Second Half Of Incubation Research Articles

Effects of Thermal Manipulation During the Second Half of Incubation on Embryo Physiology, Hatching Parameters, and Quality of Broiler Chickens in Tropical Climate of Togo

Chickens are sensitive to environmental challenges caused by temperature. The current study aimed to determine the effects of heat manipulation during embryonic development on the physiological responses of Goliath chickens. A total of 2000 hatching eggs from 48-week-old breeders were weighed, numbered, and randomly distributed equally into 4 incubators. Each incubator received 500 eggs (4 replicates of 125 eggs each). Eggs in two of the incubators were rotated hourly at a 45° angle and maintained at 37.8°C and 60% relative humidity (T0 groups). Between embryonic days (ED) 10 and 18 of incubation, the eggs from the other two incubators were heated to 38.5°C for 6 hours per day (T1 groups). The eggs were reweighed and candled, and viable eggs were moved to the hatching baskets at ED 18 of incubation. Hatching eggs were examined individually for hatching events every three hours during the final three days of incubation. On day 21, blood samples were collected from 12 chicks per group for hormonal and biochemical analyses. The evaluated blood parameters included Triiodothyronine (T3), T4 (thyroxine), cortisol, uric acid, lactate dehydrogenase, and total protein. At hatch, chicks were weighed and their quality (survival after hatching and performance standards) was evaluated. Data were collected on embryonic development, hatching window, hatching events, biochemical parameters, and hormonal concentrations. Results indicated that hatchability, chick’s weight, Tri-iodothyronine, and corticosterone were higher in the T1 group, compared to the control group. At hatch on day 21, the pipping muscle of chicks in the treated group (T1) was significantly heavier than that of the control group, while the embryonic mortality rate was significantly higher in the T0 group. In conclusion, applying heat treatment for 6 hours at 38.5°C from ED10-ED18 of embryogenesis increased significantly the hatching rate, the pipping muscle, and the chick’s weight in this study. Keywords: Embryonic development, Physiology, Slow-growing broiler, Thermal manipulation, Tropical climate

Negligible evidence for detrimental effects of Leucocytozoon infections among Emperor Geese (Anser canagicus) breeding on the Yukon-Kuskokwim Delta, Alaska.

Emperor Geese (Anser canagicus) are iconic waterfowl endemic to Alaska and adjacent areas of northeastern Russia that are considered to be near threatened by the International Union for Conservation. This species has been identified as harboring diverse viruses and parasites which have, at times, been associated with disease in other avian taxa. To better assess if disease represents a vulnerability for Emperor Geese breeding on the Yukon-Kuskokwim Delta, Alaska, we evaluated if haemosporidian parasites were associated with decreased mass or survival among adult female nesting birds captured during 2006–2016. Through molecular analyses, we detected genetically diverse Leucocytozoon, Haemoproteus, and Plasmodium parasites in 28%, 1%, and 1% of 607 blood samples screened in triplicate, respectively. Using regression analysis, we found evidence for a small effect of Leucocytozoon infection on the mass of incubating adult female Emperor Geese. The estimated mass of infected individuals was approximately 43 g (95% CI: 20–67 g), or approximately 2%, less than uninfected birds when captured during the second half of incubation (days 11–25). We did not, however, find support for an effect of Leucocytozoon infection on survival of adult female nesting Emperor Geese using a multi-state hidden Markov framework to analyze mark-resight and recapture data. Using parasite mitochondrial DNA cytochrome b sequences, we identified 23 haplotypes among infected Emperor Geese. Leucocytozoon haplotypes clustered into three phylogenetically supported clades designated as ‘L. simondi clade A’, ‘L. simondi clade B’, and ‘other Leucocytozoon’. We did not find evidence that parasites assigned to any of these clades were associated with differential mass measures among nesting adult female Emperor Geese. Collectively, our results provide negligible evidence for Leucocytozoon parasites as causing detrimental effects to adult female Emperor Geese breeding on the Yukon-Kuskokwim Delta.

