Variation in the timing of the reproductive season among breeds of sheep in relation to differences in photoperiodic synchronization of an endogenous rhythm.

Photoperiod may regulate seasonal reproduction either by providing the primary driving force for the reproductive transitions or by synchronizing an endogenous reproductive rhythm. This study evaluated whether breed differences in timing of the reproductive seasons of Finnish Landrace (Finn) and Galway ewes are due to differences in photoperiodic drive of the reproductive transitions or to differences in photoperiodic synchronization of the endogenous rhythm of reproductive activity. The importance of decreasing photoperiod after the summer solstice in determining the onset and duration of the breeding season was tested by housing ewes from the summer solstice in either a simulated natural photoperiod or a fixed summer-solstice photoperiod (18 h light:6 h dark; summer-solstice hold). Onset of the breeding season within each breed did not differ between these photoperiodic treatments, but Galway ewes began and ended their breeding season earlier than Finn ewes. The duration of the breeding season was shorter in Galway ewes on summer-solstice hold than on simulated natural photoperiod; duration did not differ between photoperiodic treatments in Finn ewes. The requirement for increasing photoperiod after the winter solstice for initiation of anoestrus was tested by exposing ewes from the winter solstice to either a simulated natural photoperiod or a winter-solstice hold photoperiod (8.5 h light:15.5 h dark). Onset of anoestrus within each breed did not differ between these photoperiodic treatments, but the time of this transition differed between breeds. These observations suggest that genetic differences in timing of the breeding season in Galway and Finn ewes do not reflect differences in the extent to which photoperiod drives the reproductive transitions, because neither breed requires shortening days to enter the breeding season or lengthening days to end it at appropriate times. These findings are consistent with the hypothesis that photoperiod synchronizes an endogenous rhythm of reproductive activity in both breeds and that genetic differences in timing of the breeding season reflect differences in photoperiodic synchronization of this rhythm.     

Evidence that the seasonally breeding Bennett's wallaby (Macropus rufogriseus rufogriseus) does not exhibit short-day photorefractoriness

Adult female Bennett's wallabies (N = 6) were maintained in artificial winter solstitial daylengths commencing 3 weeks before the winter solstice for 16 or 42 weeks. Such treatment effectively prevented the normal establishment of seasonal reproductive quiescence with animals continuing to exhibit reproductive cycles beyond the time of the normal termination of the breeding season. Animals maintained in natural photoperiods or simulated natural changes in daylength after the winter solstice all entered reproductive quiescence by early February. In the Bennett's wallaby, therefore, the breeding season does not terminate as a result of refractoriness to short daylengths. Our results indicate that the relatively small increases in photoperiod shortly after the winter solstice provide the environmental signal responsible for initiating the onset of seasonal reproductive quiescence which normally occurs 5-8 weeks after the solstice. These results contrast with the effect of fixed artificial summer solstitial daylengths on the onset of the breeding season in which breeding begins spontaneously at the normal time of year as a result of long-day photorefractoriness.     

Seasonal regulation of reproduction: altered role of melatonin under naturalistic conditions in hamsters

The seasonal reproductive cycle of photoperiodic rodents is conceptualized as a series of discrete melatonin-dependent neuroendocrine transitions. Least understood is the springtime restoration of responsiveness to winter-like melatonin signals (breaking of refractoriness) that enables animals to once again respond appropriately to winter photoperiods the following year. This has been posited to require many weeks of long days based on studies employing static photoperiods instead of the annual pattern of continually changing photoperiods under which these mechanisms evolved. Maintaining Siberian hamsters under simulated natural photoperiods, we demonstrate that winter refractoriness is broken within six weeks after the spring equinox. We then test whether a history of natural photoperiod exposure can eliminate the requirement for long-day melatonin signalling. Hamsters pinealectomized at the spring equinox and challenged 10 weeks later with winter melatonin infusions exhibited gonadal regression, indicating that refractoriness was broken. A photostimulatory effect on body weight is first observed in the last four weeks of winter. Thus, the seasonal transition to the summer photosensitive phenotype is triggered prior to the equinox without exposure to long days and is thereafter melatonin-independent. Distinctions between photoperiodic and circannual seasonal organization erode with the incorporation in the laboratory of ecologically relevant day length conditions.     

