PHYSIOLOGY OF PEPSIN-FORMING GLANDS. 
679 
Nussbaum suggested that the pepsin arose from the oesophageal gland granules. He 
found a correspondence between the number of granules in the gland-cells and the 
amount of ferment contained by the glands. 
Sewall and myself, whilst differing from Nussbaum as to the times of increase and 
decrease of granules, came nevertheless to the conclusion that the granules were con¬ 
nected with the formation of ferment. To this conclusion we came partly on general 
grounds, partly by comparing Gjuutzner’s results on the times of increase and decrease 
of pepsin with our own on the times of increase and decrease of granules. 
I have made some experiments- to determine this point. Since the oesophageal 
glands are affected during digestion to such different degrees in different Frogs, I have 
paid no especial attention to the alteration in pepsin-content of the oesophagus which 
occurs in successive hours after feeding. 
I proceeded in the following manner: hungry Frogs and Frogs fed with worm or 
sponge were taken, and (1) the granularity of the gland-cells observed ; (2) the relative 
amount of pepsin contained by equal weights of dried oesophagus estimated. 
The Frogs were taken as much as possible alike in size and general condition. Part of the oesophagus 
was observed fresh, part after treatment with osmic acid and alcohol—the rest of the mucous membrane of 
the oesophagus was dried and a definite quantity weighed out, this was treated with HOI. 0'2 per cent.— 
3 cub. centims. for O'Ol grm.—for twenty-four hours. The filtrate from this was tested for pepsin in 
the ordinary manner by Geutzner’s colorimetric method. Only those specimens which were analysed for 
pepsin on the same day were compared. 
In some cases of slight difference in amount of granules the results were not constant, 
this I attribute to the imperfection of the method ; equal weights of dried oesophagus 
do not necessarily contain equal weights of secretory gland-cells. 
In all cases where there was a marked difference in the amount of granules there 
was also a marked difference in the amount of pepsin. 
Hence, then, the greater the amount of granules the greater is the amount of pepsin 
to be obtained from the glands. I think we can fairly conclude that the granules give 
rise to the ferment. 
We can now consider the question, Do the oxyntic glands form pepsin 1 I have 
suggested above that Swiecicki was inclined to attribute the pepsin found in the 
gastric mucous membrane to an absorption of the pepsin secreted by the oesophageal 
glands. In this I cannot agree with him. The amount of ferment found in the 
mucous membrane of the stomach is far too great to allow of any explanation except 
that it is formed by the gastric glands. It is true that when equal iveights of the 
mucous membrane of the oesophagus and stomach are taken and their relative pepsin- 
content compared, the amount is found to be considerably greater in the former than in 
the latter, yet the latter contains a not inconsiderable amount. Thus in the experi¬ 
ment given below an acid extract of the gastric mucous membrane is made in the pro¬ 
portion of 1 grm. of dried tissue to 2,000 cub. centims. of hydrochloric acid—0'2 per 
cent. Of this acid-extract 18 cub. centims. are added to 2 cub. centims. of swollen 
