72 The Endocrine Organs 



it than normally. He states that in the guinea-pig the blood can be shown 

 to contain an excess of adrenalin after the sugar puncture. A. Mayer 

 found that Bernard's puncture is not effective after removal of the supra- 

 renals. Macleod and Pearce observed that adrenalin produces hyper- 

 glycemia even after division of the hepatic plexus (which is not the 

 case with the hyperglycemia caused by stimulation of the splanchnics), 

 and that only when the suprarenals are intact is it possible to produce 

 hyperglycemia by stimulation of the nerves of the liver. Adrenalin is 

 therefore in some way necessary to the activity of these nerves : possibly 

 by rendering them more sensitive. It is stated that the effect of adrenalin 

 on the liver glycogen can be prevented by atropine (Doyan and Gautier). 

 Ergotoxine also antagonises this action of adrenalin (see p. 64). 



As already stated, if the liver is rendered free from glycogen, adrenalin 

 injection will not only produce hyperglycemia and glycosuria as before, 

 but may, even in fasting animals, cause glycogen to be stored : this is 

 associated with the fact that the metabolism of proteins is increased under 

 the influence of adrenalin, and the protein is in this case the source of 

 the carbohydrate which is formed (Noel Paton). On the other hand, 

 0. Schwarz found the liver glycogen to disappear in rats in which the 

 suprarenals had been removed, although it could be restored by liberal 

 feeding with glucose and cane sugar (but not by levulose or starch). 

 If the glycogen of the liver is got rid of by phloridzin, administration 

 of adrenalin does not increase the glycosuria (A. I. Ringer). 



With the Pancreas. — Pemberton and Sweet found removal of the supra- 

 renals to cause a flow of pancreatic juice which is stopped by injection of 

 adrenalin, although under normal conditions the flow of pancreatic juice 

 seems not to be affected by the intravenous injection of adrenalin, at least 

 in moderate doses. But it is the internal secretion which is mainly 

 affected by this hormone. Herter and Wakeman found that the mere 

 swabbing of the exposed pancreas with adrenalin solution provokes 

 intense glycosuria, comparable with that produced by extirpation of the 

 organ. The glycosuria and the hyperglycemia caused by pancreatic 

 extirpation is, however, increased, as several observers have found, by 

 adrenalin, so that the whole of its effect on carbohydrate metabolism 

 is not exerted through the pancreas, but part must be directly on the 

 liver. We have, in fact, already seen that adrenalin has a direct effect on 

 the glycogen storage of the liver. But this again is influenced by the 

 internal secretion of the pancreas. Thus Zuelzer failed to get adrenalin 

 o-lycosuria when he injected pancreas extract at the same time as the 

 adrenalin ; on the other hand, he states that tying the suprarenal veins is 

 effectual in preventing glycosuria pancreatopriva. These experiments 

 show that carbohydrate metabolism is in some way regulated by the inter- 

 action of the suprarenals and pancreas upon the liver cells. Graham Lusk, 

 however, from a study of the respiratory quotient in a glucose-fed dog 

 to which adrenalin was administered, comes to the conclusion that this 



