Season of birth and morningness: comparison between the northern and southern hemispheres

The present study explored the possible role of the photoperiod at birth on morningness by collecting data in the northern (Italy) and southern (Australia) hemispheres. To assess circadian typology, the Composite Scale of Morningness (CS) was administered to a sample of 1734 university students (977 Italian and 757 Australian; 1099 females and 635 males; age 24.79?±?7.45 yrs [mean?±?SD]). Consistent with the literature, females reported higher CS scores (morningness) than males, and Australian participants reported higher CS scores than Italian participants. Allowing for the fact the seasons are reversed between the hemispheres, the results are in line with previous studies. The authors found more evening types were born during the seasons associated with longer photoperiod (spring and summer), and more morning types were born during the seasons associated with shorter photoperiod (autumn and winter), indirectly supporting an imprinting-like phenomenon played by the photoperiod at birth.     

The effect of pregnant and oestrous females on male testosterone and behaviour in the tammar wallaby

Tammar wallaby females (Macropus eugenii) are seasonally breeding marsupials with a post-partum oestrus after a highly synchronised birth period when testosterone concentrations rise in males. Chemical communication appears to be important for mating, as males show checking behaviour, sniffing the urogenital opening (UGO) and the pouch of females. This study investigates whether the presence of pregnant and oestrous females directly influences testosterone in males and if oestrous odours or secretion from the pouch or UGO are attractive. Concentrations of plasma testosterone were measured in males housed with pregnant and oestrous females during two consecutive cycles in the breeding season, and an artificially induced cycle in the non-breeding season. Males were also tested for their interest in swabs taken from the urogenital opening (UGO) or pouch of oestrous females. Testosterone increased sharply in males in the presence of pregnant and oestrous females during all cycles in both seasons, but there was no change when males were exposed to non-cycling females in lactational or seasonal diapause. Males had no preference for either oestrous or non-oestrous samples taken from the pouch or from the UGO from oestrous females. This study confirms that the increase in plasma testosterone in tammar males can be induced through the presence of pregnant and oestrous females, regardless of season and that the increase began when the females were in late-pregnancy. This confirms that the male's reproductive state is dependent on a signal from females and is not blocked through seasonal effects.     

Timing of puberty and synchronization of seasonal rhythms by simulated natural photoperiods in female Siberian hamsters

The timing of puberty is a critical life history trait of short-lived species; spring-born individuals mature rapidly and breed in the season of birth, whereas young born in mid- to late summer delay puberty until the next spring. The cues that govern the transition from rapid to delayed maturation in natural populations remain unknown. To identify ecologically relevant photoperiod cues that control timing of puberty, we monitored nine cohorts of female Siberian hamsters (Phodopus sungorus) born every 2 wk from 4 wk before to 12 wk after the summer solstice in a simulated natural photoperiod (SNP). Hamsters born by the summer solstice underwent rapid somatic growth and achieved puberty that summer; among females born 2-4 wk after the solstice, some delayed puberty by many weeks, whereas others manifested early puberty. Hamsters born 6 or more weeks after the solstice generally delayed puberty until the following spring. The transition from accelerated to delayed pubertal development in the SNP occurred at day lengths that induce early puberty when presented as static photoperiods. Despite differences in timing of birth and timing of puberty, fall and subsequent spring seasonal events occurred at similar calendar dates in all cohorts. We found no evidence that prenatal photoperiod history influenced postnatal development of female hamsters. Considered together with a parallel study on males, the present findings point to sex differences in responsiveness to natural photoperiod variations. In both sexes, incrementally changing photoperiods exert a strong organizing effect on seasonal rhythms.     

Behavioural and hormonal responses of male rhesus monkeys introduced to females in the breeding and non-breeding seasons.

Six adult male rhesus monkeys were introduced individually to an all-female group for 10 days during the mating season. The initial aggressive responses of the females were rapidly replaced by positive social behaviour, and each male achieved alpha status and had access to social and sexual partners. A repetition of this paradigm in the non-breeding season produced significantly more female aggression, and no male attained high rank or engaged in sexual or other social behaviour. Male testosterone levels rose following introduction to the females in both seasons, but were significantly higher during the breeding season. Hormonal levels following removal from the females suggest a complex interplay between social, sexual and seasonal variables and recent social experiences. The differences in female social behaviour with newly introduced males, as a function of season, suggest an explanation for the seasonal limitation of male troop transfers.     

Seasonality of births in Homo sapiens in pre-industrial Finland: maximisation of offspring survivorship?

