Ultradian Rhythm of Activity in Japanese Quail Groups under Semi-Natural Conditions during Ontogeny: Functional Aspects and Relation to Circadian Rhythm

The function of ultradian rhythms is not yet clearly elucidated. In particular, short-term rhythms are expressed during early ontogeny, especially in broods of precocial birds. We investigated the relationship between the clarity of the ultradian rhythm of the activity/rest cycle of a group of young Japanese quail (Coturnix japonica) and the level of social synchronisation and spatial cohesion between the birds within that group. The subjects were descended from two lines selected for either very pronounced rhythmic or arrhythmic circadian activity. We found a positive relationship between the clarity of the ultradian rhythm of the activity/rest cycle when birds were young and the clarity of the circadian rhythm of feeding activity when birds were older, but still immature. The temporal organisation of the behaviour of the chicks from these two lines was observed in outdoor aviaries, when they were 4, 8, 12 and 15 days old. The mean ultradian period expressed by groups of 12 chicks was variable, with a minimum of 6 minutes. The ultradian period lengthened regularly as chicks grew older, and reached approximately 40 min on day 15. The clarity of the ultradian rhythmicity of group activity was linked to the level of inter-individual social synchronisation and of spatial cohesion; the more pronounced the ultradian rhythms of a group, the greater the temporal and spatial cohesion of the chicks within the group. Moreover, these characteristics varied with the age of the chicks. Finally, chicks in the less rhythmic groups weighed less. These results stress the adaptive value of this temporal organisation strategy under natural conditions.     

Ultradian Rhythm of Activity in Japanese Quail Groups under Semi-Natural Conditions during Ontogeny: Functional Aspects and Relation to Circadian Rhythm

The function of ultradian rhythms is not yet clearly elucidated. In particular, short-term rhythms are expressed during early ontogeny, especially in broods of precocial birds. We investigated the relationship between the clarity of the ultradian rhythm of the activity/rest cycle of a group of young Japanese quail (Coturnix japonica) and the level of social synchronisation and spatial cohesion between the birds within that group. The subjects were descended from two lines selected for either very pronounced rhythmic or arrhythmic circadian activity. We found a positive relationship between the clarity of the ultradian rhythm of the activity/rest cycle when birds were young and the clarity of the circadian rhythm of feeding activity when birds were older, but still immature. The temporal organisation of the behaviour of the chicks from these two lines was observed in outdoor aviaries, when they were 4, 8, 12 and 15 days old. The mean ultradian period expressed by groups of 12 chicks was variable, with a minimum of 6 minutes. The ultradian period lengthened regularly as chicks grew older, and reached approximately 40 min on day 15. The clarity of the ultradian rhythmicity of group activity was linked to the level of inter-individual social synchronisation and of spatial cohesion; the more pronounced the ultradian rhythms of a group, the greater the temporal and spatial cohesion of the chicks within the group. Moreover, these characteristics varied with the age of the chicks. Finally, chicks in the less rhythmic groups weighed less. These results stress the adaptive value of this temporal organisation strategy under natural conditions.     

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)     

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)     

Rhythmic birds show a better social integration than arrhythmic birds

The activity rhythms of Japanese quail vary from one individual to another. Performing a divergent selection, we obtained one line of quail expressing a robust circadian rhythmicity of feeding activity (R) and one line of quail expressing circadian arrhythmicity of feeding activity (A). We questioned whether the endogenous rhythmicity of an individual could predict its integration in a group. For that, we introduced either an R- or an A-line chick into stable groups of standard chicks. First, we evaluated proximity and synchronization of the introduced chicks on the activities of the other group members. R-chicks remained spatially and temporally closer to other group members than did A-chicks. Second, we evaluated interactions of the introduced chicks and the level of their acceptance by the other group members. R-chicks were more competent to gain access to food than were A-chicks, and separation from their group stressed R-chicks more than A-chicks. Last, successive introductions assessed age effects: before, around, and after dispersal time (～11th day of chicks' life). Most differences between R- and A-chicks were observed between their 7th and 15th day of life. In conclusion, individual endogenous rhythms predict social integration.     