The role of refractorinesss to long daylength in the annual reproductive cycle of the female Bennett's Wallaby (Macropus rufogriseus rufogriseus)

The beginning of the breeding season of the female Bennett's wallaby occurs when seasonal quiescence terminates 1-2 months after the summer solstice. In this study, the role of photoperiod in terminating seasonal quiescence was examined. One week before the summer solstice, five non-lactating wallabies were transferred from natural to artificial summer solstice daylength for 5 months. The beginning of the breeding season in these animals as indicated by births, matings, and peripheral progesterone profiles was not different from that of five control animals maintained on natural photoperiod. The following year, three animals were transferred from natural to summer solstice daylength on February 25 and were held on the artificial photoperiod until September 30. Changes in plasma progesterone concentrations indicative of the beginning of the breeding season occurred on June 12-30 (range), which was significantly (P less than 0.01) advanced by 29 days when compared with six control animals. These results indicate that the decrease in daylength that occurs after the summer solstice is not required to induce the termination of seasonal quiescence at the beginning of the breeding season. Further, the beginning of the breeding season can be advanced by transferring animals to long daylength early in seasonal quiescence. Photorefractoriness to long daylengths may therefore be important in the initiation of the breeding season in this species. In further experiments, groups of six animals were transferred from natural to artificial summer solstice daylength on September 26 and December 9 and pouch young were removed 7 days after the transfer. In September, reactivation of the quiescent corpus luteum followed soon after removal of pouch young (RPY) indicating that exposure to long daylength had not induced a transition into seasonal quiescence. In December, RPY was not followed by reactivation of the quiescent corpus luteum indicating that animals were in seasonal quiescence. These results suggest that the female Bennett's wallaby may need to experience a period of shortening days after the summer solstice before exposure to long days can again initiate seasonal quiescence.     

Termination of the breeding season in the Suffolk ewe: involvement of an endogenous rhythm of reproduction

Two studies were performed to determine if the transition into anestrus in the Suffolk ewe results from the lack of a decrease in photoperiod, a signal suggested to be important in maintaining the breeding season, or from an obligatory turn-off that reflects the expression of an endogenous rhythm of reproduction. Shortly after the autumnal equinox, three groups of ovariectomized ewes bearing s.c. Silastic implants of estradiol were placed in different lighting environments. A control group was exposed to normal variations in day length until the winter solstice and held on that day length thereafter. The two experimental groups experienced either a continuously decreasing day length or a continuously increasing duration of elevated melatonin levels during each 24-h period until mid-March. Reproductive activity, assessed by circulating levels of luteinizing hormone (LH), was marginally extended (2-3 wk) in the experimental groups. In a second experiment, two groups of ovary-intact ewes were exposed to photoperiodic treatments shortly after the winter solstice. A control group was held on the winter solstice day length (10L:14D) until the end of the study in mid-March. The experimental group experienced a 3-h photoperiodic reduction to 7L:17D. It remained to determine if that abrupt decrease could maintain reproductive induction. Again, the cessation of reproductive induction was marginally delayed in the experimental group (2-3 wk). This marginal delay in the arrest of reproductive activity seen in both experiments indicates that the lack of decrease in day length around and after the winter solstice may play some role in timing the end of the breeding season. However, our inability to prevent or markedly delay the termination of reproductive activity leads to the conclusion that the primary reason for the transition into anestrus is an obligatory turn-off. This obligatory process may be the expression of an underlying endogenous rhythm of reproduction.     

Photoperiod: Its importance as an impeller of pineal and seasonal reproductive rhythms

A number of long day breeding rodents depend on seasonal changes in photoperiodic length to synchronize their breeding seasons with the appropriate time of the year. These relationships are particularly conspicuous in the Syrian hamster where day length is vitally important in determining periods of sexual activity and inactivity. The organ in the body whose activity is most closely attuned to the photoperiodic environment is the pineal gland. During periods of darkness the biochemical and secretory activity of the pineal is enhanced with the resultant production of antigonadotrophic principles which are strongly suppressive to reproductive physiology. In this manner, decreasing day lengths of the fall are involved with suppressing sexual capability in male and female hamsters. Throughout the winter months darkness (because of the shorter day lengths and the fact that hamsters remain underground in lightless burrows) holds the gonads in an atrophic condition and thereby prevents hamsters from breeding. As spring approaches the neuroendocrine reproductive axis becomes refractory to the inhibitory effects of darkness and the pineal gland and, as a consequence, the gonads recrudesce allowing the animals to successfully reproduce. The long days of the spring and summer serve to interrupt the refractory period so that when winter approaches shortening day lengths will again, by way of the pineal gland, induce gonadalinvolution. In this scheme both light and darkness are critically important in synchronizing the phases of the annual reproductive cycle of the hamster with the appropriate season of the year. Melatonin may be the pineal hormone which mediates the effects of darkness on reproductive physiology.Presented at the Eighth International Congress of Biometeorology, 9–14 September 1979, Shefayim, Israel.     

