Test, Rejection, and Reformulation of a Chemical Reactor–Based Model of Gut Function in a Fruit‐Eating Bird

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Abstract
We explored modulation of retention time in cedar waxwings (Bombycilla cedrorum) by feeding them diets varying in hexose concentration. Our goals were to (1) test three predictions of a chemical reactor-based model of how guts might respond optimally to diet shifts; (2) determine whether modulation of retention time can occur quickly, thereby facilitating rapid changes in diet; (3) tease apart the relative influence of ingestion rate and nutrient concentration on retention time; and (4) examine the degree of axial mixing in the intestine and its relationship with retention time. The model's predictions were rejected: mean retention time did not decrease, ingestion rate did not increase, and glucose assimilation efficiency did not decrease with increased hexose concentration of the diet. Instead, birds displayed maximal intake rate at intermediate sugar concentration, and mouth to cloaca mean retention times increased with hexose concentration. Significant modulation of retention time occurred quickly, within 3 h of exposure to a different diet. Birds did equally well in terms of total energy assimilated on diets differing 3.3-fold in hexose concentration (from 500 mmol/L to 1660 mmol/L) but showed reduced intake when fed food with low hexose concentration (110 mmol/L). Far more variation in retention time was explained by direct effects of ingestion rate than by direct effects of hexose concentration. Finally, a gut dispersion index that measured degree of axial mixing was positively correlated with mean retention time, indicating that higher retention times are accompanied by increased axial mixing. We propose a modification of the assumptions of the original model. The resulting "osmotic constraint" model better captures the interaction between feeding rate and digestive function in fruit-eating birds.