Daily temporal organization of laying in Japanese quail: variability and heritability

In birds, many behavioral and physiological processes that occur during reproduction show daily rhythms in response to environmental temporal constraints. In this study, the individual daily organization of laying and its genetic determinant in Japanese quail (Coturnix coturnix japonica) were analyzed. For this purpose, the oviposition time of 102 randomly chosen females, maintained in long-day photoperiodic conditions (LD 14h:10h) for 1 mon was observed and extreme phenotypes selected. Laying is characterized by two parameters: oviposition interval and laying hour.

The birds showed a specific time of laying during 24 h. All eggs were laid in the afternoon between 6.5 and 14 h after lights on (HALO). Two laying profiles were determined: 20% of females with an oviposition interval greater than 24 h (24.7±0.2 h) (the “delayed” profile) laid progressively later each day until a pause day. The remaining 80% of the females laid at the same time each day, with few pause days and an oviposition interval close to 24 h (24.0±0.2 h) (the “stable” profile). Among the females, showing this last profile, an intra-individual stability and an inter-individual variability of laying hour was established. Two extreme laying phenotypes were then determined: the “early” phenotype (E) for females laying on average between 7.5 and 9.5 HALO and the “late” phenotype (L) for females laying between 12.5 and 14 HALO.

In order to study the genetic basis of the laying hour, three females of each extreme phenotype were selected and crossed with two different males. The E and L females produced 57 F1E and 42 F1L daughters, respectively. F1 females displayed both laying profiles. However, the proportion of females displaying a “delayed” profile was higher in the L line (50%) than in the E line (29.8%). For the “stable” daughters, artificial selection induced an advance in laying hour of 4.7% for the E line and a delay of 4.7% for the L line. Realized heritability was estimated at 0.5. Moreover, the laying hour of the daughters was correlated positively to that of the mothers (N=61;r=0.45). These results support the notion of heritability of oviposition time in Japanese quail.     

Effects of mating system in Japanese quail

Summary A 17-generation selection experiment was conducted to study direct and correlated responses to mass selection under a mating system with alternating generations of full-sib inbreeding and wide outbreeding (population I) as compared with a mass selected, randomly mated population (population II). The selection criterion was an index of total egg mass to 78 days divided by adult female body weight. Estimates of heritabilities and genetic correlations are reported. Estimated heritabilities for the index were 0.38±0.04 and 0.29±0.05 in population I and II, respectively. Realized heritabilites were 0.10±0.05 and 0.12±0.03. For most traits studied the mean phenotypic values in the cyclic mated population decreased for inbred generations. Increased inbreeding levels also caused outbred generation means of population I to decrease through the first six or seven generations. After this period of adaptation to inbreeding selection response was positive for the index and positively correlated traits. Total response to selection under the cyclic inbred-outbred mating system did not exceed selection response made under random mating. However, the rate of response in the cyclically mated population exceeded that in the randomly mated population in later generations when the cyclically mated population had apparently adapted to inbreeding.Published with approval of the Director of the Montana Agricultural Experiment Station, Journal Series No. 1221     

Stable individual profiles of daily timing of migratory restlessness in European quail

Temporal characteristics of migratory behavior in birds are usually studied at the species and population levels, and rarely at the individual level. Variations among species and populations of the seasonal onset of migratory behavior have been widely investigated, but very little is known about its daily organization or whether birds are conservative in their behavior. The determination of intra- and inter-individual variability is important for the study of genetic variations and can reveal the existence of different adaptation capacities within populations. This laboratory study analyzed intra- and inter-individual variability of daily initiation and time course of nocturnal restlessness in partial-migrant European quail (Coturnix coturnix coturnix). Thirty-five quail were selected randomly from a captive stock, and their spring activity was recorded under natural daylenghs. Eighteen of the thirty-five quail presented behavioral profiles of migrant birds. Migrant birds initiated their nocturnal activity punctually, and the time courses of the nocturnal activity of 88% of them revealed intra-individual stability over six consecutive nights. All birds initiated their nocturnal activity after sunset and civil twilight, and they were more active at the beginning than the middle or end of the night, suggesting that their drive to migrate could be synchronized with particular skylight conditions. For the first time, stable individual profiles in the daily time course of migratory restlessness are shown. These results support previous findings concerning biological rhythms of quail and raise questions concerning the timing of migratory behavior.     

