Embryonic heart rate and oxygen pulse in two procellariiform seabirds,Diomedea immutabilis and Puffinus pacificus

Developmental patterns of embryonic heart rate were measured non-invasively in two procellariiform seabirds, the Laysan albatross (Diomedea immutabilis) and wedge-tailed shearwater (Puffinus pacificus), during prepipping and after pipping. The O₂ pulse, defined as the O₂ consumption per single heart beat, was calculated using the previously reported O₂ consumption for these species. The embryonic heart rate of the albatross was not changed by internal pipping (initial pipping event in this species), remained unchanged during the prolonged internal pipping period and tended to increase, although insignificantly, with the initiation of external pipping (second pipping event). Heart rate in the shearwater remained unchanged during the late prepipping stages, did not change with external pipping (initial pipping event), but increased during the prolonged internal pipping period (second pipping event) and reached a maximum on the last day of incubation. The developmental pattern of heart rate in the shearwater was very similar to that reported previously for the brown noddy, a member of the order Charadriiformes but with the same pipping sequence. Developmental patterns of embryonic O₂ pulse were also different between the two procellariiform seabirds. However, those of the shearwater and the noddy were similar. The sequence of access to atmospheric O₂ during development and hatching may be an important factor determining the developmental patterns of embryonic heart rate and O₂ pulse.Abbreviations A-VO₂ difference, arteriovenous oxygen difference - BCG ballistocardiogram of egg - CV coefficient of variation - EP external pipping - HR heart rate - IP internal pipping - mass mass of freshly laid egg - SD standard deviation