FEATURES OF PROCESSING OF WATERFOWL EGGS DURING THE INCUBATION

The purpose of this work was to conduct a comparative evaluation of existing disinfectants for their use in the second half of incubation for irrigation of waterfowl eggs. Methods. The study was conducted at the State Poultry Research Station of NAAS on hatching eggs of ducks and incubation wastes. Pre-incubation disinfection of eggs was carried out by irrigation with 5.0% solution of "Lasept-forte". Eggs were incubated according to standard regimes in modernized laboratory incubators ILU-F-0.3 and ILB-0.5. According to the requirements of the current standard for the technological process of incubation of duck eggs from the 13th day of incubation air cooling of eggs twice a day (morning and evening) for 10-15 minutes was carried out, and from the 20th day – a wet one. Modern disinfectants in various concentrations were used to irrigate duck eggs during incubation. Irrigation of duck eggs of the control group was performed with clean water. According to the results of research, the efficiency of irrigation of eggs with «Lasept-forte» and «Brovadez Plus» has been proved, and their optimal concentrations have been selected. It was found that the use of 0.5 % "Lasept-forte" or 0.05 % "Brovadez plus" for irrigation of duck eggs from the 20th day of incubation is desirable because it increases the results of incubation by 1.5- 2.5% due to the improvement of the sanitary condition in the incubator and the reduction of embryo death in the later stages of development. The pathological autopsy of incubation wastes showed that the category of waste "dead" (embryos that died after 25 days of incubation) was in the control group at 7.0%, in the experimental 6.0-6.5 %. At the same time, at the autopsy of the “dead” from these groups, there were signs of death from damage to eggs by microflora and fungi. In the control group, 64.3 % of deaths had such symptoms, in the first experimental group - 53.8 %, in the second experimental group - 33.3 %, which also proves the effectiveness of the means used.

No detectable effect of light-level geolocators on the behaviour and fitness of a long-distance migratory seabird

Light-level geolocators are increasingly popular devices for tracking migratory birds. However, to ensure that data on migratory behaviour represent natural behaviour that is not affected by potentially harmful effects of carrying such a device, their effects on behaviour and fitness should be assessed. A review of studies that tested for effects of tarsus-mounted light-level geolocators on seabirds showed that results are equivocal and often difficult to interpret due to the inclusion of only few traits and/or the lack of a proper experimental design. We therefore experimentally tested whether tarsus-mounted light-level geolocators affected a long-distance migratory seabird, the Common Tern Sterna hirundo. Using a well-matched treatment and control group, including both males and females, we tested whether light-level geolocators, deployed in the second half of incubation, affected the subsequent share of incubation, provisioning rate, reproductive performance, phenology or survival of tagged birds or their partners. In the year of deployment, we found no evidence for the behaviour of tagged birds or their partners to be affected by the geolocators. Moreover, we found no effect on their reproductive performance and departure date from the breeding colony. Finally, neither local survival to the next season, nor arrival date to the breeding colony in that season differed between tagged birds or their partners and control birds. These results suggest that a year of carrying a light-level geolocator attached to the tarsus has negligible effects on Common Terns and that such a device can be used to study their migratory behaviour without causing problems or introducing bias.

Parabronchial remodeling in chicks in response to embryonic hypoxia.

The embryonic development of parabronchi occurs mainly during the second half of incubation in precocious birds, which makes this phase sensitive to possible morphological modifications induced by O2 supply limitation. Thus, we hypothesized that hypoxia during the embryonic phase of parabronchial development induces morphological changes that remain after hatching. To test this hypothesis, chicken embryos were incubated entirely (21 days) under normoxia or partially under hypoxia (15% O2 during days 12 to 18). Lung structures, including air capillaries, blood capillaries, infundibula, atria, parabronchial lumen, bronchi, blood vessels larger than capillaries and interparabronchial tissue, in 1- and 10-day-old chicks were analyzed using light microscopy-assisted stereology. Tissue barrier and surface area of air capillaries were measured using electron microscopy-assisted stereology, allowing for calculation of the anatomical diffusion factor. Hypoxia increased the relative volumes of air and blood capillaries, structures directly involved in gas exchange, but decreased the relative volumes of atria in both groups of chicks, and the parabronchial lumen in older chicks. Accordingly, the surface area of the air capillaries and the anatomical diffusion factor were increased under hypoxic incubation. Treatment did not alter total lung volume, relative volumes of infundibula, bronchi, blood vessels larger than capillaries, interparabronchial tissue or the tissue barrier of any group. We conclude that hypoxia during the embryonic phase of parabronchial development leads to a morphological remodeling, characterized by increased volume density and respiratory surface area of structures involved in gas exchange at the expense of structures responsible for air conduction in chicks up to 10 days old.