Stable Individual Profiles of Daily Timing of Migratory Restlessness in European Quail

Temporal characteristics of migratory behavior in birds are usually studied at the species and population levels, and rarely at the individual level. Variations among species and populations of the seasonal onset of migratory behavior have been widely investigated, but very little is known about its daily organization or whether birds are conservative in their behavior. The determination of intra‐ and inter‐individual variability is important for the study of genetic variations and can reveal the existence of different adaptation capacities within populations. This laboratory study analyzed intra‐ and inter‐individual variability of daily initiation and time course of nocturnal restlessness in partial‐migrant European quail (Coturnix coturnix coturnix). Thirty‐five quail were selected randomly from a captive stock, and their spring activity was recorded under natural daylenghs. Eighteen of the thirty‐five quail presented behavioral profiles of migrant birds. Migrant birds initiated their nocturnal activity punctually, and the time courses of the nocturnal activity of 88% of them revealed intra‐individual stability over six consecutive nights. All birds initiated their nocturnal activity after sunset and civil twilight, and they were more active at the beginning than the middle or end of the night, suggesting that their drive to migrate could be synchronized with particular skylight conditions. For the first time, stable individual profiles in the daily time course of migratory restlessness are shown. These results support previous findings concerning biological rhythms of quail and raise questions concerning the timing of migratory behavior.     

Daily organization of laying in Japanese and European quail: effect of domestication

In the present study, we aimed to examine the possible effects of domestication on the daily organization of laying in female quail. To achieve this we compared laying patterns in a laboratory strain of Japanese quail with those in European quail originating from a wild population. Under LD 14:10, the same specific laying window was observed, with ovipositions occurring mainly between 7 h and 15 h after lights on. Thus, domestication did not modify the specific daily organization of laying in females laying in the afternoon. Research about the functional value of this temporal organization in the field thus seems justified. At an individual level, two distinct laying profiles were recorded, as in a previous experiment on an industrial Japanese quail strain selected for meat and egg production. However, different frequencies were noted. Whereas stable profile females (laying at the same time each day) were predominant (80.4%, N=102) in the industrial Japanese quail strain, delayed profile females (laying successively later each day) appeared dominant in the European quail (81%, N=22). The Japanese quail from the laboratory strain constituted an intermediate group with a slight dominance of the stable profile (58.8%, N=17). Thus, domestication has changed the frequencies of the laying profiles, favoring stable females, which are the most productive birds. A model involving interaction between one circadian rhythm and one ultradian rhythm can explain the observed laying profiles.     

Effect of a cranium-directed daily illumination cycle on the laying rhythm in Japanese quail

This study tests the effect on the laying rhythm of a light cycle reaching directly the encephala via a diode in Japanese quail maintained in constant darkness. In DD, all the birds expressed their free-running laying rhythm (period close to 25 h). When the diode is switched on 14 h per 24 h cycle, females showed the same organization as in LD with the same laying time. Thus, a photoperiodic cycle, where light was only perceived through the skull of the female quail, could synchronize its laying rhythm. This device with a LED is an interesting alternative solution to eye-patching or blinding of birds.     

Ontogeny of the ultradian rhythm of activity in Japanese quail

As soon as they hatch, gallinaceous chicks follow broody hens. This matriarchal unit presents a temporal organization of activity. The ontogeny of this ultradian rhythm of activity was followed in Japanese quail during their first 3 weeks of life. Under controlled laboratory conditions, 12 groups of four chicks were recorded using an activity monitoring system. They were observed between the ages of 2 and 17 days. Chicks in groups presented an ultradian rhythm of activity, with a period that increased significantly from 14.3 ± 1.4 minutes when chicks were 2 days old to 26.0 ± 1.9 minutes when they were 16 days old. The increase of ultradian periodicity was particularly pronounced during their first and third weeks of life. Finally, the ultradian period was correlated positively with body weight of the chicks. (Chronobiology International, 17(6), 767-776, 2000)     

Development of the skeleton in Japanese quail embryos

In the research fields of experimental embryology, teratological testing, and developmental engineering in avian species, a knowledge of normal embryonic development is necessary so that research may be performed efficiently and precisely. A series of normal stages based on external appearance has been established in both chicken and quail embryos. Those based on skeletal features, however, have not been elucidated. The present study newly established a series of normal stages for the development of the Japanese quail embryo skeleton. This series is composed of 15 stages determined by observing the timing of chondrification and calcification of the skeleton every 24 h, from 3 to 17 days of incubation. Cartilage and ossified bones were stained blue and red with Alcian blue 8GX and alizarin red S, respectively. These skeletogenous stages of the Japanese quail embryo will be useful as a normal control not only in studies of experimental embryology, teratological testing, and developmental engineering, but also in the analysis of mutant embryos with skeletal abnormalities.     

