J. T. Pearson

Development of heart rate irregularities in chick embryos

Heart rate (HR) irregularities in chick embryos were defined as large fluctuations (>10 beats/min) comprising irregular, brief deceleration and/or acceleration of instantaneous HR (IHR). IHR was determined directly from the arterial blood pressure while adequate gas exchange was maintained through an eggshell and chorioallantoic membrane. Five embryos were examined on each day from day 11 to day 19 of incubation. Baseline HR was stable until day 12-13, and on around day 13-14 transient, rapid deceleration of HR (termed V pattern) began to appear, with a subsequent increase in its frequency and magnitude. The acceleration patterns (lambda, avian omega, and periodic patterns) appeared later, and the IHR became increasingly irregular, with additional, spontaneous deceleration and acceleration patterns toward hatching. Additional experiments with intravenous administration of autonomic drugs clearly showed that rapid deceleration of HR was mediated by parasympathetic nervous function but did not always show clear relations of sympathomimetic and sympathetic blocking agents to the acceleration patterns.Heart rate (HR) irregularities in chick embryos were defined as large fluctuations (>10 beats/min) comprising irregular, brief deceleration and/or acceleration of instantaneous HR (IHR). IHR was determined directly from the arterial blood pressure while adequate gas exchange was maintained through an eggshell and chorioallantoic membrane. Five embryos were examined on each day from day 11 to day 19 of incubation. Baseline HR was stable until day 12-13, and on around day 13-14 transient, rapid deceleration of HR (termed V pattern) began to appear, with a subsequent increase in its frequency and magnitude. The acceleration patterns (lambda, avian omega, and periodic patterns) appeared later, and the IHR became increasingly irregular, with additional, spontaneous deceleration and acceleration patterns toward hatching. Additional experiments with intravenous administration of autonomic drugs clearly showed that rapid deceleration of HR was mediated by parasympathetic nervous function but did not always show clear relations of sympathomimetic and sympathetic blocking agents to the acceleration patterns.

Long-Term Measurement of Heart Rate in Chicken Eggs

Taking advantage of acoustocardiogram (ACG), we measured the heart rate (HR) of chick embryos continuously from day 12 until hatching and then investigated the development of HR irregularities (HRI), HR variability (HRV), and the existence of a circadian rhythm in mean HR (MHR). HRI comprised transient bradycardia and tachycardia, which first developed on day 14 and 16 in most embryos, respectively. Transient bradycardia increased in frequency and magnitude with embryonic development and occurred over periods of up to 30 min in some embryos. MHR was maximal on around days 14-15 and thereafter decreased to about 250-260 bpm on days 16-18. Baseline HRV, which is an oscillation of the MHR baseline, occurred as HR decreased from days 15-16 and became predominant on days 17-18. The magnitude of the baseline oscillations reached up to 50 bpm in some embryos and the period ranged between about 40-90 min (ultradian rhythm). A circadian rhythm of MHR was not found in late chick embryos. On days 18-19, embryonic activities were augmented and then breathing movements began to occur, disturbing ACG signals and thus making it difficult to measure the HR. Instead, the development of breathing activities was recorded. Breathing frequency was irregular at first and then increased to a maximum of about 1.5 Hz prior to hatching.

Non-invasive determination of instantaneous heart rate in developing avian embryos by means of acoustocardiogram

Previous noninvasive studies of the mean heart rate of embryonic birds have prompted an investigation into the instantaneous heart rate (IHR), which may be informative in developmental studies of cardiac rhythm. Using the acoustocardiogram (ACG), a noninvasive, long-term measuring system for embryonic IHR is developed, and the IHR in chickens during the last half of embryonic development is determined. The system, which uses a micro-computer, samples the ACG at a frequency of 50 Hz, restores the ACG wave by sinc function and calculates the IHR with an error in accuracy of less than 1 beat min−1. It was found that characteristic, transient bradycardia begins to appear late in the second week of incubation, and, with the additional development of transient tachycardia, the embryonic cardiac rhythm becomes more arrhythmic towards hatching. Simultaneous measurements of IHR with somatic movements showed no relationship between arrhythmia and embryonic activities. This system is useful, providing new evidence on long-term IHR developmental patterns.