We examined the seasonal variation in human birth and infant survival rates in pre-modern Finland. If survival probabilities of children born during different seasons of the year differ and if timing of reproduction has been affected by natural selection, periodic variation in environment could have led to reproduction during the season of best infant survival expectations. Significant seasonal variation in both birth rate and survival probability was found, but the monthly birth and survival rates of newborn were uncorrelated. Hence, if there was any tendency to maximise the reproductive success, increase in some other component of fitness than the infant survival was probably targeted. The effect of major holidays on the birth rate was proved to be notable, suggesting that although the basis for seasonal variation in birth rate was biological, sociocultural factors had an impact on the timing of reproduction in humans. The overall mortality of infant boys exceeded that of infant girls in all seasons. This difference was smallest during the time of best food supplies, indicating that the development of males was less buffered against environmental disturbances than that of females.     

Season of Birth and Morningness: Comparison Between the Northern and Southern Hemispheres

The present study explored the possible role of the photoperiod at birth on morningness by collecting data in the northern (Italy) and southern (Australia) hemispheres. To assess circadian typology, the Composite Scale of Morningness (CS) was administered to a sample of 1734 university students (977 Italian and 757 Australian; 1099 females and 635 males; age 24.79?±?7.45 yrs [mean?±?SD]). Consistent with the literature, females reported higher CS scores (morningness) than males, and Australian participants reported higher CS scores than Italian participants. Allowing for the fact the seasons are reversed between the hemispheres, the results are in line with previous studies. The authors found more evening types were born during the seasons associated with longer photoperiod (spring and summer), and more morning types were born during the seasons associated with shorter photoperiod (autumn and winter), indirectly supporting an imprinting-like phenomenon played by the photoperiod at birth. (Author correspondence: vincenzo.natale@.unibo.it)     

Influence of gonadal hormones on odours emitted by male meadow voles (Microtus pennsylvanicus).

Free-living male meadow voles (Microtus pennsylvanicus) emit odours that are attractive to females at the beginning, but not at the end, of the breeding season. The effect of gonadal hormones on female-attractant cues was examined in males born and reared in long (14 h light day-1) and short (10 h light day-1) photoperiods that simulate daylengths in the breeding and nonbreeding seasons, respectively. Gonadectomy affected the attractant properties of odours emitted by long photoperiod, but not short photoperiod, males. Long photoperiod females preferred odours of intact rather than those of gonadectomized long photoperiod males, and odours of gonadectomized long photoperiod males rather than those of intact short photoperiod males. Females did not show a preference between the odours of intact and castrated short photoperiod males. Gonadal hormone replacement in males affected female responses to the odours emitted by long photoperiod, but not short photoperiod, gonadectomized males. Long photoperiod females did not display a preference between odours of intact long photoperiod males and gonadectomized long photoperiod males treated with testosterone or oestradiol. We conclude that in spring and summer gonadal hormones increase attractiveness of male odours; this effect may require aromatization of testosterone to oestradiol. Substrates that control attractiveness of odour cues in male voles appear to be unresponsive to androgens during the nonbreeding season.     

Season of birth affects juvenile survival of giraffe

Variation in timing of reproduction and subsequent juvenile survival often plays an important role in population dynamics of temperate and boreal ungulates. Tropical ungulates often give birth year round, but survival effects of birth season for tropical ungulate species are unknown. We used a population of giraffe in the Tarangire Ecosystem of northern Tanzania, East Africa to determine whether calf survival varied by season of birth. Variation in juvenile survival according to season of birth was significant, with calves born during the dry season experiencing the highest survival probability. Phenological match may confer a juvenile survival advantage to offspring born during the dry season from greater accumulated maternal energy reserves in mothers who conceive in the long rainy season, high-protein browse in the late dry-early short rainy seasons supplementing maternal and calf resources, reduced predation due to decreased stalking cover, or some combination of these. Asynchrony is believed to be the ancestral state of all ungulates, and this investigation has illustrated how seasonal variation in vegetation can affect juvenile survival and may play a role in the evolution of synchronous births.     

Effects of maternal birth season on birth seasonality in the Canadian population during the seventeenth and eighteenth centuries

Birth records of the French-Canadian population for the period 1621-1765 were analyzed retrospectively to examine the effect of maternal birth season on the seasonal distribution of births. Preliminary examination indicated that there was a bimodal pattern in birth seasonality: a major peak in early spring, a trough in early summer, a minor peak in autumn, and a trough around December. Because this seasonality was strongly biased at the level of the first birth by the month of marriage, which was concentrated in November, the seasonality of nonfirst births (n = 32,926) was examined in relation to the four seasons of maternal birth. Mothers born in May-July showed a flatter monthly distribution of nonfirst births at a maternal age of 28 years or more. Analysis of marriage-first birth intervals indicated that mothers who married in August-October showed a lower percentage of immediate conception (intervals of 8-10 months), whereas those mothers born in May-July had a higher percentage of immediate conception. This difference in birth seasonality shown by mothers born in May-July is similar to results from early twentieth-century Japan. Some seasonal infertility factors could have affected the embryos at the earliest stage of pregnancy, modifying a part of the seasonal variation in birth rate.     