Amplitude of the plasma melatonin nycthemeral rhythms is not associated with the dates of onset and offset of the seasonal ovulatory activity in the Ile-de-France ewe

We investigated if absolute (nocturnal) or relative (nocturnal/diurnal ratio) plasma melatonin concentrations were associated with the seasonal ovulatory activity in Ile-de-France ewes. Ninety-six and 121 ewes in two different groups of the same flock were used to determine the potential existence of a relationship between melatonin concentrations at the summer and winter solstices, and the dates of onset and offset of the ovulatory activity, respectively. The dates of the first and last ovulations were estimated by assaying progesterone in plasma samples taken once weekly. Mean +/- SEM (1) plasma melatonin concentrations at the summer solstice and at the winter solstice were 302.4 +/- 19.4 and 412.0 +/- 18.7 pg x mL(-1), respectively, (2) date of onset of the breeding season 29 Jul. +/- 1.6 days, (3) date of offset of the breeding season 24 Jan. +/- 2.2 days. In spite of a large variability in the different traits studied here, the analyses of correlation and regression indicated that neither the absolute nor relative melatonin concentrations were significantly related with the dates of onset or offset of ovulatory activity. Therefore, we concluded that absolute or relative plasma melatonin concentrations are not linked to the seasonal breeding activity, in Ile-de-France ewes.     

Nocturnal variation of prolactin secretion in the Mouflon (Ovis gmelini musimon) and domestic sheep (Ovis aries): seasonal changes

Seasonal changes in nocturnal prolactin secretion and their relationship with melatonin secretion were monitored in wild (Mouflon, Ovis gmelini musimon) and domesticated sheep (breed Manchega, Ovis aries). Two groups of eleven adult females each, were maintained outdoors under natural photoperiod. Plasma concentrations of prolactin and melatonin were determined during the summer and winter solstices and the autumn and spring equinoxes. Blood samples were collected every 3h during the night hours, and 1h before and after the onset of darkness and sunrise. Maximum mean plasma concentrations of prolactin during the dark-phase in Mouflons were observed in the summer solstice, (P<0.001) and in the summer solstice and spring equinox in Manchega ewes (P<0.001). Mean plasma concentrations of prolactin were higher in the wild species (P<0.001) during the summer solstice. In contrast, during the spring equinox, mean levels of prolactin were higher in Manchega ewes than in Mouflons (P<0.05). Plasma prolactin concentrations showed a nocturnal rhythm in both breeds, with seasonal variations (P<0.001). The increase in plasma melatonin levels during the first hour after sunset was accompanied to increasing concentrations of PRL 1h after the onset of darkness, only in the autumn and spring equinox for the Mouflon, and in the summer solstice and spring equinox for the Manchega ewes. In Mouflons, the fall of plasma PRL concentrations about the middle dark-phase in all the periods studied, coincided with high levels of melatonin. A similar relation was observed in Manchega ewes only in the winter solstice and spring equinox. The current study shows that the nocturnal rhythm of prolactin secretion exhibits seasonal variation; differences in the patterns of prolactin secretion between Mouflon and Manchega sheep are taken to represent the effects of genotype.     

