Metabolic Rate of Laysan Albatross and Bonin Petrel Chicks on Midway Atoll!
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Pacific Science (1984), vol. 38, no. 2 © 1984 by the University of Hawaii Press. All rights reserved Metabolic Rate of Laysan Albatross and Bonin Petrel Chicks on Midway Atoll! 2 2 GILBERT S. GRANT ,3 AND G. CAUSEY WHiTTOW ABSTRACT: The resting metabolic rates ofLaysan albatross and Bonin petrel chicks of known age were measured on Midway Atoll in the North Pacific Ocean. The mass-specific metabolism peaked at hatching and then declined to adult levels in Laysan albatross nestlings . The mass specific metabolism of hatchling Bonin petrels was similar to that of adults, but it tripled shortly after hatching. Fasting and feeding episodes affected day-to-day changes in petrel chick metabolism. THE RESTING METABOLIC RATES of chicks rep METHODS resent the major part of their energy expen diture, as they do not fly and their level of The metabolic rates of Laysan albatross activity .is generally low (Blem 1978). The chicks ofknown age on Sand Island, Midway metabolic rate increases as the chick grows Atoll (28°13' N, 177°23' W) were measured in but the relationship between metabolic rate a building immediately adjacent to the nesting and body mass var ies during growth, the par sites. The chicks were placed in a small plexi ticular pattern of variation depending on a glas chamber or in an air-tight wooden box number of factors (Ricklefs 1974). In the (0.7 m on all sides), depending on the size of Procellariiformes, which have relatively long the chick. In the latter instance, a plexiglas nestling periods, the metabolic rate of grow cover allowed observation during a series of ing chicks is known only for Leach's storm measurements. Ambient air was drawn petrel, Oceanodroma leucorhoa (Ricklefs, through the metabolic chamber by means ofa White, and Cullen 1980a). The present note vacuum pump. The flow rate (1-5 stems from the opportunity to study the re liters ' min - 1) was adjusted to obtain a de lationship between metabolism and growth in crease of approximately 1 percent in the O 2 two other procellariiform species that differed concentration from the air inlet to the outlet. greatly in body size, and for which the meta The birds were weighed, to the nearest gram, bolic rates of the developing embryos are on a Terraillon balance, immediately after the known (Pettit et aI. 1982a, 1982b). The two final metabolic measurements. All albatross species studied (Laysan albatross, Diomedea metabolic measurements were made between immutabilis; Bonin petrel, Pterodroma hypo 1000-1800 hr, 21 January-14 Ma y 1981. leuca) share the same nesting sites in the Chamber temperatures ranged from 19.3 Northwestern Ha waiian Islands. to 26.0°C during these measurements. Mea surements could be made only until the 107th day of nestling life, when we had to leave Midway; the birds fledge at 165 days. 1 Supported by National Science Found ation Grant Metabolic measurements on Bonin petrel No.PCM 76-12351-AOIadministered by G. C. Whitlow. This study was supported also by the Sea Grant College chicks of known age were made in a small Program (NIjR-14). Manu script accepted 14 December plexiglas chamber (kept in the building ad 1983. jacent to the nesting site) at chamber tempera 2 University of Hawaii, John A. Burns School of tures of 21.3 to 29.0°C. All measurements Medicine, Department of Physiology, and the PBRC Kewalo Marine Laboratory, Honolulu , Hawaii 96822. were between 1000-1900hr, 2 March to 19 3Present Address: North Caro lina State Museum of May 1981. The flow rate (ca. 200 Natural History, P.O. Box 27647, Raleigh, N.C. 27611. 500 cm": min -1) was also adjusted to achieve 170 iiLUUaotJiCUl!iJIlLia::mtillUuiii, j·P 6 . 3 ' i i i , t :::tlrt:,W_ .l..IIWUlD Petrel Chick Metabolic Rate-GRANT AND WHITTOW 171 a decrease of approximately I percent in the measured by Pettit et al. (1982a). The O 2 concentration between the inlet and out RQ of albatross chicks averaged 0.78 flow from the chamber. Petrels were weighed (Figure I). The mass-specific metabolism on a torsion balance (OWL 5) to the nearest (V02 -;- mass) peaked near hatching 1 0.01 g, immediately after the metabolic (0.77 cm ' °2 ' g -l . hr - ) and declined measurements. Measurements were made steadily, reaching adult levels at approxi until a few days prior to fledging (mean age mately 75 days (Figure I). of fledging = 82 days; Pettit, Grant, and The resting metabolism and mass of petrel Whittow 1982). chicks from hatching to 76 days ofage (within Chicks of both species settled down within a week offledging) are shown in Figure 2. The a few minutes and remained calm in the meta RQ of petrel chicks averaged 0.76. Mass bolic chambers during a series of measure specific metabolism tripled between hatching ments that typically lasted 3-5 hours. and 2 days of age and declined to adult levels Metabolic values