Human responses to the geophysical daily, annual and lunar cycles

Collectively the daily, seasonal, lunar and tidal geophysical cycles regulate much of the temporal biology of life on Earth. The increasing isolation of human societies from these geophysical cycles, as a result of improved living conditions, high-quality nutrition and 24/7 working practices, have led many to believe that human biology functions independently of them. Yet recent studies have highlighted the dominant role that our circadian clock plays in the organisation of 24 hour patterns of behaviour and physiology. Preferred wake and sleep times are to a large extent driven by an endogenous temporal program that uses sunlight as an entraining cue. The alarm clock can drive human activity rhythms but has little direct effect on our endogenous 24 hour physiology. In many situations, our biology and our society appear to be in serious opposition, and the damaging consequences to our health under these circumstances are increasingly recognised. The seasons dominate the lives of non-equatorial species, and until recently, they also had a marked influence on much of human biology. Despite human isolation from seasonal changes in temperature, food and photoperiod in the industrialised nations, the seasons still appear to have a small, but significant, impact upon when individuals are born and many aspects of health. The seasonal changes that modulate our biology, and how these factors might interact with the social and metabolic status of the individual to drive seasonal effects, are still poorly understood. Lunar cycles had, and continue to have, an influence upon human culture, though despite a persistent belief that our mental health and other behaviours are modulated by the phase of the moon, there is no solid evidence that human biology is in any way regulated by the lunar cycle.     

Seasonal, sex, and interspecific differences in activity time budgets and diets of patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus), living sympatrically in northern Cameroon

I examined seasonal, sex, and interspecific differences in activity time budgest and diets of patas (Erythrocebus patas) and sympatric tantalus monkeys (Cercopithecus aethiops tantalus) on the basis of 5-day data sets collected in three and two different seasons, respectively, by the method of focal animal sampling. The seasons included species-specific mating and birth seasons. As compared with not only the birth season but also conspecific females, both patas resident male and tantalus male spent less time feeding and more time resting, day and night, in their respective mating seasons. Given that day-resting time includes time for vigilance for non-resident males and receptive females, this may reflect that males should minimize time spent feeding to allow maximum participation in other fitness-increasing activities such as mating-relating activities asSchoener (1971) predicted. In both species, the males consumed fruits containing less protein but more calories and showed a high feeding rate to compensate for the shorter time spent feeding in the mating season. In contrast, females consumed protein-rich food types (i.e. animals, protein-rich seeds, leaves, and flowers) in the birth season to meet the high demand for protein due to pregnancy and lactation. Given that the season for males was considered to be not a calendar but a reproductive “season” (i.e. mating or birth season), both sexes of patas spent more time moving and less time day- and night-resting than did the tantalus counterparts irrespective of the “season”. Patas subsisted on fruits, gums, and supplementarily lipid-rich seeds as an energy source and animal matters and protein-rich seeds as a protein source. In contrast, tantalus subsisted on fruits and lipid-rich seeds as energy and flowers and leaves as protein.     

Influence of season and sex on the inhibitory effect of ovine follicular fluid on plasma gonadotrophins in gonadectomized sheep

The inhibitory effects of follicular fluid on FSH secretion were similar in gonadectomized male and female sheep, and in the anoestrous and breeding seasons. Significant suppression of LH was variable and was observed only at the highest dose of follicular fluid when suppression rarely exceeded 50% of pretreatment values. Basal plasma FSH and LH concentrations were higher in castrated males than in ovariectomized females in both seasons. Plasma FSH concentrations in gonadectomized males and females and LH concentrations in the males were lower in the anoestrous than the breeding season. Therefore, in the absence of the gonads, sex and photoperiod can influence hypothalamic control of basal pituitary gonadotrophin secretion in males and females, whereas the feedback effect of non-steroidal factors in follicular fluid (inhibin) on FSH secretion is not influenced by photoperiod or sex.     