The circadian rhythm of thermoregulation in Japanese quail

Japanese quail exhibit a robust circadian rhythm in body temperature. This rhythm is readily entrainable by 24 h light-dark (LD) cycles and persists under constant conditions. Because both the pineal organ and the eyes have been implicated as major components of the circadian system of birds, the role of these organs in generating the rhythm of body temperature was investigated. Pinealectomy, when performed alone, had little effect on the body temperature rhythm of quail either under LD or under constant darkness (DD). Most birds subjected to optic nerve section alone remained rhythmic in DD although the robustness of the rhythm was decreased, and 25% became arrhythmic. Birds subjected to both pinealectomy and optic nerve section behaved similarly to birds subjected to optic nerve section alone. However, complete eye removal, when performed alone or in combination with pinealectomy, caused all birds to become arrhythmic in DD. The data support the hypothesis that the eyes are the loci of circadian pacemakers in quail that act, via both neural and hormonal outputs, to preserve the integrity of (self-sustaining or damped) circadian oscillators located elsewhere.     

Thermoregulatory and metabolic responses of Japanese quail to hypoxia

Common responses to hypoxia include decreased body temperature (T_b) and decreased energy metabolism. In this study, the effects of hypoxia and hypercapnia on T_b and metabolic oxygen consumption (V.O₂) were investigated in Japanese quail (Coturnix japonica). When exposed to hypoxia (15, 13, 11 and 9% O₂), T_b decreased only at 11% and 9% O₂ compared to normoxia; quail were better able to maintain T_b during acute hypoxia after a one-week acclimation to 10% O₂. V.O₂ also decreased during hypoxia, but at 9% O₂ this was partially offset by increased anaerobic metabolism. T_b and V.O₂ responses to 9% O₂ were exaggerated at lower ambient temperature (T_a), reflecting a decreased lower critical temperature during hypoxia. Conversely, hypoxia had little effect on T_b or V.O₂ at higher T_a (36 °C). We conclude that Japanese quail respond to hypoxia in much the same way as mammals, by reducing both T_b and V.O₂. No relationship was found between the magnitudes of decreases in T_b and V.O₂ during 9% O₂, however. Since metabolism is the source of heat generation, this suggests that Japanese quail increase thermolysis to reduce T_b. During hypercapnia (3, 6 and 9% CO₂), T_b was reduced only at 9% CO₂ while V.O₂ was unchanged.     

Glutathione S-transferases in the Japanese quail: Tissue distribution and purification of the liver isozymes

Cytosolic glutathione S-transferase (GST) activities toward 1-chloro-2,4-dinitro-benzene (CDNB), 1,2-dichloro-4-nitrobenzene (DCNB), ethacrynic acid (EA), 1,2-epoxy-3-(p-nitrophenoxyl)propane (EPNP), trans-4-phenyl-3-buten-2-one (t-PBO), δ³-androstene-3,17-dione (ASD) and trans-stilbene oxide (t-SO); cytosolic glutathione peroxidase activity toward cumene hydroperoxide (CuOOH); and microsomal GST activity toward CDNB were examined in liver, kidney, brain, and lung of adult male and female Japanese quail. In all cases, the renal specific activity per milligram protein was higher than the hepatic activity and was the highest among the four tissues examined. No consistent sex differences in GST activity were observed. The GSTs were purified from quail liver cytosol by S-hexylglutathione and glutathione affinity chromatography. Total GSTs eluted from the S-hexylglutathione affinity column were further separated by chromatofocusing, and the microheterogeneity of the GST isozymes was shown by high-resolution native isoelectrofocusing (IEF) in polyacrylamide slab gels and by SDS-PAGE. Five subunits were identified: QL1 (30.5 kDa), QL2 (27.2 kDa), QL3a (26.8 kDa), QL3b (26.5 kDa), and QL4 (25.5 kDa). Western blot analysis revealed that QL1 and QL2 reacted with antibodies raised against the rat Mu class GSTs (Yb1 and Yb2), and QL3a and QL3b reacted with those raised against the Alpha class (rat Ya and mouse a). Substrate specific activity of each isoform was determined with CDNB, DCNB, CuOOH, EA, t-PBO, ASD, and t-SO. QL3a and QL3b have high reactivity toward CuOOH, while QL1 and QL2 showed high activity toward t-SO. The N-terminal amino acid sequence of QL2 was identical to that of the chicken Mu class GST subunit CL2. However, no sequence was obtained with QL1 due to possible N-terminal blockage. © 1996 John Wiley & Sons, Inc.     

