The secretion of glucocorticoids (GCs) is a classic endocrine response to stress. Despite that, it remains controversial as to what purpose GCs serve at such times. One view, stretching back to the time of Hans Selye, posits that GCs help mediate the ongoing or pending stress response, either via basal levels of GCs permitting other facets of the stress response to emerge efficaciously, and/or by stress levels of GCs actively stimulating the stress response. In contrast, a revisionist viewpoint posits that GCs suppress the stress response, preventing it from being pathologically overactivated. In this review, we consider recent findings regarding GC action and, based on them, generate criteria for determining whether a particular GC action permits, stimulates, or suppresses an ongoing stressresponse or, as an additional category, is preparative for a subsequent stressor. We apply these GC actions to the realms of cardiovascular function, fluid volume and hemorrhage, immunity and inflammation, metabolism, neurobiology, and reproductive physiology. We find that GC actions fall into markedly different categories, depending on the physiological endpoint in question, with evidence for mediating effects in some cases, and suppressive or preparative in others. We then attempt to assimilate these heterogeneous GC actions into a physiological whole. (Endocrine Reviews 21: 55‐ 89, 2000)

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How do glucocorticoids influence stress responses? Integrating permissive, suppressive, stimulatory, and preparative actions.

Publisher Summary This chapter reviews the present empirical and theoretical work on antipredator decision making. The ways in which predators influence the behavioral decisions made by their prey is now the subject of a large and growing literature. Some notable recent advances include clear demonstrations that antipredatory decision making (1) may influence many aspects of reproductive behavior, (2) has demonstrable long-term consequences for individual fitness, and (3) may influence the nature of ecological systems themselves. There have also been many advances in the theory of antipredator behavior, which should provide a sound conceptual basis for further progress. Attention is needed for further work on the effects that predator and prey have on the other's behavioral decisions. The range of reproductive behaviors influenced by the risk of predation also requires much more investigation. Work on the long-term costs of antipredator decision making needs more empirical documentation and greater taxonomic diversity. Work on the ecological implications of antipredatory decision making has only scratched the surface, especially with regard to population-level effects and species interactions. Theoretical investigations should also play a prominent role in future work.

Stress and Decision Making under the Risk of Predation: Recent Developments from Behavioral, Reproductive, and Ecological Perspectives

https://www.healthpromoting.com/sites/default/files/FastingMolecularMechanisms.pdf

Fasting: Molecular Mechanisms and Clinical Applications

https://www.webpages.uidaho.edu/fish511/Readings/Readings%202010/Romero_2009%20%20Allostasis.pdf

The Reactive Scope Model - a new model integrating homeostasis, allostasis, and stress.

Actions of glucocorticoids at a seasonal baseline as compared to stress-related levels in the regulation of periodic life processes.

During long-term fasting in birds and mammals, protein utilization initially decreases (phase I), is thereafter maintained at a low value (phase II), and then further increases (phase III). To delineate hormonal and biochemical changes responsible for these modifications, the effect of food deprivation for 50 days was studied in 6 male king penguins captured at the beginning of their natural breeding fast. During phase II, both rate of mass loss and plasma uric acid concentration remained at low levels, whereas plasma beta-hydroxybutyrate concentration increased. In phase III there was by contrast a 2.5-fold increase in the rate of mass loss, an eightfold increase in plasma uric acid, and an 80% drop in plasma beta-hydroxybutyrate. Plasma corticosterone was low and steady in phase II and increased three times in phase III. During the overall fast, there were no significant variations in plasma insulin, but there was a fourfold increase in plasma glucagon and a decrease in plasma thyroxine and triiodothyronine. These findings suggest that protein sparing (phase II) requires low levels of corticosterone, insulin, and thyroid hormones, whereas the further increase in protein utilization (phase III) is due to an increase in plasma corticosterone. The high plasma glucagon concentration in phase III is presumably responsible for a transient increase in plasma glucose observed at this stage; such increase in glucagon could enhance gluconeogenesis from amino acids.

Fasting in king penguin. I. Hormonal and metabolic changes during breeding

The stress that might result in animals from the routine handling that most experimental studies involve, e.g., weighing, injecting, and blood sampling, is usually assumed to be minimal when the animals look quiet. However, the intensity of this stress remains largely ignored. We have developed a system that allows blood samples to be taken from freely behaving geese without entering the animal room. In these entirely undisturbed geese, the humoral indexes of stress, i.e., blood levels of catecholamines, corticosterone, and lactate, were as low or even lower than the lowest values previously reported for birds. Remarkably, the mean basal values for epinephrine and norepinephrine were 90-fold and 5-fold, respectively, below the lowest values in the literature. Stress-induced variations in pH that would have concealed detection of nutrition-induced changes in pH were eliminated. In contrast, even though the birds looked quiet during a short 5-min routine handling procedure, to which they had been accustomed for weeks, there was a dramatic increase in the level of humoral indexes of stress. These increased severalfold within only 2 min, and the return to initial values could take up to 1 h. Acid-base balance was also disrupted. Thus, in studies on animals, the absence of stress cannot be deduced from only behavioral observations. Only a system for taking blood without human interference may enable stress-free investigations.

Stress in birds due to routine handling and a technique to avoid it

During seasonal cycles in ground squirrels, as in many other species, there are periods of spontaneous loss of appetite, very low calorie intake and a 30% loss in body mass. Measurements of nitrogen balance during early and later stages of the mass loss phase of the cycle (-1.2 +/- 6.7 and +13.1 +/- 8.8 mg/24 h, respectively) showed a total sparing of protein, indicating a selective use of fat. However, when no food at all was available, nitrogen balance was negative (-45 +/- 5 mg/24 h). Provided that they have access to some food, ground squirrels are therefore able to compensate for any protein utilization, while at the same time selectively losing large amounts of fat. It appears that a factor related to spontaneous reduction in food intake enables these animals to achieve the total sparing of protein that eludes dieting humans on comparably low caloric intakes.

Protein sparing on very low calorie diets: ground squirrels succeed where obese people fail.

Plasma glucose and insulin levels were measured in chronically catheterized, freely moving, undisturbed geese, which were offered a free standard meal after an overnight fast. The insulin level markedly rose within the first minute after the start of food ingestion, whereas plasma glucose did not increase. This early insulin response was not correlated with the size of the meal. In contrast, both postabsorptive insulin response and plasma glucose changes were dependent on meal size. When a small amount of food (2-6 g) was eaten, insulin returned to basal level within 30 min, whereas plasma glucose remained unchanged. Larger meals (15-20 g) maintained plasma insulin at a higher level and induced a sustained rise of plasma glucose. These results indicate that there is a cephalic phase of insulin secretion at the beginning of the meal in birds as previously described in mammals.

H. Karmann

Papers

Fasting in king penguin. I. Hormonal and metabolic changes during breeding

Stress in birds due to routine handling and a technique to avoid it

Protein sparing on very low calorie diets: ground squirrels succeed where obese people fail.

Early insulin response after food intake in geese