Interaction between ascites susceptibility and CO2 during the second half of incubation of two broiler lines: the effect on post-hatch development and ascites mortality

1. The aim of this study was to investigate if genetic predisposition to ascites interacts with changed incubation conditions, and how this might affect the post-hatch performance and ascites susceptibility. 2. An ascites sensitive (A) and resistant (E) broiler line were incubated under standard or high CO2 conditions (up to 4%) from embryonic d 10 onwards. After hatch, chicks were exposed to cold from the 15th day of the rearing period to increase the incidence of ascites. 3. The A line had a higher post-hatch body weight from week three, higher blood pCO2 from d 21, higher haematocrit at d 35 and d 42, and higher plasma corticosterone concentration from d 21 onwards, compared with the E line, regardless of incubation conditions, supporting the given selection criteria. Ascites mortality did not, however, differ between lines. 4. Incubation under high CO2 conditions during the second half of incubation increased the ascites mortality, decreased body weight from week 4 onwards, affected venous blood pCO2, decreased blood pO2 from d 31, increased haematocrit at d 35 and d 42, and lowered the thyroxine and triiodothyronine concentrations at most sampling days. These effects were observed in both lines. The results suggested a metabolic programming of CO2 incubated chickens which affected ascites susceptibility.

High eggshell temperatures during incubation decrease growth performance and increase the incidence of ascites in broiler chickens

High eggshell temperatures (EST; ≥38.9°C) during the second half of incubation are known to decrease the body and organ development of broiler hatchlings. In particular, relative heart weights are decreased by a high EST, and this may increase the incidence of metabolic disorders that are associated with cardiovascular development, such as ascites. The current study investigated the effects of a high EST on chick quality, subsequent performance, and the incidence of ascites later in life. Eggs were incubated at a normal (37.8°C) or high (38.9°C) EST from d 7 of incubation onward. After hatching, the chickens were housed per EST in pens, and a normal or cold temperature schedule was applied during the grow-out period. Hatchability, hatchling quality, BW, feed conversion ratio, total mortality, mortality associated with ascites, slaughter characteristics, and ascites susceptibility at 6 wk of age were evaluated. Except for total ventricle weight, no interaction was found between EST and the grow-out temperature. Hatchability was comparable between the EST treatments, but the percentage of second-grade chickens was 0.7% higher at the high EST. Yolk-free body mass was 3.0 g lower, and heart weights were 26% lower at hatch in the high compared with the normal EST treatment. Body weight continued to be less during the grow-out period after the high EST incubation. However, breast meat yield was 1.0% higher in the high than in the normal EST. Feed conversion ratio did not differ between EST treatments. Total mortality was 4.1% higher and mortality associated with ascites was 3.8% higher in the high compared with the normal EST treatment. The ratio between the right and total ventricle was 1.1% higher in the high compared with the normal EST treatment at slaughter age. In conclusion, a high EST from d 7 of incubation onward decreased hatchling quality and growth performance, but increased breast meat yield. Furthermore, high EST incubation increased the incidence of ascites, which may be related to the reduced heart development at hatch.

Regulatory capacities of a broiler and layer strain exposed to high CO 2 levels during the second half of incubation

It has been shown that during embryonic chicken ( Gallus gallus) development, the metabolism of broiler embryos differs from that of layers in terms of embryonic growth, pCO 2/pO 2 blood levels, heat production, and heart rate. Therefore, these strains might adapt differently on extreme environmental factors such as exposure to high CO 2. The aim of this study was to compare broiler and layer embryos in their adaptation to 4% CO 2 from embryonic days (ED) 12 to 18. Due to hypercapnia, blood pCO 2 increased in both strains. Blood bicarbonate concentration was ~ 10 mmol/L higher in embryos exposed to high CO 2 of both strains, while the bicarbonates of broilers had ~ 5 mmol/L higher values than layer embryos. In addition, the pH increased when embryos of both strains were exposed to CO 2. Moreover, under CO 2 conditions, the blood potassium concentration increased in both strains significantly, reaching a plateau at ED14. At ED12, the layer strain had a higher increase in CAII protein in red blood cells due to incubation under high CO 2 compared to the broiler strain, whereas at ED14, the broiler strain had the highest increase. In conclusion, the most striking observation was the similar mechanism of broiler and layer embryos to cope with high CO 2 levels.

