On the Histogenesis of Gastric Glands. 515 
of a few zymogenic cells in the deepest part, but the neck segment, 
like the corresponding part of the fundic gland, has become split 
up into several tubules, which display the two cells characteristic 
of the neck of the fundic gland. While the aberrant development 
of zymogen granules is confirmatory, I lay most emphasis on the 
early parallelism in form-development, and especially on the 
presence, distribution and remarkable persistence of parietals 
in the cardiac tubules, as demonstrating their affinity with the 
fundie tubules. 
By all these findings we are inevitably brought to one conclusion 
with reference to the genetic relation of cardia and fundus. The 
cardia represents a part of the fundus which has undergone partial 
involution. The regressive changes have manifested themselves 
first in the general chronologic retardation of ontogenetic develop- 
ment in the left part of the old cardio-fundic region; second, in 
the phylogenetic disappearance of the fundic segment of the tubule 
with its zymogenic cells,—the curious, delayed appearance of 
zymogenic granules at one stage of the developing cardia repre- 
senting, probably, the last vestige of this segment; third, in the 
ontogenetic transformation of the cervix of the old fundus gland, 
with its mucous and parietal cells, into the tubule of the adult 
cardia with its mucous chief cell. This last process I have not 
seen, but it must needs occur, for we have the first and last terms 
of the series before us. 
The zone of fundic tubules, lying between undoubted cardia 
and the ordinary fundic zone, represents an intermediate stage 
in the regressive process. 
The facts of development, then, show that the cardiac glands 
are not primitive, but regressive structures. For the early stages 
do not resemble, in any way, the condition found in lower verte- 
brates, where, for example, even the fundus glands possess no 
definitely identified parietals. Moreover, no fact in the develop- 
mental history so much as suggests a possible derivation from 
the oesophagus. 
Thus, our embryological findings seem to confirm strongly 
the conclusion as to the origin of the cardiac glands reached by 
Bensley (’02), working from the standpoint of adult comparative 