Birth season glucocorticoids are related to the presence of infants in sifaka (Propithecus verreauxi)

The responses of plural breeding mammals to environmental stressors are little understood in free-ranging populations, but recent studies of singular breeders suggest that ecological factors and social milieu influence the variable physiological stress responses observed among individuals. Our previous studies examining faecal glucocorticoid (fGC)–behaviour interactions in plural breeding male sifaka (Propithecus verreauxi) show that fGC elevations coincide with specific dispersal events, particularly the eviction of subordinates by resident alpha males. This study examined the utility of fGC assays for assessing the stress responses of male sifaka to demographic changes in the population during the birth season. Analyses were based on 889 faecal samples collected over five birth seasons from 124 adult males residing in 55 different groups at Beza Mahafaly, Madagascar. Results show that fGC levels in males are unrelated to age, residence, group stability or rank, but are significantly and positively correlated with the presence of infants, annual elevations in weekly mean fGC concentrations in males paralleling increasing numbers of infants born annually in the population. These data are the first to show that in seasonal plural breeding species such as sifaka, elevated fGC in males reflects specific events related to reproduction rather than states or social context during the birth season.     

粪便激素水平反映不同出生时期雄性布氏田鼠的繁殖策略

布氏田鼠是我国内蒙古草原的主要害鼠之一,具有明显的季节繁殖特征,不同季节出生个体可能具有不同的繁殖策略,但尚缺乏内分泌证据支持。本研究采用标志重捕法,连续监测大型自然围栏中不同年龄雄性布氏田鼠的繁殖状态和应激水平的季节变化,分析各年龄组雄鼠的繁殖发育策略。结果表明,越冬雄鼠可保持较高的睾丸下降率和睾酮水平至8月初;而部分5月生雄鼠的繁殖期睾酮水平较接近越冬鼠,但8月初已降至年内最低水平,而6月及以后出生雄鼠睾酮始终处于较低水平;这说明越冬鼠的繁殖状态可贯穿繁殖期始终,只有部分5月生雄鼠可能参与当年繁殖,而6月及以后出生雄鼠则不能在当年繁殖。越冬鼠皮质醇水平高于当年鼠,繁殖期高于非繁殖期,这可能是越冬鼠由于繁殖需要而保持较高应激状态,从而造成繁殖盛期后死亡率升高。这些结果说明,不同时期出生的雄性布氏田鼠具有不同的出生后性腺发育模式和繁殖策略,反映出生存与繁殖之间的权衡。     

Seasonal differences in self-grooming in meadow voles, Microtus pennsylvanicus

We determined whether seasonal differences exist in the amount of time meadow voles, Microtus pennsylvanicus, self-groom when they encounter the scents of conspecifics. To do so, we used voles that were born and reared under long photoperiod (LP) and short photoperiod (SP). LP voles represent those found in free-living populations during the spring and summer breeding season, whereas SP voles characterize those found in free-living populations during the fall and winter nonbreeding season. Experiment 1 showed that LP male and female voles self-groomed more in response to odors of LP opposite-sex conspecifics as compared to those of other LP and SP conspecifics, suggesting that they may be self-grooming to signal sexual interest or excitement to potential mates. Experiment 2 demonstrated that SP males self-groomed more in response to scents of LP female voles and those of SP males as compared to scents of LP males and SP females, whereas SP females spent similar amounts of time self-grooming in response to scents of LP males, LP females, SP females, and SP males. These seasonal differences in self-grooming may reflect differences in the messages produced by groomers when they broadcast their odors as well as differences in the meaning of such odors to opposite-sex conspecifics. Alternatively, these data may be associated with seasonal differences in sexual motivation of the groomers when exposed to scents of particular conspecifics.     

Comparison of reproductive investment in native and non‐native populations of common wall lizards reveals sex differences in adaptive potential

Non‐native animals can encounter very different environments than those they are adapted to. Functional changes in morphology, physiology and life‐history following introduction show that organisms can adapt both fast and efficiently. It remains unclear, however, if female reproductive characters and male sexually selected behaviour show the same adaptive potential. Furthermore, the invasion success and evolutionary trajectory of non‐native species might often depend on the ability of the sexes to coordinate shifts in characters associated with reproductive strategy. The common wall lizard, Podarcis muralis, has been repeatedly introduced from southern Europe to England over the past 80 years. Lizards in England experience a cool, seasonal climate that effectively restricts recruitment to the first clutch of the season, whereas in their native range up to three clutches per season recruit. As a consequence, both females and males in non‐native populations should benefit from reducing or even eliminating their reproductive investment in second clutches. Using a combination of field data and experiments, we show that non‐native females produce relatively larger and heavier first seasonal clutches and smaller and lighter second seasonal clutches compared to native females. In contrast, non‐native and native males do not differ in their territorial and sexual behaviour later in the season. An adaptive shift in male seasonal reproductive investment may be constrained because males use breeding females as cues for sexual behaviour. If this is so, we expect a general pattern across climatic regimes whereby female reproductive investment evolves first, with responses in males lagging behind.     