Interaction between ascites susceptibility and CO during the second half of incubation of two broiler lines. Effect on embryonic development and hatching process

1. Because CO2 during the second half of incubation is known to influence air cell and blood gases, and embryo development, it is postulated that post-hatch development and ascites sensitivity could also be influenced. 2. An ascites susceptible (A) and an ascites resistant (E) broiler line were incubated under standard incubation or high CO2 conditions (up to 4%) from embryonic day (ED) 10 onwards. The embryonic development and the hatching process of these two lines were compared when incubated under standard or high CO2 conditions from over the second half of incubation. 3. The A line, selected for high post-hatch growth rate, exhibited a higher relative embryo weight from ED10 until ED16, which was supported by a higher air cell pCO2, lower air cell pO2, higher corticosterone and thyroid hormones and earlier hatching time. 4. Incubation under high CO2 increased air cell pCO2, retarded yolk consumption, and decreased glycogen concentration in the liver at hatch. Hatchability decreased in both lines when incubated under high CO2, due to an increased late mortality of embryos that died before IP. 5. These results suggest that the development and metabolism of CO2-incubated embryos differ from control incubated embryos.

The importance of carbonic anhydrase II in red blood cells during exposure of chicken embryos to CO2

The importance of carbonic anhydrase (CA) during exposure of chicken embryos to CO(2) during the second half of incubation was investigated. The protein abundance and activity of CAII in erythrocytes was significantly higher in CO(2)-exposed embryos compared to normal conditions. Daily injections of acetazolamide (ATZ), an inhibitor of CA, increased blood P(CO2) and decreased blood pH in both control and CO(2)-incubated embryos. ATZ increased blood bicarbonate concentration in embryos exposed to normal atmosphere and in day-12 embryos exposed to high CO(2). The tendency of an increased blood potassium concentration in ATZ-injected embryos under standard atmospheric conditions might indicate that protons were exchanged with intracellular potassium. However, there was no evidence for such an exchange in CO(2)-incubated ATZ-treated embryos. This study shows for the first time that chicken embryos adapt to CO(2) during the second half of incubation by increasing CAII protein expression and function in red blood cells. This response may serve to "buffer" elevated CO(2) levels.

Involvement of thyroid hormones in chicken embryonic brain development

Thyroid hormones (THs) play an important role in vertebrate brain development by stimulating and coordinating cell proliferation, migration and differentiation. Several TH-responsive genes involved in these processes have been identified, but the information is mainly derived from studies of late brain development, while relatively little is known about the more early stages, prior to the onset of embryonic TH secretion. We have chosen the chick embryo to investigate the role of THs in both late and early brain development. T 4 and T 3 are found in chicken brain from the earliest stages tested (day 4). Indirect clues for the involvement of T 3 in brain development are found in the ontogenetic expression profiles of proteins regulating its bioavailability and action, including TH transporters, deiodinases and TH-receptors. All of them are expressed in whole embryos tested on day 2 of incubation and in developing brain tested from day 4 onwards. Their distribution patterns vary over time and according to the brain area and cell type studied. Hypothyroidism induced during the second half of incubation disturbs cell migration in the cerebellum, providing more direct evidence for the requirement for THs during the later stages of brain development. Direct morphological proof for the requirement for THs during the first half of incubation is still missing, but microarray analysis of telencephalon shows a clearly divergent gene expression profile in hypothyroid embryos. In vivo knockdown of TH transporters and deiodinases in chick embryos cultured ex ovo provides an excellent tool to study the role of THs in early brain development in more detail.