Importance of photoperiodic signal quality to entrainment of the circannual reproductive rhythm of the ewe

An endogenous circannual rhythm drives the seasonal reproductive cycle of a broad spectrum of species. This rhythm is synchronized to the seasons (i.e., entrained) by photoperiod, which acts by regulating the circadian pattern of melatonin secretion from the pineal gland. Prior work has revealed that melatonin patterns secreted in spring/summer entrain the circannual rhythm of reproductive neuroendocrine activity in sheep, whereas secretions in winter do not. The goal of this study was to determine if inability of the winter-melatonin pattern to entrain the rhythm is due to the specific melatonin pattern secreted in winter or to the stage of the circannual rhythm at that time of year. Either a summer- or a winter-melatonin pattern was infused for 70 days into pinealectomized ewes, centered around the summer solstice, when an effective stimulus readily entrains the rhythm. The ewes were ovariectomized and treated with constant-release estradiol implants, and circannual cycles of reproductive neuroendocrine activity were monitored by serum LH concentrations. Only the summer-melatonin pattern entrained the circannual reproductive rhythm. The inability of the winter pattern to do so indicates that the mere presence of a circadian melatonin pattern, in itself, is insufficient for entrainment. Rather, the characteristics of the melatonin pattern, in particular a pattern that mimics the photoperiodic signals of summer, determines entrainment of the circannual rhythm of reproductive neuroendocrine activity in the ewe.     

Dynamic expressions of hypothalamic genes regulate seasonal breeding in a natural rodent population

Seasonal breeding is a universal reproductive strategy in many animals. Hypothalamic genes, especially type 2 and 3 iodothyronine deiodinases (Dio2/3), RFamide‐related peptide 3 (Rfrp‐3), kisspeptin (Kiss‐1) and gonadotropin‐releasing hormone (GnRH), are involved in a photoperiodic pathway that encodes seasonal signals from day length in many vertebrate species. However, the seasonal expression patterns of these genes in wild mammals are less studied. Here, we present a four‐year field investigation to reveal seasonal rhythm and age‐dependent reproductive activity in male Brandt's voles (Lasiopodomys brandtii) and to detect relationships among seasonal expression profiles of hypothalamic genes, testicular activity, age and annual day length. From breeding season (April) to nonbreeding season (October), adult male voles displayed a synchronous peak in gonadal activity with annual day length around summer solstice, which was jointly caused by age structure shifts and age‐dependent gonadal development patterns. Overwintered males maintained reproductive activity until late in the breeding season, whereas most newborn males terminated gonadal development completely, except for a minority of males born early in spring. Consistently, the synchronous and opposite expression profiles of Dio2/3 suggest their central function to decode photoperiodic signals and to predict the onset of the nonbreeding season. Moreover, changes in Dio2/3 signals may guide the actions of Kiss‐1 and Rfrp‐3 to regulate the age‐dependent divergence of reproductive strategy in wild Brandt's vole. Our results provide evidence on how hypothalamic photoperiod genes regulate seasonal breeding in a natural rodent population.     

The coincidence of light and melatonin with a specific phase of the circadian pacemaker is important for the timing of seasonal breeding in the ewe

The timing of reproductive activity in seasonal breeding sheep relies on daily photoperiodic signals being relayed to provide information on the time of year. Although light and melatonin are involved, the exact mechanism is not understood. In this experiment, three groups of 6 Romney Marsh ewes, a highly seasonal breed, were provided with 8 weeks of short nights (9.6-9.8 h, by artificially advancing dawn) around the winter solstice, near the end of their natural breeding season. One group of animals was infused to a physiological level with melatonin for 5 h during the afternoon prior to the onset of dark, while a second group was identically infused but for 5 h from the time of lights on. A third group received the short-night treatment only. Following the short-night treatment, all groups were exposed to long nights (> 14 h, by delaying dawn) until the summer solstice. Ovarian activity, assessed by progesterone monitoring twice weekly, showed that the noninfused and the morning-infused groups displayed renewed reproductive activity in response to the short-night/long-night treatment. There was no renewed ovarian activity in the afternoon-infused group, indicating that the time of day that melatonin is present, rather than the duration of melatonin exposure, is an important signal in the control of reproductive timing. Measurements of a marker of the endogenous circadian pacemaker, by melatonin measurements under acutely extended darkness, revealed that the short-night treatments phase advanced the onset of the pacemaker in all groups such that the afternoon phase of the pacemaker was coincident with light. The results provide strong support for the model that proposes that an afternoon-located sensitive phase of the pacemaker is responsible for the relay of photoperiodic signals in the timing control of seasonal breeding. The model proposes that the reproductive axis be primed during short nights when the sensitive phase is coincident with light in the afternoon so ovarian activity can be induced when the sensitive phase is located within the longer nights of autumn and coincident with endogenous melatonin.     

