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Metabolic and Behavioral Responses to Fasting in the White-Crowned Sparrow (Zonotrichia Leucophrys Gambelii) Author(S): Ellen D

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Metabolic and Behavioral Responses to Fasting in the White-Crowned Sparrow (Zonotrichia Leucophrys Gambelii) Author(S): Ellen D Metabolic and Behavioral Responses to Fasting in the White-crowned Sparrow (Zonotrichia leucophrys gambelii) Author(s): Ellen D. Ketterson and James R. King Reviewed work(s): Source: Physiological Zoology, Vol. 50, No. 2 (Apr., 1977), pp. 115-129 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/30152551 . Accessed: 22/08/2012 16:44 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp . JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to Physiological Zoology. http://www.jstor.org METABOLIC AND BEHAVIORAL RESPONSES TO FASTING IN THE WHITE-CROWNED SPARROW (ZONOTRICHIA LEUCOPHRYS GAMBELII)1 ELLEN D. KETTERSON2AND JAMES R. KING Laboratoriesof Zoophysiology,Washington State University, Pullman, Washington99164 (Accepted1/21/77) During winter, ground-feeding bird species may occasionally be forced to fast when weather conditions (e.g., snowstorms, icing) make food temporarily inaccessible. To describe the response of White-crowned Sparrows to fasting, we (1) investigated variations of body weight and temperature in fasting and fed birds confined in a cold room for up to 64 h, and (2) periodically measured oxygen consumption and locomotor activity of fasting and fed birds through a 39-h night/day/night cycle while simulating natural conditions of temperature and daylength. Our data indicate that the un- usual White-crowned Sparrow might survive 3 nights and 2 days without food at air temperatures near freezing, but typical individuals could not be expected to survive longer than 1 day and 2 nights. Males lose weight at a greater rate than females, but are able to endure fasting longer because they are initially heavier. This suggests that males may be more resistant to winter storms and food deprivation, and that this attribute may have contributed to the evolution of geographic variation in the sex ratio in this species. Physiological and metabolic responses to fasting included a nighttime decline in body temperature (ca. 3 C) that was greater than that observed in fed birds, and a 21% decline in oxygen consumption on the second night of the fasting cycle when compared to the first night. The drop in body temperature would be sufficient, in terms of the van't Hoff effect, to account for the decline of metabolic rate, but visual inspection of the fasting birds at night suggested that adjustments in ptiloerection and posture may also play a role in effecting the decline. During the day, locomotor activity was 1.5-10 times greater in fasted than fed birds. Oxygen con- sumption, however, did not differ, indicating (as one possibility) that fasting birds may be able to utilize heat generated by exercise to meet some of the cost of thermo- regulation. The short-term adjustment to starvation under natural conditions may thus include a moderately reduced energy requirement and a partial diversion of energy expenditure from thermoregulation to foraging. Several recent investigations (e.g., avian species that overwinter even in Fretwell 1972, Ivacic and Labisky 1973) temperate latitudes are periodically con- have emphasized the synergistic effects fronted by low environmental temper- of climatic and trophic stresses on the atures and by snow cover that pre- survival of homeotherms in extreme vents foraging or reduces its yield. environmental conditions. Individuals of Small-bodied ground-feeding passerines are likely to be especially vulnerable to IThis investigationwas supportedby grants from mortality under such conditions. We the National Institutes of Health (5T01 GM 1276) selected such a the White-crowned and National Science Foundation (BMS 72 02347). bird, For technical assistance, critical suggestions, or Sparrow (Zonotrichia leucophrys gam- stimulating discussions, the authors thank J. belii), as the subject of an investigation Boehme, G. S. Campbell, G. Doell, C. Maxwell, on metabolic and behavioral S. Mahoney,and J. Mugaas. focusing responses to simulated conditions of 2Present address: Department of Biological winter in low air Sciences, Bowling Green State University, Bowling severity, particular Green, Ohio 43403. temperatures and food deprivation. We 115 116 ELLEN K. KETTERSON AND JAMES R. KING chose this species not only because of mated from the ratio of the caloric its size and foraging characteristics, but values of fat stores and metabolic rate also because comparatively much is estimated