Rhythmic Birds Show a Better Social Integration than Arrhythmic Birds

The activity rhythms of Japanese quail vary from one individual to another. Performing a divergent selection, we obtained one line of quail expressing a robust circadian rhythmicity of feeding activity (R) and one line of quail expressing circadian arrhythmicity of feeding activity (A). We questioned whether the endogenous rhythmicity of an individual could predict its integration in a group. For that, we introduced either an R- or an A-line chick into stable groups of standard chicks. First, we evaluated proximity and synchronization of the introduced chicks on the activities of the other group members. R-chicks remained spatially and temporally closer to other group members than did A-chicks. Second, we evaluated interactions of the introduced chicks and the level of their acceptance by the other group members. R-chicks were more competent to gain access to food than were A-chicks, and separation from their group stressed R-chicks more than A-chicks. Last, successive introductions assessed age effects: before, around, and after dispersal time (～11th day of chicks' life). Most differences between R- and A-chicks were observed between their 7th and 15th day of life. In conclusion, individual endogenous rhythms predict social integration. (Author correspondence: sophie.lumineau@univ-rennes1.fr)     

Ultradian and circadian compartmentalization of respiratory and metabolic exchanges in small laboratory vertebrates

Carbon dioxide emission (VCO2) taken as an index of respiratory and metabolic exchanges, was continuously recorded during 4-30 consecutive days in 100 quail, 87 chicks, 347 rats, 665 mice and 70 guinea-pigs which were under controlled environmental parameters. Harmonic analysis, fast Fourier transform, chi-square periodograms, peak and trough intervals were computed with VCO2 values obtained with CO2 concentrations sampled every 20 min on the CO2 recordings. In LD 12:12 alternation, circadian rhythms were observed in all quail, chicks, rats and mice, but only in 80% of the guinea-pigs. Ultradian VCO2 rhythms, with periods which show statistically significant interspecies differences, were assessed. For each of the 5 species these computed periods, which were the same in LL and DD, were: 1.17 h for quail and chickens, 1.25 h for rats, 1.50 h for mice and 1.0 h for guinea-pigs. In LD 12:12 these periods were different during L and D in quail, chicks, rats and mice, but not in guinea-pigs. The amplitudes of these ultradian variations were, according to the species, 10-20% of their mean VCO2 levels. These ultradian rhythms persist in the absence (or masking) of circadian rhythms, e.g. in LD 12:12 in 20% of guinea-pigs and in LL in 87% of Japanese quail and in 23% of Sprague-Dawley rats. Moreover, these ultradian rhythms persist during starvation, locomotor activity restraint and ageing. These ultradian VCO2 cycles which are related to rest-activity variations appear to be basic physiological rhythms with a genetic origin.     

Ultradian activity rhythms in large groups of newly hatched chicks (Gallus gallus domesticus)

A clutch of young chicks housed with a mother hen exhibit ultradian (within day) rhythms of activity corresponding to the brooding cycle of the hen. In the present study clear evidence was found of ultradian activity rhythms in newly hatched domestic chicks housed in groups larger than natural clutch size without a mother hen or any other obvious external time-keeper. No consistent synchrony was found between groups housed in different pens within the same room. The ultradian rhythms disappeared with time and little evidence of group rhythmicity remained by the third night. This disappearance over time suggests that the presence of a mother hen may be pivotal for the long-term maintenance of these rhythms. The ultradian rhythm of the chicks may also play an important role in the initiation of brooding cycles during the behavioural transition of the mother hen from incubation to brooding. Computer simulations of individual activity rhythms were found to reproduce the observations made on a group basis. This was achievable even when individual chick rhythms were modelled as independent of each other, thus no assumptions of social facilitation are necessary to obtain ultradian activity rhythms on a group level.     

Inter-individual variability during morning choruses in Japanese quail males (Coturnix coturnix japonica)

We compared the organization of dawn choruses in five groups of Japanese quail, Coturnix coturnix japonica, males (four groups of kin-related males and one group of unrelated males). We documented the structure of chorus groups: one male sang much more than the others during the dawn peak in four of the five groups, and this regardless of the number of males in the group. The dominant singers were then removed from two similar-sized groups, one of kin-related (K3) and the other of unrelated (NK) males. The quantity of song emitted by the remaining NK males was modified, but not that emitted by the K3 males.     