Hypothalamic regulation of testicular function in Japanese quail

Summary Some of the components of the system that controls the function of the testes of the Japanese quail were identified by means of midline or bilateral electrolytic or radio-frequency coagulation of various regions of the hypothalamus. The destruction of two identifiable regions of the ventral hypothalamus resulted in different forms of testicular dysfunction. Extensive medial or bilateral damage to the infundibular nuclear complex, ventral regions thereof (including the tuberal nuclei), the entire median eminence or its posterior division resulted in regression of both the gametogenic and endocrine components of the testis and regression of the cloacal gland, an anal gland known to depend on testosterone for normal function. Damage to more anterodorsal regions of the infundibular nuclear complex resulted, in a single bird, in regression of the seminiferous tubules, but had no noticeable effect on the cells of Leydig; the cloacal gland of this bird, although reduced somewhat in size, appeared histologically active. Unilateral damage to the infundibular nuclear complex or the median eminence, or bilateral destruction of the antero—or posteroventral portions of the infundibular nuclear complex did not affect testicular function. The differing testicular dysfunctions are discussed in relation to the release of LH and FSH from the adenohypophysis.This investigation was supported by NIH Predoctoral Fellowship GM-33,458 to the author, and by Research Grants from the National Science Foundation (GB-11905) and the National Institutes of Health (NB-06187) to Professor Donald S. Farner. This paper is based on a dissertation (Stetson, 1971b) submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Zoology at the University of Washington. The hormones used in this investigation were generously supplied by the Endocrinology Study Section of the National Institutes of Health.     

Hypothalamic regulation of gonadotropin release in female Japanese quail

Summary The control mechanisms of the hypothalamo-hypophysio-ovarian axis were investigated in breeding Japanese quail by means of electrolytic or radiofrequency lesions in various regions of the hypothalamus. Depending on the site of damage to the hypothalamus, two forms of ovarian dysfunction were observed. Lesions in anterior regions of the infundibular nuclear complex or in the anterior division of the median eminence resulted in regression of the ovary and oviduct; all follicles of the ovaries of these birds measured less than 3 mm in diameter. These conditions are interpreted as indicative of cessation of the release of folliclestimulating hormone from the adenohypophysis. Lesions in medial, ventral and posterior regions of the infundibular nuclear complex or in the preoptic region of the hypothalamus resulted in cessation of ovulation, but did not cause complete regression of the ovary and oviduct. In these birds the ovaries and oviducts were reduced somewhat in weight, but the ovarian follicles remained large and laden with yolk. This condition is thought to result from the cessation of the release of luteinizing hormone from the adenohypophysis.This investigation was supported by NIH Predoctoral Fellowship GM-33,458 to the author, and by Research Grants from NIH (NB-06187) and NSF (GB-11905) to Professor Donald S. Farner. This paper is based on a thesis (Stetson, 1971) submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy in Zoology at the University of Washington. The hormones used in this investigation were generously supplied by the Endocrine Study Section of the National Institutes of Health.     

A third allele at the phosphoglucose isomerase locus of the Japanese quail

Phosphoglucose isomerase electrophoretic patterns of the Japanese quail were found to be controlled by three alleles at an autosomal locus. In the laboratory quail population, the frequency of the alleles PGI^F, PGI^S1 and PGI^S2 was 0.175, 0.465 and 0.360, respectively.     

Alterations in histones of the liver and oviduct of Japanese quail during aging

Age-related changes occur in histones of the liver and oviduct of the female quail. In the liver an extra histone band, named HCX, between H2A and H4, is observed that increases with age. In the oviduct, a large complex of histones is seen which is tissue-specific. This complex declines with increasing age. The changes in the histones of the oviduct of adult and old birds in response to estradiol and progesterone are age-related. In the adult, the histone-complex increases after administration of either one of the hormones. In old birds, however, it is seen only after progesterone administration. Thus, the alterations in histones in the birds are not only tissue- and age-related, but also vary in response to steroid hormones.     

The role of the subfornical organ in drinking induced by angiotensin in the Japanese quail,Coturnix coturnix japonica

Summary Synthetic 5-valine angiotensin II (AII) induced copious drinking when applied directly to the subfornical organ (SFO) in the Japanese quail. Reliable response was obtained with as little as 1 ng of AII. The amount of water intake increased dose-dependently from 5 ng to 1 ng. A latent period of 73.0 ± 11.0 seconds at 100 ng was noted. The electrical destruction of the SFO significantly reduced the amount of water intake induced by both intravenous and intracranial AII injections. The decrease was proportional to the extent of the SFO lesion. It is conceivable, therefore, that the SFO plays an important role in elicitation of drinking by AII in birds as suggested in mammals.     

