The Metabolic Significance of Differential Absorption of Radiant Energy by Black and White Birds

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The Metabolic Significance of Differential Absorption of Radiant Energy by Black and White Birds THE METABOLIC SIGNIFICANCE OF DIFFERENTIAL ABSORPTION OF RADIANT ENERGY BY BLACK AND WHITE BIRDS FRANK HEPPNERl Department of Zoology University of California Davis, California The surface color of homeotherms is com- The undyed birds exposed to radiant energy monly thought to have two possible functions: consumed a mean of 2.00 ml OJmin, while concealment (Cott 1957) and communication the dyed birds exposed to radiant energy had ( Lorenz 1935; Tinbergen 1948). Hamilton a mean consumption of 1.44 ml Oz/min. By and Heppner ( 1967) proposed a third possible assuming a thermal quotient of 4.8 Cal/ml 02 function for dark, or black, homeotherm color- (King and Famer 1961), the heat production ation: the reduction of the metabolic cost of of the blackened birds was 6.9 cal/min, and homeothermy by maximization of absorption the heat production of the white birds was of radiant solar energy. This hypothesis pro- 9.6 Cal/mm, a difference of 2.7 cal/min. It is posed that the integument o,f black animals this difference that must be accounted for in absorbs more of the visible and near infrared calculating the difference in the amount of components of solar energy than does the radiant energy absorbed by the blackened and integument of lighter-colored animals, and white birds. that this greater absorption may permit them to reduce their heat 108s~when ambient tem- DETERMINATION OF TOTAL AMOUNT peratures are below their lower critical tem- OF ENERGY FALLING ON SURFACE perature. As reported earlier, the rate of energy falling Hamilton and Heppner reported that black- on the surface of a bird in the metabolism dyed, white Zebra Finches (Poephilu castan- chamber was 1.23 cal/cm2-min, as determined Otis) demonstrated a significant metabolic by a 59junction Eppley pyroheliometer. The economy over undyed birds when both were metabolism chamber, which was made of exposed to artificial sunlight while in a mod- Plexiglas, had a lid fitted with an ultraviolet erately cold (10%) metabolism chamber. I transmitting glass filter. During a metabolism now report the results of experiments relating test, the chamber was submerged in a water the metabolic differences between blackened bath, with 1 cm of water over the top surface and white birds observed by Hamilton and of the chamber lid. Heppner to the difference in amount of radi- The spectrum of the light energy that fell on ant energy absorbed by the black and white a bird in the chamber, after passing through the birds in our experiments, and discuss a physi- lid and the water over the lid, was determined cal mechanism whereby dark-colored birds by employing a Beckman DK-2 spectrophoto,m- might achieve a metabolic advantage over eter set up in a spectroradiometry configura- lighter counterparts. tion. A Sylvania DWY “Sun-Gun” lamp, which was the artificial sun in the metabolism experi- DETERMINATION OF DIFFERENCES ment, was mounted over the entrance port of IN ABSORBED ENERGY the spectrophotometer. Spectrum measure- Of paramount importance is the quantitative ments were then made (fig. 1) over the range relationship between the difference in require- 0.359-2.960~ of: (A) the light of the lamp ments for chemical energy of the white and alone, (B ) the light passing through the lid blackened birds, and the different rates of of the metabolism chamber, which was energy absorption from the artificial sun. This mounted between the lamp and the spectro- relationship can be determined by comparing photometer, and (C) the light passing through the difference in heat production, expressed both the lid and a l-cm layer of tap water in calories, between the white and blackened which was created by making a rim of mask- birds, with the difference in the amount of ing tape around the edge of the chamber lid energy absorbed at the surface of birds in the and then pouring water on the lid. The area two color conditions. under the curves can be expressed as the amount of energy falling on‘ a unit of surface 1 Present address: Department of Zoology, University of Rhode Island, Kingston, Rhode Island 02881. per unit of time. The lid had little effect on The Condor, 72 :50-59, 1970 50 METABOLIC SIGNIFICANCE OF ABSORPTION OF RADIANT ENERGY 51 100 90 1 70- : 2 60- t, YI so- : ; 40- WAVELENGTH OJ,, 08 I I I I I, - .3 .5 .7 .9 1.1 I.3 1.5 1.7 1.9 2.1 2.3 2.5 FIGURE 1. Spectrum measurements of Sun-Gun WAVELENGTH I/L, artificial sun (A, Sun-Gun; B, Sun-Gun through filter; C, Sun-Gun through filter and water). FIGURE 2. Absorbance characteristics of the sur- faces