Breeding success and nest-site characteristics of Red-breasted Flycatchers Ficedula parva in a primeval forest

Capsule Predation is a major factor influencing the breeding success of Red-breasted Flycatchers under natural conditions. Aims To examine the breeding success of Red-breasted Flycatchers in relation to nest-site characteristics and time of breeding. Methods Data were collected during seven breeding seasons under natural conditions in Białowieża National Park. We used survival time analysis to estimate changes in survival probability over time and breeding success in relation to first-egg date and nest-site characteristics. Results Offspring fledged successfully from 51% of clutches, but the Kaplan–Meier estimate of survival was lower at 0.43. Of the unsuccessful clutches, 82% were lost to predators. The highest probability of loss was during egg-laying and in the second half of incubation. Breeding success was influenced by the height of the nest above the ground, but no relation to other nest-site characteristics was found. Conclusion To avoid predation Red-breasted Flycatchers build nests in various sites that are more open than most cavity-nesting species. The position of the nest-sites allows the sitting female to observe her surroundings and to escape rapidly from the nest, giving the chance for a replacement clutch.

Changes in Acid-Base Balance and Related Physiological Responses as a Result of External Hypercapnia During the Second Half of Incubation in the Chicken Embryo

This study investigated the effect of high CO2 (4%) from embryonic day (ED)10 until ED16 on the acid-base balance and related parameters in the chicken embryo. From ED10 to ED16, blood was taken from a vein from the chorioallantois membrane and was analyzed for pH, partial pressure of CO2, partial pressure of O2 (pO2), [HCO3(-)], [K+], and [Ca2+]. Allantoic fluid was taken for measurement of pH, NH3-N, phosphate, and calcium concentration. The right tibia was ashed, and calcium was measured with atomic absorption spectroscopy. Embryos exposed to high CO2 showed a consistent higher blood pH than control embryos. Notwithstanding this alkalosis, bicarbonate concentration was significantly higher in the CO2 group from ED12 until ED16. Potassium concentration in the blood was significantly higher in the CO2 group from ED11 until ED16. The pH of the allantois was significantly higher on ED14 and ED15. Ammonia N concentration was significantly higher in the CO2-incubated embryos on ED12 and ED13, whereas phosphate did not differ between groups. Calcium per tibia dry weight did not differ between incubation conditions. We can conclude that embryos adapt to high CO2 during the second half of incubation by increasing blood HCO3(-). It appears that this increase in HCO3(-)is mainly the result of the stimulated intracellular exchange of H+ with K+, although temporary reabsorption of HCO3(-)by the kidney cannot be excluded.

Effect of Four Percent Carbon Dioxide During the Second Half of Incubation on Embryonic Development, Hatching Parameters, and Posthatch Growth

In this study, broiler embryos were exposed during the second half of incubation [embryonic day (ED) 10 until ED18] to 4% CO(2). The CO(2) was set to reach 2% on ED11 and 4% from ED12 onward. Two experiments were conducted with the same setup. Embryo weight was measured and partial pressure of CO(2) and O(2) in the air cell was analyzed at several embryonic ages. Times of internal pipping, external pipping, and hatching were recorded. Chicks were raised until d 7 posthatch. Plasma corticosterone, triiodothyronine, and thyroxine concentrations were determined. Embryonic growth was not retarded and hatchability did not decrease in the CO(2)-incubated group, demonstrating that chicken embryos can tolerate high (4%) concentrations of CO(2) between ED10 and ED18. In the first experiment, partial pressure of CO(2) in the air cell was significantly higher in the CO(2) group on ED11, ED12, ED13, and ED14, but disappeared thereafter. This difference was not observed in the second experiment. A change in the hatching process of the CO(2) group was seen. Relative growths of newly hatched chicks until d 7 posthatch were equal in the CO(2) group and the control group. However, corticosterone and thyroxine concentrations were significantly higher in the CO(2)-incubated chicks on d 7 posthatch.