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.     

Influence of different photoperiods on development of gonad, cloacal gland and circulating thyroid hormones in male Japanese quail Coturnix coturnix japonica.

One day old chicks of Japanese quail were exposed to different photoperiods (LD, 8:16, 13.5:10.5, 16:8 and LL) and observations (testes weight, cloacal gland size, body weight and circulating thyroxine and triiodothyronine) were taken at the age of 3, 5, 7, 9 and 16 weeks. Results indicate that immediate reproductive development occurred in birds exposed to long photoperiods (greater than 12 hr). Growth under LD 8:16, was not apparent till 7th week and by 16 weeks, degree of gonadal development was similar in all the birds, irrespective of photoperiodic treatment. Whereas body weight of the intermediate and long day (LD 13.5:10.5, 16:8 and LL) treated birds increased upto 5th week and remained constant thereafter. But the chicks maintained under short day length (LD 8:16), showed spontaneous increase till the end of the study and birds were much heavier compared to all other groups. Plasma T4 concentration increased with increasing age till 9th week and remained unaltered thereafter. On the other hand T3 level did not change till 7th week followed by a decline. It is suggested that the initiation and degree of gonadal growth in quail depends on the availability of daily photoperiod, until the achievement of full breeding condition. Peak level of T4 observed in 9 week old birds may be involved in the development of photorefractoriness at that age.     

Post-embryonic development of chicks of the Japanese quail under conditions of the limited volume

According to our results, chicks of the Japanese quail can survive within the first 10 days of a life under conditions of the limited volume when their mobility is completely limited. However, their development was impaired in comparison with chicks housed in the common cages. Besides, the delay in development of chick's plumage has been observed. The analysis of body mass in test groups has shown that practically all chicks were smaller than chicks from control group. It is the result of feed consumption decrease (about 30% from control group) caused by the restriction of mobility of chicks. But after their transferring to the common cages at the end of the experiment, body mass in test group rapidly reached the level of body mass in control group. Research has shown that percent of alive of quail chicks maintained in the chamber of animals of complex "Incubator-2", was higher than in first two experiments. Therefore, works on improvement of devices for the maintenance of quail chicks continue, and also new adaptations develop for the maintenance of chicks in their early post-embryonic period (till 10 days) with reference to conditions of weightlessness.     

Mucuna pruriens seed powder feeding influences reproductive conditions and development in Japanese quail Coturnix coturnix japonica

This study was designed to test whether Mucuna pruriens, a natural source of l-dihydroxyphenylalanine (l-DOPA, a dopamine precursor) feeding, can influence development and reproductive conditions in the high food value bird, Japanese quail, Coturnix coturnix japonica. Experiments were performed in both male and female Japanese quail. One-week-old quail chicks were divided into three groups of 36 birds each. Group I was provided with normal diet and served as control. Group II was provided with food mixed with l-DOPA (50 mg/15 g food) and Group III was provided with food mixed with M. pruriens seed powder (480 g/kg food). At the age of 3 weeks (when birds were sexually distinguished) Group I was divided into two sub-groups IA (male) and IB (female) of six birds each. Similarly, Groups II and III were sub-divided into IIA (male), IIB (female) and IIIA (male), IIIB (female), respectively, of six birds each. Observations were made up to the age of 5 weeks. Male experimental groups (IIA and IIIA) showed significantly increased testicular activity, cloacal gland volume, body weight (BW), plasma testosterone and LH level in comparison to control (IA). Similarly female experimental groups (IIB and IIIB) showed significantly greater weight of reproductive organs (uterus, ovary, oviduct and ovarian follicle), BW, egg weight and size and number of follicles. On the other hand, plasma prolactin level was significantly low in comparison to control (IB). Results suggest that M. pruriens is a rich natural source of l-DOPA and the development and reproduction in Japanese quail might be associated with the dopaminergic system of the brain.     