Cardiac rhythms in chick embryos during hatching

Avian embryos develop within a hard eggshell which permits the measurement of heart rate while maintaining an adequate gas exchange through the chorioallantoic membrane. Heart rate has been determined from cardiogenic signals detected either noninvasively, semi-invasively or invasively with various transducers. Firstly, we reviewed these previously-developed methods and experimental results on heart rate fluctuations in prenatal embryos. Secondly, we presented new findings on the development of heart rate fluctuations during the last stages of incubation, with emphasis on the perinatal period, which remained to be studied. Three patterns of acceleration of the instantaneous heart rate were unique to the external pipping period: irregular intermittent large accelerations, short-term repeated large accelerations and relatively long-lasting cyclic small accelerations. Besides these acceleration patterns, respiratory arrhythmia, which comprimised oscillating patterns with a period of 1-1.5 s, appeared during the external pipping period. Furthermore, additional oscillating patterns with a period of 10-15 min were found in some externally pipped embryos.

Cardiac rhythms in developing chicks

Instantaneous heart rate (IHR) of chicks was determined by electrocardiogram measured non-invasively from the day of hatch to day 6 for continuity of investigation of HR fluctuations from embryos and for ascertainment of HR diurnal rhythms. In Experiment I, IHR was determined for 1-h periods twice a day, in daytime and at night, to investigate development of heart rate fluctuations (variability and irregularities). Chick IHR was substantially more arrhythmic than embryonic HR and spontaneous acceleration dominated HR fluctuations. Chick HR fluctuations were categorized into three types; [1] Type I as a widespread baseline HR (20-50 bpm) due to respiratory arrhythmia, with a mean oscillatory frequency of 0.74 Hz (range 0.4-1.2 Hz); [2] Type II as low frequency oscillations of baseline HR, at a mean of 0.07 Hz (range 0.04-0.10 Hz), and [3] Type III as non-cyclic irregularities, dominated by frequent transient accelerations. In Experiment II, continuous measurements of HR were made under conditions of a natural photoperiod, thermoneutrality and with feed available throughout the first week after hatching and circadian rhythms of HR were ascertained. HR was very variable in the daytime (250-500 bpm), due in part to feeding and activity, and decreased to a diurnal low (200-350 bpm) at night when mean HR was relatively stable. HR fluctuations persisted throughout the diurnal cycle.

Developmental patterns of O2 consumption, heart rate and O2 pulse in unturned eggs.

The effects of failure to turn eggs on the developmental patterns of oxygen consumption (MO2), heart rate (fH) and O2 pulse during the second half of incubation of individual chicken eggs were examined. The MO2 of unturned eggs increased at a significantly lower rate than the control toward the end of prenatal incubation, and the plateau MO2 between day 17 and 19 was significantly lower than the control. Lack of turning also resulted in significant changes in the developmental patterns of fH and O2 pulse. It is suggested that the effects of lack of egg-turning on the developmental patterns of MO2 may be attributable to lower embryonic growth rate in addition to impairment of gas exchange through the chorioallantoic gas exchanger.

Heart rate in developing ostrich embryos

1. A non-invasive condenser microphone was used to detect cardiogenic, acoustic pressure changes (acoustocardiogram, ACG) over the eggshell in order to determine embryonic heart rate (HR) of ostriches in a commercial hatchery. 2. HR measured for 36 eggs at 36.3C was maintained at about 185 bpm during the middle stage of development (days 19 to 23) and then decreased with embryonic development. 3. The daily changes in HR were not related to egg mass, but HR of high water vapour conductance (GspH2O) eggs was found to decrease less during the last stages of incubation relative to low and medium GspH2O groups. 4. The averaged HR at 80% of incubation period was close to the value predicted from the allometric equation determined previously for embryos of domesticated precocial birds.