Micro‐environment of olive ridley turtle nests deposited during an aggregated nesting event

AbstractThe hatching success of nests deposited by olive ridley turtles Lepidochelys olivacea during aggregated nesting events (‘arribada’) is typically low and the underlying mechanisms are not clear. In this study, temperature, oxygen and carbon dioxide partial pressures (PO2 and PCO2) of in situ nests as well as nests relocated into a hatchery with clean sand were monitored throughout incubation. Hatching success of hatchery nests was significantly higher than in situ nests (83.1 vs. 21.6%) and mainly resulted from higher mortality of early‐stage embryos. During the first half of incubation, temperature and PCO2 were higher (by 0.6°C and 0.7 kPa, respectively) and PO2 was lower (by 1.1 kPa) within in situ relative to hatchery nests. Because embryo metabolism does not interfere significantly with nest gas contents during the first half of incubation, these results suggest that the greater content of organic matter and/or microorganisms in the sand surrounding in situ nests had an effect on nest gas contents. As PO2 and PCO2 differences were relatively small, microbial activity (such as fungal and bacterial infection) may have caused the early embryo mortality found in situ. Moreover, our results suggest that during the second half of incubation, neither PO2 nor PCO2 reached threshold levels that resulted in the death of embryos or hatchlings. Overall, this study showed a clear benefit of using clean sand to increase hatchling production in arribada beaches and highlights the importance of further investigating the relationship between nest micro‐environment, sand microbial activity and embryo development under natural conditions during these unique nesting events.

Effects of hatchery shading and nest depth on the development and quality of Chelonia mydas hatchlings: implications for hatchery management in Peninsular, Malaysia

One of the decisions made by hatchery managers around the world is what degree of shading and nest depth are required to maximise the production of high-quality hatchlings at optimal sex ratios. The primary objectives of this study were to determine the effects of (1) hatchery shading and nest depth on nest temperatures and emergence lag, and (2) nest temperatures and nest depth on hatchling sex ratio and quality. In 2001, 26 Chelonia mydas clutches from Ma’Daerah beach, Terengganu, Malaysia, were relocated alternatively at depths of 50 cm and 75 cm into a 70%-shaded and a 100%-shaded hatchery. Data loggers were placed into the centre of each relocated clutch to record the temperature every hour over the course of incubation. When the hatchlings emerged, a sample of the clutch was run, measured and weighed and a separate sample was examined histologically for sex characteristics. Nest temperatures ranged between 28°C and 30°C and generally showed increases over the second half of incubation due to metabolic heating of the clutch. There was no significant correlation found between nest temperature and any of the hatchling parameters measured. Hatchlings from 75-cm-deep nests had a longer emergence lag (46.4 (±10.2) h) than hatchlings from 50-cm-deep nests. Hatch and emergence success were similar to those of natural populations and hatchling sex ratios were male dominant, with an average of 72% males. There was a poor correlation between mean middle-third incubation temperatures and sex ratio. Hatchlings from 75-cm-deep nests had similar running speeds but lower condition index than their conspecifics from 50-cm-deep nests.

Developmental expression of glycocomponents in the chick chorioallantoic membrane.

Widespread interest has focused on the research of the chorioallantoic membrane (CAM) and its functional contribution to gaseous exchange, calcium reabsorption, water and electrolyte transport during chick embryogenesis. Nevertheless, very little information is available on the glycoconjugate components of this extra-embryonic structure. In the present study, we investigated by lectin histochemistry, the glycosylation pattern expressed in the CAM epithelia during embryonic development. Occurrence of sialic acid-associated glycoproteins was detailed by either specific lectins, which discriminate alpha2,3 and alpha2,6 sialoderivatives, or sialidase digestion combined with appropriate lectins to identify the sialic acid acceptor sugars. Lectin affinities proved to depend greatly on differentiation of the CAM epithelia which showed highest expression of binding sites during the second half of incubation up to hatching. Differences emerged between the chorionic and the allantoic epithelium, regarding qualitative, quantitative and temporal expression of sugar moieties. A cell type-specific distribution of glycocomponents was found in the chorionic epithelium where lectin binding sites were specifically located in the villus cavity cells. In the allantoic epithelium, high and heterogeneous occurrence of sialoglycoconjugates as well as specific presence of fucose residues were evidenced mostly in the granule cells. We conclude from these findings that various glycoconjugates in the CAM could participate in different physiological functions characteristic of the chorionic and the allantoic epithelium.