The interaction between photoperiod and nutrition and its effects on seasonal rhythms of reproduction in the ewe

In sheep, the seasonal patterns of reproductive activity are driven primarily by the annual photoperiodic cycle, but can also respond to other environmental factors, such as nutrition, yet little is known about the mechanisms underlying this interaction. This study was designed to define the interaction between photoperiodic and nutritional cues on seasonal patterns of ovarian activity, and to determine if there is a central interaction between these cues. Groups of Ile-de-France ewes were maintained in two nutritional states (restricted and well fed) under a simulated annual photoperiod of 8-16 h of light per day over two breeding seasons. At the end of the first breeding season, half of the animals of each group were ovariectomized (OVX) and fitted subcutaneously with estradiol implants. Low nutritional status shortened the season of ovarian activity, determined from the pattern of progesterone concentrations, by modifying the timing of seasonal transitions between periods of ovarian activity and anestrus. The same results were observed for the seasonal rhythm of neuroendocrine activity, assessed in the OVX ewes, from the pattern of luteinizing hormone concentrations. These results were then confirmed for neuroendocrine activity induced by a photoperiodic treatment. We conclude that nutrition centrally modulates the interpretation of photoperiod to affect seasonal reproductive transitions. The mechanisms of this interaction are discussed in the paper.     

Social cues can play a role in timing onset of the breeding season of the ewe

Female Suffolk sheep were pinealectomized around the vernal equinox to eliminate the major environmental input to the reproductive system (photoperiod) and then either isolated from, or maintained with, pineal-intact gonad-intact sheep. The ewes were ovariectomized and treated with constant-release oestradiol implants and reproductive state was monitored by measuring serum LH concentrations. Pinealectomized ewes that were isolated from the normal flock showed a 2 1/2-month delay in onset of the seasonal rise in LH values compared with that of pineal-intact controls (18 November vs 5 September). On the other hand, pinealectomized ewes that were maintained with the flock showed an onset of the seasonal rise in LH that was not delayed. These results suggest a timekeeping role for social cues for timing onset of the breeding season in an animal that normally relies on photoperiodic signals for temporal regulation of the seasonal reproductive cycle.     

Seasonal LH profile in ovariectomized cattle

Two experiments were conducted to examine seasonal changes in circulating LH concentrations in ovariectomized heifers. In experiment 1, four Holstein heifers were ovariectomized in April 1977 during middiestrus. Blood samples were collected daily for 30 days surrounding each equinox and solstice for one year to examine changes in plasma LH levels at the time of seasonal photoperiod changes. The LH concentrations were highest during the winter solstice period and lowest during the summer solstice period. In addition, samples taken at two-week intervals indicated a distinct LH profile with maximal LH concentrations during November-April and minimal concentrations during May-October. In experiment 2, eight Holstein heifers were ovariectomized in June-July, 1979 and given an estradiol or a control implant in October. A distinct LH profile for the interval extending from January, 1980 to February, 1981 was found in the heifers that were not treated with estradiol. Concentrations were maximal during December-April and minimal during May-November. The LH profile followed a similar pattern in the estradiol-treated heifers; however, the overall profile was at a higher level. These data indicate that underlying seasonal reproductive mechanisms are present in cattle even though the species ovulates and breeds throughout the year.     

The onset of seasonal quiescence in the female Bennett's wallaby (Macropus rufogriseus rufogriseus)

The breeding season of the non-lactating Bennett's wallaby terminates when animals enter the state of seasonal quiescence. To examine this transition, pouch young were removed from females at intervals which were 3, 4 or 8 weeks (6, 11 and 8 animals respectively) after the winter solstice. Within 48 days, 3, 1 and 1 females gave birth respectively, indicating that these animals were not in seasonal quiescence when pouch young were removed. Those animals which did not give birth were either in seasonal quiescence or had undergone a non-pregnant cycle. To differentiate between the 2 possibilities, techniques which would ensure the detection of pregnant and non-pregnant cycles were assessed in 8 females during the breeding season. As has been previously reported for the wallaby, changes in peripheral progesterone concentrations and the vaginal smear occurred during pregnant and non-pregnant reproductive cycles. In addition, mating was detected by marking the male with a mixture of coloured crayon and paraffin wax. It was concluded that reproductive cycles in female wallabies could be monitored by collecting blood samples 2 times each week for progesterone determination and daily examination of females for mating marks. These techniques were then used to study the onset of seasonal quiescence in 9 females. All animals continued to show reproductive cycles after the winter solstice and it was not until 10 weeks after the winter solstice that all animals were in seasonal quiescence. This represents an increase in the duration of the breeding season over that previously reported for this species.     

