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Conductance and Structure of Eggs of Adelie Penguins, Pygoscelis Adeliae, and Its Implications for Incubation ’

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Conductance and Structure of Eggs of Adelie Penguins, Pygoscelis Adeliae, and Its Implications for Incubation ’ The Condor92304-312 0 The CooperOrnithological Society 1990 CONDUCTANCE AND STRUCTURE OF EGGS OF ADELIE PENGUINS, PYGOSCELIS ADELIAE, AND ITS IMPLICATIONS FOR INCUBATION ’ MICHAEL B. THOMPSON~ School of Biological Sciences,Victoria Universityof Wellington,P.O. Box 600, Wellington,New Zealand KENNETH N. GOLDIE Faculty of Science, Victoria Universityof Wellington, P.O. Box 600, Wellington,New Zealand Abstract. Structureand conductanceof eggshellsof Adelie Penguins,Pygoscelis adeliae, were studied to addressthe question:How is the eggshelladapted to cope with the extreme aridity of the Antarctic, and in particular, what is the significanceand function of the organic cuticleon the eggshell?Adelie Penguinshell structurehas a basicavian pattern with relatively straight, unbranched pores that are occluded by an organic cuticle on the outside of the shell. Mean shell conductance of 15.4 mg.day-l.Torr- ’ of six intact eggs is significantly lower than one would predict on the basis of fresh egg mass, and conductanceafter cuticle removal (18.6 mg day-l. Torr+) from three eggsis significantlyhigher. Theoretical consid- eration of tensions of 0, and CO, in intact eggsand cuticle-removed eggsindicates that, whereas 0, tensions inside the egg may not be stressfulto the embryo prior to hatching if the cuticle remained intact throughout incubation, CO, tensionsmay be. We concludethat the cuticle reduces loss of water during the early part of incubation, but that its erosion during incubation increasesconductance thereby maintaining CO, tensions close to levels expectedin eggsof this size. Calculationsof functional pore radius from conductancemea- suresfor cuticle-removed eggsare almost identical to real mean pore radius measuredfrom pore casts. Key words: Eggshell; conductance;pore structure:Adelie Penguin; Pygoscelisadeliae; incubation;egg; Antarctic. INTRODUCTION mour 1987). Conductance is modified by selec- The porosity of avian eggshellis extremely im- tion in such a way that desiccation does not en- portant becausethe avian embryo is isolatedfrom danger the embryo but that water loss during its environment by the shell, and the only ex- incubation is sufficient for proper air space for- changesduring incubation occur by diffusion of mation and the ability to supply sufficient oxygen gasesthrough pores in the shell (Wangensteen et and get rid of excesscarbon dioxide is not com- al. 1970/197 1). Oxygen is taken up and carbon promised. The eggshell must also be strong dioxide and water vapor given off during incu- enoughfor the brooding bird not to break it when bation. Shell conductance is one of the factors incubating, yet not so strong that the chick can- which determines the rate at which gasesmay be not break free (Ar et al. 1979). transmitted acrossthe shell and is the inverse of As gas exchange through the eggshell occurs resistance(Ar et al. 1974). Conductance depends, by diffusion, conductance is inversely propor- in part, on the number and structure of pores tional to the partial pressuregradient of the gas (Rahn and Ar 1974). Pore structure may change acrossthe shell (Paganelli 1980). Selection acts through incubation by embryonic erosion of the to modify conductance of eggshellsof birds at inner surface of the shell (e.g., Booth and Sey- high altitude, where partial pressure of oxygen and water vapor is very different from that at sea level, by acting on functional pore area (Wan- gensteenet al. 1974, Carey et al. 1983). Thus, it I Received 1 May 1989. Final acceptance1 February is clear that conductance of avian eggshells 1990. 2 Present address:Zoology (A08), School of Biolog- matches the physiological requirements of the ical Sciences,University ofsydney, N.S.W. 2006, Aus- egg with its normal incubation environment. tralia. As water moves through the shell as vapor (Ar 13041 ADELIE PENGUIN EGGS 305 TABLE 1. Physicalcharacteristics of six Adelie Penguineggs used in the study.