Developmental allometry of pulmonary structure and function in the altricial Australian pelican Pelecanus conspicillatus

SUMMARY Quantitative methods have been used to correlate maximal oxygen uptake with lung development in Australian pelicans. These birds produce the largest altricial neonates and become some of the largest birds capable of flight. During post-hatching growth to adults, body mass increases by two orders of magnitude (from 88 g to 8.8 kg). Oxygen consumption rates were measured at rest and during exposure to cold and during exercise. Then the lungs were quantitatively assessed using morphometric techniques. Allometric relationships between body mass (M) and gas exchange parameters (Y) were determined and evaluated by examining the exponents of the equation Y=aMb. This intraspecific study was compared to interspecific studies of adult birds reported in the literature. Total lung volume scales similarly in juvenile pelicans (b=1.05) as in adult birds (b=1.02). However, surface area of the blood–gas barrier greatly increases (b=1.25), and its harmonic mean thickness does not significantly change (b=0.02), in comparison to exponents from adult birds (b=0.86 and 0.07, respectively). As a result, the diffusing capacity of the blood–gas tissue barrier increases much more during development (b=1.23) than it does in adult birds of different sizes (b=0.79). It increases in parallel to maximal oxygen consumption rate (b=1.28), suggesting that the gas exchange system is either limited by lung development or possibly symmorphic. The capacity of the oxygen delivery system is theoretically sufficient for powered flight well in advance of the birds need to use it.

Development and regulation of heart rate in embryos and hatchlings of gulls (Larus schistisagus and Larus crassirostris) in relation to growth

Abstract We compared the developmental patterns of mean heart rate in Larus crassirostris and L. schistisagus embryos and chicks with other avian species of different hatchling developmental modes. We proposed that, since mean heart rate is inversely related to fresh egg mass in all birds, larger species reached a higher fraction of their hatchling mean heart rate by the end of the early phase of incubation and that heart rate contributions to supplying the increasing metabolic demands during mid and late incubation phases were less important than in smaller avian species. Mean heart rate was essentially independent of age throughout the mid-incubation phase (33% of normalised incubation until pipping), but increased with time during early (L. schistisagus only investigated) and late-incubation phases in both species. The O2 pulse of L. schistisagus embryos and chicks increased linearly with yolk-free body mass (log-log) after the early-phase of incubation until shortly before pipping, but was independent of body mass in the periods before and after. We conclude that a high heart rate in this first period is probably more important for increasing the circulation of nutrients to the embryo at a stage when extra-embryonic circulation to the yolk sac is limited by the size of the growing area vaculosa. Furthermore, large increases in mean heart rate during the late-incubation phase are probably important for increasing the cardiac output to hatching levels with onset of endothermy. However, mean heart rate is stable over the mid-incubation while O2 pulse increases, suggesting that increases in stroke volume and other circulatory adjustments may be entirely responsible for the largest increases in O2 transport during incubation of large avian species.

Development of cardiac rhythms in altricial avian embryos.

Mean heart rate (MHR) was determined during incubation and in hatchlings of 14 altricial avian species to investigate (1) if there is a common developmental pattern of heart rate in altricial embryos and (2) if heart rate changes during incubation are correlated with changes in embryonic growth rate. On the basis of normalized incubation MHR increased approximately linearly in 12 of 14 species from as early as 30-40% of incubation to that of pipped embryos. The MHR of hatchlings was equal to or higher than that of pipped embryos in seven species. Passerine embryos and hatchlings maintained higher MHR in comparison to parrots of similar egg mass, which may reflect phylogenetic differences in development. Embryonic MHR increased at a higher rate while embryonic growth rates were highest during the first 40% of incubation in tit, budgerigar and crow embryos than during subsequent development when relative growth rates decreased. MHR became independent of yolk-free wet mass at a smaller fraction of hatchling mass in budgerigar and crow than in the tit, suggesting that MHR is more likely to increase continuously after 40% of incubation in small altricial species than larger species.