Gas exchange and energy metabolism of the ostrich ( Struthio camelus) embryo

We measured oxygen consumption ( V ̇ O 2 ) and carbon dioxide emission ( V ̇ CO 2 ) rates, air-cell gas partial pressures of oxygen (P AO 2) and CO 2 (P ACO 2), eggshell water vapour conductance and energy content of the ostrich ( Struthio camelus) egg, ‘true hatchling’ and residual yolk, and calculated RQ and total oxygen consumption ( V ̇ O 2 tot ) for ostrich eggs incubated at 36.5°C and 25% relative humidity. The V ̇ O 2 pattern showed a drop of approximately 5% before internal pipping. V ̇ O 2 just prior to internal pipping agrees with allometric calculations. Despite the higher incubation temperature compared to other studies, and the resultant shorter incubation duration (42 days), V ̇ O 2 tot (91.7 l kg −1) was similar to a previously reported value. RQ values during the second half of incubation (approx. 0.68) were lower than expected for lipid catabolism. Prior to internal pipping, P AO 2 and P ACO 2 were 98 and 48.3 torr (13.1 and 6.4 kPa), respectively. The growth pattern of the ostrich embryo is different from the typical precocial pattern, showing a time delay in the rapid growth phase. As a result, the lowered overall energy expenditure for tissue maintenance, as compared to other species, is reflected in the low yolk utilization and high residual yolk fraction of the whole hatchling dry mass. These could also result from the relatively short incubation period of the ostrich egg, thereby evading desiccation by excess water loss.

Developmental patterns of heart rate in altricial avian embryos and hatchlings

The aims of this study were to determine the patterns of development of heart rate (fH) in altricial avian embryos and hatchlings, and then to examine how fH is regulated to meet metabolic requirements in altricial embryos. Embryonic mean heart rate (fH-) in 12 altricial species (Passeriformes and Psittaciformes) increased during pre-pipping incubation in all species except the cockatiel (Nymphicus hollandicus), in which fH- tended to decrease prior to pipping. The rate of increase in fH- tripled during the pipping phase in all species, and fH- was significantly higher during the pipping period and in hatchlings than during pre-pipping development. The O2 pulse (O2 consumed per cardiac beat) of altricial embryos increased in direct proportion to embryo mass (loge/loge base), although fH- was often low prior to pipping, implying that stroke volume increases in the second half of incubation. We conclude that fH contributes more than other factors towards supplying the metabolic demands of the embryo during the middle of incubation and the final pipping phase, but less during the intervening period of late incubation.

Development of heart rate in the precocial king quail Coturnix chinensis.

Our aim was to examine changes in heart rate (fh) during the embryonic and posthatching periods of the smallest precocial avian species, Coturnix chinensis. In experiment I, repeated measurements of mean fh were made for individual quail by ballistocardiogram (BCG) during incubation, and by both piezo-electric film and electrocardiogram (ECG) during the posthatching period (resting and thermoneutral conditions). Mean fh of all embryos increased during the second half of incubation and the first week posthatching, but a few embryos experienced a very brief period of decreased fh prior to internal pipping. After the first week, fh of posthatching quail was maintained at high levels (550-650 beats min-1), then decreased with age and increase in body mass. The maximal fh of quail chicks represents a greater posthatching increase in fh than is found in larger precocial chickens, this difference being attributable to the higher demands of thermoregulation at small body masses in the quail. In experiment II, the mean fh of quail embryos (day 2-16) was recorded by ECG, and embryonic stage, yolk-free embryo mass (wet and dry) and water content were measured. Mean fh was linearly related to embryo mass throughout incubation, except on the day prior to internal pipping, when the fh of a few embryos declined below this linear relationship. Measurements of instantaneous fh of late incubation embryos, young and adult quail all showed spontaneous fluctuations in fh. Two main frequency components of fh fluctuations were identified for the first time in an avian species. Low-frequency (mean 0.09 Hz, 12.6 s) and high-frequency (1.4 Hz, 0.9 s) oscillations in both young chicks and adult quail were detected and are considered to reflect baroreflex mediation of fh and respiratory sinus arrhythmia, respectively.