All animals face the possibility of limitations in food resources that could ultimately lead to starvation-induced mortality. The primary goal of this review is to characterize the various physiological strategies that allow different animals to survive starvation. The ancillary goals of this work are to identify areas in which investigations of starvation can be improved and to discuss recent advances and emerging directions in starvation research. The ubiquity of food limitation among animals, inconsistent terminology associated with starvation and fasting, and rationale for scientific investigations into starvation are discussed. Similarities and differences with regard to carbohydrate, lipid, and protein metabolism during starvation are also examined in a comparative context. Examples from the literature are used to underscore areas in which reporting and statistical practices, particularly those involved with starvation-induced changes in body composition and starvation-induced hypometabolism can be improved. The review concludes by highlighting several recent advances and promising research directions in starvation physiology. Because the hundreds of studies reviewed here vary so widely in their experimental designs and treatments, formal comparisons of starvation responses among studies and taxa are generally precluded; nevertheless, it is my aim to provide a starting point from which we may develop novel approaches, tools, and hypotheses to facilitate meaningful investigations into the physiology of starvation in animals.

Starvation physiology: reviewing the different strategies animals use to survive a common challenge.

Morphometric estimates of body condition are widely used by ornithologists, but which estimates work best is a matter of debate. We review morphometric approaches (body mass, ratio and residual condition indices, predictive regression models, fat scoring, and abdominal profiles) for estimating body condition (defined as fat mass) in birds. We describe the strengths and weaknesses of each approach. Across diverse indices and species (~200 estimates total), the mean r
 2 relating condition indices to mass of body fat was 0.55, and 64% of the r
 2 values were greater than 0.50. But despite their generally good performance, condition indices sometimes perform poorly (i.e., r
 2 is low). The data indicate that: (1) no single index was clearly best, (2) on average body mass alone, fat scores, and predictive multiple regression equations explained slightly more than 50% of the variation in fat content, (3) on average, ratio and residual indices explained slightly less than 50% of the variation in fat content, and (4) body mass alone, a variable that can be easily and reliably measured, is as good or nearly as good an indicator of fat content as any other condition index. We recommend that: (1) morphometric indicators of condition be empirically validated, (2) researchers publish their body composition data in sufficient detail that they can be used in future analyses exploring the relative merits of different condition indices, and (3) multiple regression directly on measured traits be used instead of condition indices whenever the condition index is not empirically validated.

Morphometric indices of body condition in birds: a review

The validity of the haematocrit or packed cell volume as an indicator of condition in wild birds has recently been questioned. We reviewed over 300 published papers on haematocrit values for wild birds. These studies show that changes in haematocrit could be caused by a number of different natural factors that include age, sex, geographical elevation, energy expenditure, parasitism, nutrition and genetics. Haematocrit also increased with age from hatching, due to increased erythropoiesis, so that adult birds generally have greater haematocrit values than nestlings or juveniles. Haematocrit values were either independent of elevation or increased with elevation. A meta-analysis of 36 studies showed no difference in haematocrit between the sexes. Relationships between haematocrit value and both energy expenditure and parasitic infection vary between studies. In temperate climates, haematocrit tended to be higher in winter than in summer, which may be due to dehydration or increased oxygen demand caused by thermogenesis, moult or acquisition of reproductive status. Our review indicates that the use of haematocrit as a sole indicator of condition or health could lead to incorrect conclusions if natural factors that can affect haematocrit are not taken into consideration.

Sources of variation in haematocrit in birds

The T-cell-mediated immune response (CMI) of birds, measured with the phytohaemagglutinin skin test, is in most cases positively correlated with their body mass. This correlation, however, does not...

Effects of experimental food restriction and body-mass changes on the avian T-cell-mediated immune response

Hypotheses of parasite-mediated sexual selec- tion (PMSS) propose that elaborate male ornaments have evolved due to female preferences. Females would benefit from mating with more ornamented males if males' ornamentation signals their health status and ability to provide parasite resistance genes for the offspring. Carotenoid-based plumage coloration of birds has been hypothesised to honestly reflect an individual's health status due to trade-off in allocation of carotenoids between maintenance and signalling functions. The prediction of this hypothesis, namely that individuals with brighter plumage are able to mount stronger immune responses against novel antigens and reveal generally better health state, was tested in captive male greenfinch- es (Carduelis chloris). Greenfinches with brighter yellow breast feathers showed stronger humoral immune re- sponse against novel antigen (SRBC) while no relation- ship between plumage coloration and an estimate of cell- mediated immune responsiveness (PHA response) was detected. Elaborately ornamented individuals had better general health state as indicated by the negative correla- tions between plumage brightness and heterophil haemo- concentration. Consistent with the concept of PMSS, these results suggest that carotenoid-based plumage coloration in greenfinches honestly signals immunocom- petence and health status.

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Carotenoid-based plumage coloration of male greenfinches reflects health and immunocompetence

Populations of scavenging seabird species in the North Sea may fluctuate with an artificial food source: the availability of fishery waste. To document this impact, it is necessary to assess the birds' nutritional status during periods with decreased fishing activity. Reference data for this purpose was collected from 22 herring gulls investigated during laboratory fasting. After 6 d of food deprivation and body mass losses exceeding 15%, the first birds entered starvation phase 3. Comparatively, this is a rather weak fasting capacity. Plasma levels of total protein and thyroid hormones decreased and β‐hydroxybutyrate increased with fasting duration. The leucocyte proportions were shifted from lymphocytes to heterophils. After 3 d of refeeding, most of the fasting changes were reversed. Plasma enzyme activities increased and hematocrit, hemoglobin, and erythrocyte numbers decreased in both fasting and control birds, most likely as a result of experimental stress and repeated blood sampling. Gluco...

https://www.jstor.org/stable/10.1086/316675

Ommo Hüppop

Papers

The Influence of Fasting on Blood and Plasma Composition of Herring Gulls (Larus argentatus)