Refractoriness to short and long days determines the end and onset of the breeding season in subtropical goats

The objective was to determine whether refractoriness to short and long days were involved in the end and onset of the breeding season, respectively, in goats adapted to subtropical latitudes. Ovariectomized does given a subcutaneous implant constantly releasing estradiol-l7 β (OVX+E) were used in two experiments. Plasma LH concentrations were determined twice weekly. In Experiment 1, the control group remained in an open-shed pen (natural day length and ambient temperature). Two experimental groups were placed in light-proof buildings (with natural temperature variations). One group was exposed to natural simulated increasing days (winter to spring), whereas the other was exposed to a winter solstice photoperiod (10 h of light) from December 21 to April 28. In Experiment 2, the control group remained under natural day length and ambient temperature. One experimental group was exposed to natural simulated decreasing days (summer to autumn), whereas the other group was exposed to a summer solstice photoperiod (14 h of light) from June 21 to October 20. In Experiment 1, the breeding season was not prolonged in does maintained in the winter solstice day length. Mean dates of decrease in LH secretion (end of the breeding season) did not differ significantly between does exposed to natural (February 3 ± 5 d) or natural simulated photoperiod (January 26 ± 14 d) and those exposed to constant short days of winter solstice (February 4 ± 10 d). In Experiment 2, the onset of the breeding season was not delayed in does maintained in the summer solstice day length. Mean dates of increase in LH secretion (onset of the breeding season) did not differ significantly between does exposed to natural (September 7 ± 8 d) or natural simulated photoperiod (September 18 ± 10 d) and those exposed to constant long days photoperiod of summer solstice (September 24 ± 4 d). In goats adapted to a subtropical environment, we concluded that: 1) the end of breeding season was due to refractoriness to short days, and not the inhibitory effect of increasing day length; and 2) the onset of the breeding season was due to refractoriness to long days, and not a stimulatory effect of decreasing day length.     

Refractoriness to inhibitory day lengths initiates the breeding season of the Suffolk ewe

Recent evidence indicates that the breeding season of the Suffolk ewe ends because of loss of response to a day length that was previously inductive. This condition of photorefractoriness could potentially also initiate reproduction, as is the case in several long-day breeding rodents. In this study we determined if ewes enter their breeding season because they become refractory to the long ambient photoperiods of late summer. On the summer solstice, 3 groups of ovariectomized ewes (n = 6) bearing s.c. Silastic implants of estradiol (OVX + E) were placed in different day length treatments: 1) natural photoperiod; 2) artificial photoperiod, stimulating natural day lengths; or 3) artificial photoperiod equivalent to that of the summer solstice (16.25L). Entry into the breeding season is associated with a striking (greater than 30-fold) increase in circulating levels of luteinizing hormone (LH). Timing of the onset of the breeding season was not delayed in ewes maintained on the summer solstice photoperiod; LH levels rose simultaneously in all groups. We conclude that ewes normally begin breeding not because they are actively driven to do so by decreasing or short days, but because they become refractory to prevailing long days. Because the pattern of circulating melatonin, which is known to transduce the photoperiodic message, remained entirely appropriate to day length, we believe that the mechanism responsible for photorefractoriness resides in the postpineal processing of the melatonin signal.     

Influence of season and estradiol on secretion of luteinizing hormone in ovariectomized cows

The hypotheses that secretion of luteinizing hormone (LH) varies with season and that estradiol may modulate the seasonal fluctuation in secretion of LH in cows were investigated. Seven mature cows were ovariectomized approximately 30 days before initiation of the experiment. Three of the ovariectomized cows (OVX-E2) were administered a subcutaneous estradiol implant that provided low circulating levels of 17 beta-estradiol. The remaining 4 cows (OVX) were not implanted. From December 21, 1982, to September 20, 1984, blood samples were collected sequentially (at 10-min intervals for 6 h) at each summer and winter solstice, and each spring and autumn equinox. In addition, from March 17, 1983, to March 17, 1984, sequential samples were collected midway between each solstice and equinox. Concentration of LH was measured in all samples, and concentration of estradiol was measured in pools of samples. An annual cycle in mean serum concentration of LH and amplitude of LH pulses was detected in both groups of cows. The seasonal pattern did not differ in the two treatment groups. Serum concentration of LH and amplitude of LH pulses were highest around the spring equinox, decreased gradually to the autumn equinox, and then increased and peaked again during the following spring equinox. Frequency of LH pulses and concentration of estradiol in serum did not vary with season. Circulating concentrations of LH and amplitude of pulses tended to be higher in OVX-E2 than OVX cows throughout the experimental period. Frequency of pulses of LH was lower in OVX-E2 than OVX cows throughout the experiment. Concentrations of estradiol were higher in the implanted cows.(ABSTRACT TRUNCATED AT 250 WORDS)     

