Plasma Metabolite Levels Predict Individual Body-Mass Changes in a Small Long-Distance Migrant, the Garden Warbler

Abstract

-Change in body mass is an important measurement in many studies addressing changes in energy stores or condition. Usually, change in body mass is measured in birds caught twice, but this has a number of drawbacks (e.g. low number of retraps, retraps not representative of all first captures, adverse effects of first capture on body-mass development). Therefore, we investigated whether plasma metabolites correlate with body-mass change, and which metabolites could be used to predict body-mass change in birds caught once. In an experiment, 20 Garden Warblers (Sylvia borin) were given different amounts of food to induce stable, increasing, and decreasing body masses. Most of the eight plasma metabolites we examined were significantly correlated with the change in body mass between early morning and midday, the time of blood sampling, but not with body mass or various measures of activity. Metabolites that are known to characterize resorption were elevated in birds gaining body mass and metabolites characteristic of fasting were elevated in birds losing body mass. Triglycerides and ,B-hydroxy-butyrate together explained 61% of the variation in body-mass change (triglycerides alone 44% and ,3-hydroxy-butyrate alone 51%). These metabolites may be used to predict body-mass change in birds caught once, provided that the reliability and sensitivity of this method are checked in field tests. Received 27 September 1993, accepted 30 January 1994. BIRDS MAY FACE DEFICITS in their energy balance during the nonactive period of the day, during periods of several days, or even over a series of weeks. They usually build up energy stores in anticipation of such periods and as an adaptive compromise to different selective pressures (e.g. Lima 1986, Lehikoinen 1987, Moreno 1989, Ekman and Hake 1990, Rogers and Smith 1993). This results in fluctuating energy stores between foraging and nonforaging periods (usually day and night) superimposed on longer periods of net energy storage and net energy expenditure (King 1972). For instance, cold and short winter days, bad weather, incubation, feeding young, and migration may all be associated with decreasing energy stores and preceded by energy storage. Body mass is a crude but widely used estimate of energy stores because it is easy and harmless to measure. If corrected for size, body mass may be used as an indication of condition (e.g. Blem 1990). Changes in body mass may give a fairly accurate picture of the fluctuations in energy stores (e.g. Rogers and Rogers 1990). The measurement of changes in body mass of free-living birds is important in a variety of contexts. For instance, it may allow the assessment of the effects of food availability on energy storage in different habitats, under different weather conditions, and under different degrees of competition or predation (e.g. Bibby et al. 1976, Bibby and Green 1983, Buchanan et al. 1985, Lima 1985, Lindstrom 1990, Moore and Yong 1991). Body-mass changes may indicate periods of constraints and periods of adaptive mass variations during the annual cycle (Moreno 1989). Evaluation of body-mass changes may allow study of the pattern and environmental influences of energy storage during stopovers by migrants (e.g. Bibby et al. 1976, Mehlum 1983a, b, Biebach 1985, Biebach et al. 1986, Alerstam and Lindstrom 1990). In free-living birds, changes in body mass generally are measured in birds caught at least twice, usually at the same place. This, however, has a number of drawbacks: (1) changes in body mass are measured only in the subsample of retraps and this may reduce the sample size drastically, especially when trap shyness is a factor; (2) a certain group of birds may not be recaught and, thus, the sample of retraps may not be representative (e.g. Bibby et al. 1976, Biebach et al. 1986, Veiga 1986, Winker et al. 1992); and (3) first capture may have an adverse