Effects of corticosterone treatment on responses to fasting in Japanese quail

Plasma concentrations of corticosterone, a glucocorticoid hormone, have been reported to increase during fasting in some species of birds. Although Japanese quail are widely used in laboratory studies, corticosterone responses to fasting have not been described for this species. We therefore investigated the effects of 24 h of fasting on plasma corticosterone concentrations in quail. Previous work has shown that the corticosterone response to handling in quail may be affected by treatment with corticosterone, so we also measured corticosterone responses to 24 h of fasting in quail treated with corticosterone in their drinking water, and determined the effects of corticosterone on food intake immediately after a fast. Plasma corticosterone concentrations were unaffected by 24 h of fasting in controls and in three out of four groups of birds with varying corticosterone intakes. Fasting in birds can be divided into three phases, and our results suggest that quail remained in phase I during the 24 h fast. Food intake in the 3 h immediately after fasting was higher in four groups of birds with varying corticosterone intakes than in the control group, and was greatest in quail with the highest corticosterone intake. The current results suggest that corticosterone can stimulate food intake in quail following a period of food deprivation.     

Aorta elastin turnover in normal and hypercholesterolemic Japanese quail

The turnover and degradation of mature elastin from the aortae of Japanese quail were estimated following with l-[U-¹⁴C]lysine by measuring the changes in specific activity of l-[U-¹⁴C]lysine and ¹⁴C-labelled desmosine and isodesmosine (crosslinking amino acids derived from lysyl residues) in elastin over a 39-week period. Only 5% of the variation in radioactivity could be attributed to changes in time. Therefore, it was concluded that the best estimates of mature elastin turnover are only quantifiable in years. Dietary cholesterol in amounts sifficient to induce plaque formation and fragmentation of the elastic lamina in the aorta did not significantly influence turnover time. It would appear that once the total pool of elastin in aorta is stabilized as mature fibers it is not subject to proteolysis or resynthesis of sufficient magnitude to result in measurable turnover.     

A first-generation microsatellite linkage map of the Japanese quail

A linkage map of the Japanese quail (Coturnix japonica) genome was constructed based upon segregation analysis of 72 microsatellite loci in 433 F(2) progeny of 10 half-sib families obtained from a cross between two quail lines of different genetic origins. One line was selected for long duration of tonic immobility, a behavioural trait related to fearfulness, while the other was selected based on early egg production. Fifty-eight of the markers were resolved into 12 autosomal linkage groups and a Z chromosome-specific linkage group, while the remaining 14 markers were unlinked. The linkage groups range from 8 cM (two markers) to 206 cM (16 markers) and cover a total map distance of 576 cM with an average spacing of 10 cM between loci. Through comparative mapping with chicken (Gallus gallus) using orthologous markers, we were able to assign linkage groups CJA01, CJA02, CJA05, CJA06, CJA14 and CJA27 to chromosomes. This map, which is the first in quail based solely on microsatellites, is a major step towards the development of a quality molecular genetic map for this valuable species. It will provide an important framework for further genetic mapping and the identification of quantitative trait loci controlling egg production and fear-related behavioural traits in quail.     

AFLP linkage map of the Japanese quail Coturnix japonica

The quail is a valuable farm and laboratory animal. Yet molecular information about this species remains scarce. We present here the first genetic linkage map of the Japanese quail. This comprehensive map is based solely on amplified fragment length polymorphism (AFLP) markers. These markers were developed and genotyped in an F2 progeny from a cross between two lines of quail differing in stress reactivity. A total of 432 polymorphic AFLP markers were detected with 24 TaqI/EcoRI primer combinations. On average, 18 markers were produced per primer combination. Two hundred and fifty eight of the polymorphic markers were assigned to 39 autosomal linkage groups plus the ZW sex chromosome linkage groups. The linkage groups range from 2 to 28 markers and from 0.0 to 195.5 cM. The AFLP map covers a total length of 1516 cM, with an average genetic distance between two consecutive markers of 7.6 cM. This AFLP map can be enriched with other marker types, especially mapped chicken genes that will enable to link the maps of both species and make use of the powerful comparative mapping approach. This AFLP map of the Japanese quail already provides an efficient tool for quantitative trait loci (QTL) mapping.