Seasonal variations in daily rhythms of activity in athletic horses

Circadian rhythms reflect extensive programming of biological activity that meets and exploits the challenges and opportunities offered by the periodic nature of the environment. In the present investigation, we recorded the total activity of athletic horses kept at four different times of the year (vernal equinox, summer solstice, autumn equinox and winter solstice), to evaluate the presence of seasonal variations of daily activity rhythms. Athletic Thoroughbred horses were kept in individual boxes with paddock. Digitally integrated measure of total activity of each mare was continuously recorded by actigraphy-based data loggers. Horse total activities were not evenly distributed over the day, but they were mainly diurnal during the year. Daily activity rhythms showed clear seasonal variations, with the highest daily amount of activity during the vernal equinox and the lowest during the winter solstice. Interestingly, the amount of activity during either photophase or scotophase changed significantly throughout the year. Circadian analysis of horse activities showed that the acrophase, the estimated time at which the peak of the rhythm occurs, did not change during the year, it always occurred in the middle of the photoperiod. Analysing the time structure of long-term and continuously measured activity and feeding could be a useful method to critically evaluate athletic horse management systems in which spontaneous locomotor activity and feeding are severely limited. Circadian rhythms are present in several elements of sensory motor and psychomotor functions and these would be taken into consideration to plan the training schedules and competitions in athletic horses.     

The effects of season,daylight saving and time of sunrise on serum cortisol in a large population

Cortisol is critical for maintenance of health and homeostasis and factors affecting cortisol levels are of clinical importance. There is conflicting information about the effects of season on morning cortisol and little information on the effects of sunlight on population cortisol assessment. The aim of this study was to assess whether changes in median serum cortisol occurred in a population in conjunction with changing seasons, daylight saving time (DST) or time of sunrise. We analysed serum cortisol results (n?=?27 569) from a single large laboratory over a 13-year period. Subjects with confounding medications or medical conditions were excluded and data analysed in 15-minute intervals. We assessed the influence of traditional seasons, seasons determined by equinox/solstice, DST and time of sunrise on median cortisol. The median time of cortisol collection did not vary significantly between seasons. Using traditional seasons, median cortisol was lowest in summer (386?nmol/L) and spring (384?nmol/L) with higher cortisol in autumn (406?nmol/L) and winter (414?nmol/L). Median cortisol was lowest in the summer solstice quarter with significant comparative increases in the spring equinox quarter (3.1%), the autumn equinox quarter (4.5%) and the winter solstice quarter (8.6%). When cortisol was modelled against time, with adjustment for actual sunrise time on day of collection, for each hour delay in sunrise there was a 4.8% increase in median cortisol (95% CI: 3.9–5.7%). In modelling to explain the variation in cortisol over the morning, sunrise time was better than season in explaining seasonal effects. A subtle cyclic pattern in median cortisol also occurred throughout the months of the year. A 3-year trial of DST allowed comparison of cortisol in DST and non DST periods, when clock time differed by one hour. There was modest evidence of a difference in acrophase between DST and non DST cortisol (p?=?0.038), with DST peak cortisol estimated to occur 58?minutes later than non-DST peak. In summary, we found that time of sunrise and time of cortisol collection were the most important factors influencing median cortisol. For each hour later that the sun rose there was an almost 5% increase in median cortisol. There was significant seasonal variability with lowest cortisol noted in summer coinciding with the earliest sunrise time. This is an important finding which is consistent with the understanding that light is the major zeitgeber in entrainment of the human circadian cortisol rhythm. Our data suggest this rhythm is resistant to the arbitrary changes in clock time with daylight saving.