Increased yolk testosterone facilitates prenatal perceptual learning in Northern bobwhite quail (Colinus virginianus)

Prenatal learning plays an important role in the ontogeny of behavior and birds provide a useful model to explore whether and how prenatal exposure to hormones of maternal origin can influence prenatal learning and the development of behavior. In this study we assessed if prenatal exposure to yolk testosterone can influence auditory learning in embryos of Northern bobwhite quail (Colinus virginianus). We experimentally enhanced testosterone concentrations in bobwhite quail eggs prior to incubation. The embryos from these T-treated eggs as well as control embryos that had received the vehicle-only or were non-treated were exposed to an individual bobwhite hen's maternal call for 120 min over the course of the day prior to hatching. All chicks were tested at 24 h following hatching for their auditory preference between the familiar bobwhite maternal call versus an unfamiliar bobwhite maternal call. T-treated chicks spent significantly more time in proximity to the familiar call compared to the unfamiliar call and also showed shorter latencies to approach the familiar call than control birds. Increased emotional reactivity, i.e. propensity to express fear responses, was also found in T-treated chicks. Baseline heart rates recorded in a second group of T-treated embryos and control embryos did not differ, which suggests no effect of yolk testosterone on baseline arousal level. To our knowledge this is the first demonstration of the influence of prenatal exposure to testosterone on auditory learning.     

Pair and group breeding behaviour of a communal gallinule,the pukeko, Porphyrio P. Melanotus

Pukeko, Porphyrio porphyrio melanotus breed as pairs or groups and the breeding behaviour of individuals in all units is described and compared. Courtship, including copulation, occurred between all adult members in a territory although male-female copulations were most frequent. Low ranking yearlings were excluded from copulation. Participation in copulation by more than two birds was common, and, along with homosexual behaviour, is believed to help synchronize sexual cycles allowing several females to lay in the same nest at the same time. Only adults incubated, while all birds in a territory assisted with chick care, the degree of participation varying with status. In pair territories, older chicks helped feed younger chicks. Stable kin groups were more coordinated and showed less competitive aggression among members than non-kin groups of unstable membership. The significance of these results is discussed and it is argued that in a situation where an adult group is necessary for the defence of a breeding territory, then it is in the interests of all these individuals to participate in all breeding activities.     

Epigenetic Maternal Effects on Endogenous Rhythms in Precocial Birds

Development involves interactions between genetic and environmental influences. Vertebrate mothers are generally the first individuals to encounter and interact with young animals. Thus, their role is primordial during ontogeny. The present study evaluated non‐genomic effects of mothers on the development of rhythms of precocial Japanese quail (Coturnix c. japonica). First, we investigated the influence of mothering on the ontogeny of endogenous rhythms of young. We compared circadian and ultradian rhythms of feeding activity of quail reared with or without adoptive mothers. More brooded than non‐brooded quail presented a circadian and/or an ultradian rhythm. Thus, the presence of the mother during the normal brooding period favors, in the long term, expression of rhythms in the young. Second, we investigated the influence of rhythmic phenotype of the mother on the development of endogenous rhythms of young by comparing quail brooded by circadian‐rhythmic adoptive mothers (R) to quail brooded by circadian‐arrhythmic adoptive mothers (A). More R‐brooded than A‐brooded quail expressed circadian rhythmicity, and circadian rhythm clarities were greater in R‐brooded than A‐brooded quail. Ultradian rhythmicity did not differ between R‐ and A‐brooded quail, nor between R and A adoptive mothers. Thus, the rhythmic phenotypes of quail mothers influence the rhythmic phenotypes of their young. Our results demonstrate that mothers of precocial birds influence epigenetically the ontogeny of endogenous rhythms of the young they raise.     

