There has been considerable recent interest in the development of the circulation in the zebrafish. Optical techniques typically used to visualize changes in heart size allow measurement of stroke volume during early vertebrate development, but this approach is complicated in zebrafish larvae because of the heart's irregular shape and its significant change in morphology during the first 6 d of development. By use of a threedimensional integration of the early zebrafish heart and vessels, we have greatly reduced measurement error of stroke volume and cardiac output and have determined the cross-sectional growth of major vessels in the developing zebrafish larvae. A dramatic 500%-600% increase in cardiac output (from 10 to 50-60 nL min Ϫ1 ) occurs on days 5 and 6 postfertilization in Danio rerio. Cross-sectional area of key vessels (dorsal artery, caudal artery, dorsal vein) as well as between-individual variation significantly decreased over the first 6 d of development. Associated with the decrease in cross-sectional area is a significant increase in red blood cell velocity on days 5 and 6 postfertilization. Together, the three-dimensional data of the cardiac and vascular systems have shown that the most profound physiological and developmental changes occur in days 5 and 6, which corresponds with the appearance of the adult form of the heart and the transition from diffusive to convective O 2 supply to internal tissues.
The developmental environment influences a wide variety of phenotypic traits in the adults of many vertebrates (i.e., developmental plasticity). In this study, we test to see if developmental environment (E DEV) interacts with the adult behavioral environment (E BEHAV) in determining behavioral phenotypes. We reared Zebrafish (Danio rerio) from eggs in either continuously hypoxic or normoxic conditions. We then tested aggression and avoidance (i.e., hiding) levels of fish from each developmental treatment in both environments. Developmental environment was a significant source of variation in avoidance behavior while the stimulus environment did not influence avoidance. Without a period of acclimation we found that E BEHAV and an E DEV X E BEHAV interaction were both significant sources of variation. However, when the fish were allowed to physiologically acclimate to the environment for 16 h, aggression level was highest for fish tested in the environment in which they developed. In that case the E DEV X E BEHAV interaction was the only significant source of variation. These results demonstrate that a more complete understanding of phenotypic response can be gained by incorporating environmental conditions across multiple time scales.
Relative to normoxia-reared fish, hypoxia-reared zebrafish Danio rerio achieved 24Á9 and 21Á4% slower maximal swim velocities in both normoxic and hypoxic waters, respectively. Hypoxiareared fish also produced 26Á1 and 63Á9% less lactate during resting conditions across both normoxic and hypoxic waters, respectively. During exercise, this trend continued as hypoxiareared fish produced 68Á2 and 55Á1% less lactate across both normoxic and hypoxic waters, respectively. This reduction in performance, rather than representing a purely pathological (maladaptive) response to hypoxia, appears to represent a fundamental shift in the metabolic response to hypoxia.
SUMMARY
Our understanding of avian embryonic cardiovascular regulation has been based on studies in chickens. The present study was undertaken to determine if the patterns established in chickens are generally applicable to the emu, a ratite bird species. We studied cardiovascular physiology over the interval from 60% to 90% of the emu's 50-day incubation period. During this period,embryonic emus exhibit a slight fall in resting heart rate (from 171 beats min-1 to 154 beats min-1) and a doubling of mean arterial pressure (from 1.2 kPa to 2.6 kPa). Exposures to 15% or 10%O2 initially decreased heart rate during the first period of emu incubation studied [60% of incubation (60%I)] but increased heart rate in the 90%I group. Arterial pressure responded to hypoxia with an initial depression(-1.6 kPa) at 60%I and 70%I but showed no response during the later periods of incubation (80%I and 90%I). In addition, tonic stimulation of both cholinergic and adrenergic (α and β) receptors was present on heart rate at 70%I, with the cholinergic and β-adrenergic tone increasing in strength by 90%I. Arterial pressure was dependent on a constant β-adrenergic and constant α-adrenergic tone from 60%I to 90%I. A comparison with embryonic white leghorn chickens over a similar window of incubation revealed that emus and white leghorn chickens both possess an adrenergic tone on heart rate and pressure but that only emus possess a cholinergic tone on heart rate. Collectively, these data indicate that the maturation of cardiovascular control systems differs between white leghorn chickens and emus, inviting investigation of additional avian species to determine other patterns.
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