Initiation of the breeding season in ewe lambs and goat kids with melatonin implants

Sixty-eight female and 4 male lambs of the Chios breed born in autumn (September–October) and 48 female and 4 male kids of the Damascus breed also born in autumn (November), were used to evaluate the effect of melatonin implants (Regulin^®) on the initiation of the breeding season. For each species and sex, half of the animals were either left untreated to serve as controls (C) or received ear implants (females one and males two implants) of melatonin (MEL) in May (spring). Each C and MEL group of females was kept separately and away from males for 5 weeks after implantation and then, C and MEL males were joined with the respective groups for 45 (sheep) and 35 days (goats). Animals not conceiving during the early June–July breeding season, were mated again in September (natural breeding season). The reproductive performance and the 60-day milk yield of animals giving birth during the two seasons were compared. Early in the season (mid November–December), significantly more MEL than C ewes and goats gave birth (75.5% versus 23.5%, p < 0.01; 70.8% versus 37.5%, p < 0.05, respectively). For both species, most of the other characteristics examined were significantly affected by season, but not by treatment. Females giving birth early in the season were approximately 80 days younger at parturition and recorded a lower body weight at mating than those giving birth in the natural February–March season. In the first mating period, the animals mated and conceived had similar body weights to those not mated. The natural, compared with the early June–July mating season, was associated in sheep with higher litter weight at birth (7.0 kg versus 5.8 kg, p < 0.05) and in sheep and goats with a higher number (p < 0.05) of offspring born live (1.78 versus 1.38; 2.0 versus 1.61, respectively) and weaned (1.74 versus 1.36; 1.82 versus 1.35, respectively). Total litter size at birth and 60-day milk yield after weaning, were not significantly different between the two seasons. It could be concluded that melatonin implants administered during the last month of spring in autumn-born female Chios sheep and Damascus goats, advanced the initiation of their breeding season by about 80 days when joined with young males also treated with melatonin implants. The overall reproductive performance and milk yield of animals breeding early, was satisfactory and comparable to that of animals breeding in the natural breeding season.     

Individual differences in aggressiveness of female hamsters: response to intact and castrated males and to females

The aggressive behaviour of female hamsters was studied while they were housed in large enclosures with males and in brief tests with males or females. Some females are not aggressive with any male, whereas others are very aggressive toward all males in both testing conditions. Females that are not aggressive toward intact males may be very aggressive toward castrated males or females. When the animals are housed together for long periods of time, males dominate only if they are much heavier. Male dominance takes a relatively long time to establish and often there is an equivocal period characterized by reversals of dominance. Female dominance is rapidly established. Unless the male is much heavier, the female determines the presence or absence of agonistic behaviour.     

Reproductive strategies and the influence of date of birth on growth and sexual development of an aseasonally-breeding ungulate: Reeves' muntjac (Muntiacus reevesi)

Unlike most species of deer, Reeves' muntjac does not have a seasonal reproductive cycle. Births are equally distributed throughout the year and, irrespective of season of birth, females start breeding when they have reached a minimum body weight of about 10 kg. A significantly smaller proportion of females is born in the autumn/winter than the spring/summer, but there is no effect of maternal age or condition on foetal sex ratios for first or subsequent pregnancies. Whilst aseasonal breeding increases female productivity, males are only able to hold territories encompassing the ranges of a number of does for a relatively short period of time. It is argued that for muntjac there is much less inter-sexual variation in lifetime reproductive success than for seasonally-breeding polygynous cervids. Hence, although muntjac are sexually dimorphic and polygynous, females do not invest preferentially in male offspring.
Young males can be fertile from 36 weeks of age, when their first antlers are still in velvet. Whilst season of birth has no effect on the rate of sexual development in females, it does for males, with autumn-born male fawns attaining sexual maturity earliest. This accelerated development occurs in the period following independence from the mother, and there is no evidence of a maternal cost in producing male fawns. Since the canine tusks are the main weapons for intra-sexual conflict, there could be clear gains in terms of reproductive success for males that attain sexual maturity, adult weight and adult tusk size quickly, even though their first antlers are much smaller than those of older bucks. The hypothesis is presented that, for muntjac, the majority of a male's lifetime reproductive success is achieved relatively early in life when the canine tusks are in pristine condition, and that bucks lose their territories, and associated access to oestrous does, once the canines are broken.