvature of the stomach. As the stomach enlarges,
it slowly rotates 90° in the clockwise direction around its lon-
gitudinal axis.1 The ventral border moves to the right, and the
dorsal border (greater curvature) moves to the left, changing
the position of stomach. The original left side moves to the
ventral surface, and the right side moves to the dorsal surface.
During rotation and growth of the stomach, the cranial region
moves to the left and slightly inferiorly, and its caudal region
moves to the right and superiorly. After rotation, the stom-
ach assumes its final position in the upper abdomen, with
its long axis almost transverse to the long axis of the body.
The rotation and growth of the stomach explains why the left
vagus nerve supplies the anterior wall of the adult stomach
and the right vagus nerve innervates the posterior wall of the
stomach.2
Arterial blood supply to the distal esophagus, stomach, and
proximal duodenum is derived from the branches of the celiac
axis. The stomach is drained by the left gastric vein, right gas-
tric vein, right and left gastroepiploic veins, and short gastric
veins. These veins have no valves and can provide collateral
blood flow when any portion of the portal system is obstructed.
Esophageal and gastric varices usually involve the left gastric
and short gastric veins.
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The lymphatic drainage from the stomach enters the tho-
racic duct via the celiac nodes. Ultimately, the lymphatic drain-
age enters the venous system in the neck at the junction of the
left internal jugular and left subclavian veins. Because of this
anatomic relationship, gastric malignancies in adults may pres-
ent with left supraclavicular lymph node metastasis.
Vagal and sympathetic fibers innervate the entire stomach
by about 9 weeks of gestation.3 Two major networks of nerve
fibers are intrinsic to the gastrointestinal tract: the myenteric
plexus (Auerbach’s plexus), which can be found between the
outer longitudinal and middle circular layers of muscle, and
the submucosal plexus (Meissner’s plexus), located between the
middle circular muscular layer and the mucosa. Collectively,
these neurons constitute the enteric nervous system. Catechol-
amines have been demonstrated in sympathetic fibers in Auer-
bach’s plexus by week 10 and in Meissner’s plexus by week 13.4
HISTOLOGY 
Development of fetal human gastric mucosa occurs very early
during fetal life. The first pit/gland structures are observed at 11
to 12 weeks of gestation. Between 11 and 17 weeks, the strati-
fied surface epithelium is replaced by a simple mucous colum-
nar epithelium, and gastric glands develop further. At this stage,
the progenitor zone of the pit/gland structure is already local-
ized in the isthmus,5 as in adult mucosa.1,6
The gastric mucosa is organized in vertical tubular units
consisting of an apical pit region, an isthmus, and the actual
gland region that forms the lower part of the vertical unit. The
progenitor cell of the gastric unit gives rise to all epithelial cells.
The mucus-producing pit cells migrate up toward the gastric
lumen, and acid-secreting parietal cells (oxyntic; oxys is Greek
for acid) migrate downward to the middle and lower regions of
the gland. Chief (zymogenic) cells secrete pepsinogen and pre-
dominate at the base of glands. Neuroendocrine cells, including
enterochromaffin cells (serotonin), enterochromaffin-like (ECL)
cells (histamine), and D cells (somatostatin), are also present at
the base of the gland.
The glands of the different anatomic parts of stomach are lined
with different types of cell (Figure 25-2). The cardiac glands are
mostly populated by mucus-secreting or endocrine cells. The
cardiac pits are irregular and shallow; the ratio of the lengths
of pits:glands is approximately 1:1. In the body of the stomach,
including the fundus, the glands are long and deep with straight
pits. The ratio of the lengths of pits:glands is approximately 1:4.
The gastric gland has parietal (oxyntic) cells that secrete hydro-
chloric acid and intrinsic factor, chief (zymogen, peptic) cells
that secrete pepsinogens, endocrine cells, and mucous neck cells.