Ultradian and circadian CO2 emission variations in nocturnal and diurnal animals exposed to a light stimulus

1. Carbon dioxide emission (VCO2) has been continuously recorded in three laboratory animal species (Sprague-Dawley rats, Japanese quail, Hartley guinea-pigs) which differ by their nocturnal and diurnal activities. A 100 lux stimulus has been delivered at various time intervals. 2. A regular alternation of 12, 3 or 1.5 hr light (L) and darkness (D) gives VCO2 circadian and ultradian rhythms of 24, 6 or 3 hr periods, respectively, in quail and rats. 3. Such circadian and ultradian LD rhythms are not induced in all guinea-pigs. 4. The amplitudes of the VCO2 responses are greatest at D----L when the animals have a maximum diurnal activity and at L----D when their maximum activity is nocturnal. 5. Interactions between circadian and ultradian rhythms are seen in all LD experiments, as well as in continuous light (LL) or continuous dark (DD). 6. No more well-marked or even inverted VCO2 responses to the light stimuli may occur after several days of exposure to these LD alternations.     

Ontogenetic development of conditioned food aversion in chickens

Development of conditioned food aversion (CFA) was studied in 25-, 35-, and 45-day Leghorn chicks. Food-deprived birds had 10-min access to normal food on days 1 and 2, and to green coloured food CS on day 3. Injection of LiCl (0.15 M, 3–4% body weight) administered 10 min after CS on day 3 served as the US. Control groups were injected with the same volume of saline. Retention was tested on day 4 in three 3-min presentations of normal food alternating with two presentations of green food. Number of pecks and amount of food consumed were measured. Significant neophobic rejection of green food was observed in 25- but not in 35- and 45-day-old control birds. Neophobia in the youngest chicks was further accentuated by CFA which could be observed in pure form in the 45-day-old experimental group. Comparison of pecking rate and food intake showed that CFA in younger birds was accomplished predominantly by inhibition of pecking, and in older birds also by reduction of peck volume. It is concluded that protection against poisoning in chickens shifts between 25 and 45 days of posthatching age from neophobia to CFA.     

Activity Monitoring of the Inhabitants in Tauwema, a Traditional Melanesian Village: Rest/Activity Behaviour of Trobriand Islanders (Papua New Guinea)

The inhabitants of Tauwema village represent a traditionally living society. Altogether, 39 inhabitants belonging to seven families were included in this study. Families or people living in one household were preferentially chosen for monitoring particularly the effects of social zeitgebers and synchronization within families. They were observed continuously for 7 consecutive days using microelectronic actometers that register locomotor activity with a sampling period of 2 minutes and a resolution of 7 bit. The activity data obtained showed that in young infants circadian patterns develop out of ultradian components dominating in the first months of life. The rhythmicity of the adults was well-related to the natural light-dark cycle, combined with a strong social component which is reflected in a comparatively small intra- and inter-individual variability in the time of the end of the main sleep span in the morning, while the variability in the beginning of the main sleep period in the evening is much greater. The mean sleep duration of the younger infants (up to 11 months; n =4) varied between 9 and 12 h per day and that of the adults ( n = 23) between 7 and 10 h. Gender-specific differences occurred in married couples with wives having a longer sleep duration in 7 out of 9 cases.     

Activity Monitoring of the Inhabitants in Tauwema,a Traditional Melanesian Village: Rest/Activity Behaviour of Trobriand Islanders (Papua New Guinea)

The inhabitants of Tauwema village represent a traditionally living society. Altogether, 39 inhabitants belonging to seven families were included in this study. Families or people living in one household were preferentially chosen for monitoring particularly the effects of social zeitgebers and synchronization within families. They were observed continuously for 7 consecutive days using microelectronic actometers that register locomotor activity with a sampling period of 2 minutes and a resolution of 7 bit. The activity data obtained showed that in young infants circadian patterns develop out of ultradian components dominating in the first months of life. The rhythmicity of the adults was well-related to the natural light-dark cycle, combined with a strong social component which is reflected in a comparatively small intra- and inter-individual variability in the time of the end of the main sleep span in the morning, while the variability in the beginning of the main sleep period in the evening is much greater. The mean sleep duration of the younger infants (up to 11 months; n =4) varied between 9 and 12 h per day and that of the adults (n = 23) between 7 and 10 h. Gender-specific differences occurred in married couples with wives having a longer sleep duration in 7 out of 9 cases.