(See text for calculationof mass,volume, and surfacearea.) Thickness measures are meansof the mean of four measuresat eachof 20 or 2 1 equidistantpositions between the blunt and pointedpoles of eachegg. Mass (g) Volume (cc) surfacearea (Crn~) Thickness(mm) x 117.5 108.66 110.97 14.8 0.57 SD 10.9 10.08 6.95 1.7 0.06 Range 102.0-l 30.0 94.37-120.26 101.09-118.97 12.7-16.6 0.47-0.64 et al. 1974) nest humidity is an important pa- ber and maintained in a constant temperature rameter in determining how much water is lost cabinet for 36 hr at 25.5”C. Three copper con- from an egg during incubation. Eggs incubated stantan thermocouples were placed adjacent to in conditions creating unusual vapor pressure the eggsin different parts of the chamber. Eggs gradients, e.g., high altitude (Rahn et al. 1977, were removed from the chamber and weighed to Carey 1980, Carey et al. 1983), buried (Seymour 1 mg on the electronic pan balance after 0.5, 1, and Ackerman 1980) dry (Rahn and Hammel 1.5, 2, 3, 6, 12, 24, 30, and 36 hr. They were 1982), or in wet environments (Ackerman and replaced within 30 set of removal. Temperature Platter-Rieger 1979, Sotherland et al. 1984) have was a constant 25.5”C in the first measurement conductancesdifferent from those of eggsof sim- of conductance and averaged 25.6 f O.l”C (n = ilar size from “normal” environments (i.e., in 30) in the second measurement. Rate of mass open nests,close to sealevel), and in which water loss fell until it was constant after 6 hr, so cal- loss is maintained within acceptable limits (ca. culations were based on the rate of movement 15%, Ar and Rahn 1980). of water vapor acrossthe shell (Ii&,,) over the Arguably, one of the driest environments in final 30 hr of measurement. the world in which birds breed is that of the Three of the eggswere than washed for 90 set Antarctic. The dry conditions result from ex- in Janola (active ingredient 31.5 g.l-I sodium tremely cold temperatures. Adelie Penguins, Py- hypochlorite; Reckitt and Coleman [N.Z.], goscelisadeliae, nest at Cape Bird on Ross Island Avondale, Auckland) at 42°C to remove the cu- adjacent to the Antarctic mainland in early No- ticle (Deeming 1987). Completeness of cuticle vember (Spurr 1975) when conditions are still removal was checked using a binocular dissect- very cold. A previous study showedthat eggshell ing microscope.The eggswere rinsed in tap water of this speciesis adapted to low humidities with at 50°C for several minutes and then in tap water a reduced shell conductance and fewer than pre- at room temperature. All six eggs(three washed dicted pores (Rahn and Hammel 1982). We fur- and three intact) were re-equilibrated to 25.5”C ther studied the significance of the organic cover overnight before being placed back into the bell of the shell (we refer to it as the cuticle), which chamber over fresh silica gel. All eggs were occurs commonly throughout the Sphenisci- weighed at the same 10 time intervals as before. formes (Tyler 1965) and was postulated by Rahn Conductance to water vapor was calculated and Hammel (1982) to be a factor contributing using a modification of Fick’s first law of diffu- to the lower than expected shell conductance of sion: Adelie Penguin eggs. G ~20 = M,&P,,, (1) METHODS (Ar et al. 1974) Six eggsfrom different clutchesof P. adeliae were where G,,, is the conductance of the shell to collected at Cape Bird (70”13’S; 166”28’E) at the vapor (mg.day-1 .Tort-I), MHz0 is rate of move- very beginning of incubation (9-l 1 November ment of vapor acrossthe shell (mg.day-I), and 1987). They were kept cool (but not frozen) until AP,,, is the partial pressure gradient of vapor returned to Wellington in February 1988, when acrossthe shell (Torr). P,,, was assumed to be their length and breadth were measured to 0.1 zero outside the egg(over silica gel) and saturated mm with a dial vernier caliper (Table 1). inside the egg. Thus at 25.5”C P,,, is 24.5 Torr All eggswere equilibrated to 25.5”C overnight and at 25.6”c is 24.6 Torr (Weast and Astle 198 1). before being placed over silica gel in a bell cham- Eggs had lost mass prior to arrival in Wel- 306 MICHAEL B. THOMPSON AND KENNETH N. GOLDIE lington so fresh eggmass (W) was predicted from crographs. Other shell was also boiled in 10% length (L) and breadth (B) using the equation: NaOH (by weight) for 10 min to eliminate all organic components (especially shell mem- W = 0.548L.B2 (2) branes),rinsed in distilled water, and dried. Some (Hoyt 1979). of the NaOH treated shell was examined using This value was used to predict shell conductance the SEM, and some was set in resin for prepa- (G,,J using: ration of casts as described by Tompa (1980). Resin blocks were cut to reveal eggshell,which G HZO= 0.384 W0.814 (3) was then etched in 2M HCl, rinsed in distilled (Ar and Rahn 1978). water, dried over silica gel, and mounted for SEM Egg volume (V) was calculated using: analysis as described for the shell. Pore length and diameter at 10 equidistant points along the V = 0.507L.B2 (4) pore were measured from micrographs of pore (Hoyt 1979) casts. and this value was used to calculate surfacearea Mean values are given f one standard devia- (S) of each egg using: tion. Means are compared using Student’s t-test and statistical significanceis assumedifP < 0.05. S = [4.393 + 0.394.L,(B-‘)](V0.667) (5) (Hoyt 1976). RESULTS On completion of conductancemeasures, each The six eggsaveraged 68.6 f 2.7 mm long (range egg had 5-mm intervals marked with graphite =65.5-71.6mm)and55.8 f 1.7mmwide(range pencil around its long axis and was cut along this = 53.3-57.9 mm).
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