from laboratory data (sum- known of its physiology and energy marized by King 1972; see also Chaplin metabolism (King 1964; Merkel 1966; 1974; Ketterson and Nolan 1976). De Jong 1976; Robinson, Campbell, and King 1976), and because it is a METHODS species that displays geographic varia- Our observations consist of measure- tion in its sex ratio winter. during ments of (1) body temperature and rates are Northerly populations predominantly of weight loss in fasting birds (experi- the decreas- male, proportion gradually mentals) and birds feeding voluntarily southward and Me- ing (King, Farner, (controls) when both were kept indoors waldt We that this dif- 1965). thought at constant temperature, and (2) oxygen ferential distribution of the sexes in consumption and locomotor activity of winter be on the might interpretable fasting birds and their controls when basis of sexual differences in to response both were subjected to varying thermal cold stress and starvation. conditions simulating their natural win- of Investigations weight loss during ter environment on overcast, windless food called fast- deprivation (hereafter days and nights at the northern limit of and the survival time of ing) fasting their winter range (Mahoney 1976). All individuals in the laboratory are not new birds were winter acclimatized in out- Latham (Kendeigh 1934, 1945; 1947; door aviaries and had been in captivity Koskimies Kabat 1950; Jordan 1953; for 1-3 yr. et al. 1956; Hine and Flakas 1957; Tester and Olson 1959; Hanson and CONSTANT AIR TEMPERATURES (COLD ROOM) Kossak 1963; Brenner 1965; Brenner and Treatment.-To determine how long Malin 1965; Ivacic and Labisky 1973). fasting could be endured and the ac- From these investigations we know that companying changes of body weight and heavier individuals (and species) tend to body temperature, 32 White-crowned endure fasting longer than lighter ones Sparrows were denied food (but given (Kendeigh 1945; Ivacic and Labisky water) until considered near death, 1973) and that some species exhibit while weight and body temperature sexual differences in survival time when were periodically recorded. Control was fasting (Kendeigh 1945, Latham 1947; provided by 12 birds supplied with Jordan 1953; Ivacic and Labisky 1973). chick starter mash and millet seed as Energy metabolism during prolonged well as water, but otherwise similarly fasting (greater than 1 night) has been treated. We assigned subjects randomly studied frequently in domesticated birds to these categories. All birds were sub- (e.g., Philips, Ashworth, and Brody jected in a cold room to the following 1932; Henry, Magee, and Reid 1934; microclimatic conditions: air tempera- Benedict and Lee 1937; Dukes 1937; ture range, 6.5-8 C; windspeed, 0.09- Barott et al. 1938; Smith and Riddle 0.18 m/s; wall temperature, 4.3-4.8 C; 1944; Infantellina and Rubino 1951), relative humidity, 60%-63%; photo- but much less frequently in wild species period, 10L:14D, lights on at 07:00. (Koskimies 1950; Ivacic and Labisky Observations were made between Janu- 1973). In the field, the endurance of ary 1 and March 1, 1975. fasting by overwintering small passerines Around noon on day 1 of an observa- (weighing less than 30 g) has been esti- tion sequence (N = 11), four White- EFFECTS OF FASTING IN WHITE-CROWNED SPARROW 117 crowned Sparrows were taken from an longed fasting in thermally realistic outdoor aviary, weighed (nearest 0.1 g), conditions. Nevertheless, in spite of our measured (wing length, chord), and intentions to spare their lives, two birds placed in cages in the cold room. Four were dead at the end of the sequence; hours later (ca. 16:30) they were re- four died within 48 h later despite special moved one at a time and transported to treatment consisting of glucose feedings, an adjoining room where we determined confinement at room temperature, and body temperature by inserting a Schult- exposure to prolonged photoperiod to heis quick-responding thermometer into allow voluntary feeding at odd hours; the cloaca for 15 s (98% response time in a 10 lived but were given the same special water bath). Body temperature measure- treatment just described, and in our ments were made as rapidly as possible opinion five of the 10 would have died in an effort to minimize the influence of without it; and 16 were able to feed disturbance (Southwick 1973). The usual themselves at the end of the sequence interval between the first and fourth when subjected to the experimental tem- readings was 3 min. The birds were then perature and photoperiod. Of these last weighed again and returned to the cold 16, three were certainly terminated too room. We considered
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