obtained for each individual at 20 days of age (Figure 2). on a particular day were similar and were averaged. Gas samples were collected in glass sy DISCUSSION ringes, the barrels of which were coated with mineral oil, and analyzed with a Scholander The mass-specific oxygen uptake of micro-gas analyzer. All gas values are con the Laysan albatross hatchling was I-45 verted to standard temperature and pressure 0.77 em" °2 , g -l . hr - percent higher for the dry gas (STPO). Two methods were than the adult rate ofoxygen consumption of 1 used in the field to ascertain the accuracy of the Laysanalbatross (0.53 cm' °2 ' g-l . hr - ; the gas analyzer: (I) the composition of fresh Grant and Whittow 1983) and also greater air, measured by the gas analyzer, averaged than that of the adult wandering albatross 3 20.90 percent O 2 and 0.04 percent CO 2 (the (Diomedea exulans; 0.5 em O 2 • g - 1 • hr - 1; expected values), and (2) analysis of exhaust Brown and Adams 1984). The hatchling 1 gases from the combustion of ethanol in the of Bonin petrels (1.1 cm ' ' g -l . hr - ; \'0 2 °2 chamber. The respiratory quotient (RQ) of Pettit et al. 1982b) was similar to that ofadults 1 combusted ethanol should be 0.67. We (1.1 em:' °2 , g -l . hr - ; Grant and Whittow measured an ethanol RQ of 0.66 with the 1983). Mass-specific metabolism peaked to Scholander micro-gas analyzer, providing levels nearly three times hatchling and adult confirmation of its accuracy. The resting values during the first few days of nestling life metabolism was calculated from the mea in petrels but not in albatrosses. This funda ) sured oxygen consumption (\'02 and RQ. mental difference between the albatross and Chamber temperatures were measured with a petrel may be related to differences in the re calibrated Kane-May Limited Depen lative incubation periods and embryonic datherm instrument. growth in the two Procellariiformes (Pettit et al. 1982a, 1982b). According to Ricklefs (1974), the maximal mass-specific metabolic RESULTS rate occurs earlier in the precocial than in the The resting metabolism and mass of alba altricial chick. By this criterion, the Laysan tross chicks from hatching to 107 days of albatross is more precocial than is the Bonin age are presented in Figure I. Hatchling petrel. However, by other criteria (e.g., yolk (age < I day) albatross metabolism was content ofthe egg) petrels are considered to be measured by the open flow system and com semiprecocial (Nice 1962, Carey, Rahn, and pared to values obtained with the closed man Parisi 1980, Ricklefs, White, and Cullen ometric system used by Pettit et al. (I982a). 1980b) while albatross are semialtricial (Nice The \'0 2 for a hatchling measured via the open 1962). Clearly, additional information is 3 1 system average 190cm 0 2 ·hr- , compared needed on the metabolism during growth of 3 1 with a mean value of 159cm 0 2·hr- other species. 172 PACIFIC SCIEN CE, Volume 38, April 1984 Hatchling Fledgling 3600 • • • U) •• U) • ca 2400 • ~ + 1200 0 Embryo Ad utt Nestli ng • • • • • • - -- · . -- - • • ._-+ N 1000 CI .> .8 ~ \ , • • I• I ~ • • • • C¥ • • + ~ .7 • .6 u -u .6 eu N CI. CI • • U) .=- • I • • U) .3 + U) ca ~ 0 -4 0 25 75 125 165 Age (days) I F IGURE I. Mass (g), metabolism (Yo,; ern" °2 ' hr" "), RQ, and mass-specific metabolism (Vo,; em? °2 , g-I . hr - ) of Laysan albatros s embryos, nestlings, and adults in relation to age. Adult values (from Grant and Whittow 1983) are presented as mean (horizontal bar) ± two standa rd deviat ions (vertical bar). Embryo and hatchling values are from Pettit et al. (1982a). Curves are fitted by eye. The bro ken line indicates extrapolated values (see text). ,j, ! .I A AA • iii i ., I;;;; lA- M i., .. p i I .i ;.tti :; ,ga ;. #1 'kt:. Petrel Chick Metabolic Rate- G RANT AND WHITTOW 173 -6 0 12 24 36 48 60 72 Hatchling Fledgling 300 • • • • • • • • U') 200 U') fa • + ::E 1 0 0 0 Em btyo Nestling Adult • • • • • • • N 200 .-- c • .> t o .8 • • ••• •• • • • • ho-•• • •• • • • C¥ • • •• .7 + CI:: • . 6 • - 3 • • •• ....... .... ••• I • + o 12 24 36 48 60 72 Age (days) 3 FIGURE 2. Mass (g), metabolism (Yo, ; cm . °2 ' hr - ' ), RQ, and mass-specific metabolism (Yo,; ern" .°2 , g-I .hr- ') of Bonin petrel embryos, nestlings, and adults. Adult values (from Grant and Whittow 1983)are . presented as mean (horizontal bar) ± two standard deviations (vertical bar) . Embryo and hatching values are from Pettit et al. (1982b). Cur ves are fitted by eye. The broken line indicates extrapol ated values (see text). 174 PACIFIC SCIENCE, Volume 38, Apri11984 T ABLE I O XYGEN COST OF D EVELOPMENT IN THREE P ROCELLARII FOR M C HICKS TOTAL O 2 FLEDGLING OXYGEN COST COST OF CHICK MASS-HATCHLI NG PER GRAM OF DEVELOPMENT MASS CHICK* SPECIES (liters) (g) (liters-g - 1) Oceanodroma leucorhoa 209.3 t 40.51 5.17 Pterodroma hypoleuca 393.4 I77§,1I 2.22 Diomedea immutabilis 4,43 9 1,792 * '** 2.48 • Total O2 cost/fledgling mass- hatchling mass.