of white and black-dyed Zebra Finches (solid line, black bird; dashed line, white bird). the energy passing through it (curve 1B ) over the spectrum range measured, but the water cut off radiation beyond 1.406 p and shifted tions at the five locations were then averaged the energy peak to about 1.050 .U (curve 1C). at 0.100 ,Aintervals and a compo,site curve was This curve represents the spectrum of the hand drawn for each bird (fig. 2). light energy falling on the surface of a bird The shapes of both curves are generally sim- in the metabolism chamber. From the pyro- ilar, both birds tending to be “white” beyond heliometer measurements, it was known that 1.060 p, but there is a sharp difference in the energy falling on the bird had an absolute absorbance from 0.350 to 0.800 p, with the value of 1.23 cal/c&-min. This value is blackened bird absorbing more, as would be equivalent to the area under curve 1C. expected. If the fraction of energy absorbed by the In figure 3 the emission spectrum of the surface is determined for each wavelength, lamp is again shown with the curves produced and this fraction then multiplied by the frac- by multiplying the absorption fractions of the tion of total energy emitted by the source at black and white birds at 0.106 p intervals times that wave-length, a new curve will be ob- the height of the emission curve at the same tained, the area under which will be the wavelength. The area under these curves, energy absorbed by the surface. This area can compared with the area under the curve rep- be measured by a mechanical integrator (pla- resenting the energy falling on the bird, indi- nimeter), and compared with the area under cates what fraction of the energy falling on the curve of the emission spectrum, the ab- the bird is actually absorbed. solute value of which is known ( 1.23 cal/cms- The areas under the three curves were mea- min). The new value obtained will be the sured with a planimeter. It was found that energy absorbed by the surface of the bird. the area under the curve representing the en- The absorption characteristics of the birds ’ ergy absorbed by the black bird was 33.2 per surfaces were determined by setting up the cent of the total energy falling on the bird, Beckman DK-2 spectrophotometer in a spec- and the area under the curve for the white troreflectometer configuration. Live white bird, 16.0 per cent. Zebra Finches and newly dyed, blackened From the pyroheliometer measurements, Zebra Finches were anaesthetized with a gas the area under the curve representing energy anaesthetic and strapped to the sample port falling on the bird is equivalent to 1.23 cal/ of the integrating sphere of the spectrophotom- cm2-min. The black bird absorbed 33.2 per eter. The reflection and absorption charac- cent of this energy, or 0.41 cal/cm2-min, and teristics were compared to a white reference the white bird 16.0 per cent, or 0.20 cal/cm2- block of magnesium carbonate. Because there mm. were slight differences in absorption at differ- Finally, to determine the difference in total ent locations on the back, measurements were energy absorbed by the black and white birds, taken at five locations. The absorption frac- it is necessary to know the area upon which 52 FRANK HEPPNER 100 90 ;;e;;;Sfolling 80 . Energy absorbed 5 70 by black bird. ti Energy absorbed ; 60 by white bird. Ly g 50 f 40 4 / ‘, 1.23 cal/cm2Amin \ ff 30 20 WAVELENGTH (+L’ FIGURE 3. Energy falling on the surface of birds, and energy absorbed by black and white birds. the radiation is falling. This cannot be mea- conservative. The effective absorbing area sured precisely, because there are reflections of the birds is probably larger than the 14.7 from the bottom of the chamber that will ir- cm2 obtained by projecting the dorsal surface radiate the ventral surface of the bird, postural of the bird on a plane, since there will be some variations, and differences in feather erection reflections in the experimental universe that which will alter the effective area exposed to will impinge on all surfaces of the bird, direct radiation. An estimate can be obtained although the intensity of these reflections is by projecting the dorsal view of the bird on probably small. A larger surface area than a plane surface, and measuring the area of estimated here would increase the absolute this projection. The dorsal projected area of value of the difference in energy absorbed by three birds (excluding the tail) was 13.8, 15.2, black and white birds. Even with a conserva- and 15.2 cm2, or a mean of 14.7 cm2. These tive value, it appears that the difference in areas were obtained by measuring the pro- energy absorbed is sufficient to account for jected area with a planimeter.
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