Influence of plane of winter nutrition on plasma concentrations of prolactin and testosterone and their association with voluntary food intake in red deer stags (Cervus elaphus)

Blood samples were taken at weekly or 3-weekly intervals from 12 red deer stags, from 3 weeks of age until 3 years of age. Six of the stags were fed to appetite throughout the study and six were maintained on 70% as much during each winter and to appetite each summer. All plasma samples were analysed for testosterone and prolactin. Food intake was measured daily. Annual rhythms of testosterone, prolactin and food intake were evident. Peaks of prolactin, on an increasing photoperiod tended to occur slightly before peaks of food intake and troughs of food intake on a decreasing photoperiod occurred at or slightly after peaks of testosterone. The reduced plane of winter nutrition tended to delay rather than alter the amplitude of the annual cycles of testosterone, prolactin or food intake. Several of the unrestricted stags showed biannual rhythms of testosterone, prolactin and food intake accompanied by additional cycles of coat development and antler growth. These additional cycles tended to be 6 months out of rhythm with normal cycles. It is considered that this is due to the stags being stimulated by a particular day-length irrespective of whether it is decreasing or increasing and that plane of nutrition may influence the control of phase blockers.     

Photoperiod requirements for puberty differ from those for the onset of the adult breeding season in female sheep

Reproductive responses to photoperiod were directly compared in mature ewes and in their spring-born twin female lambs. All females were ovariectomized and treated with oestradiol implants before transfer into artificial photoperiod; serum LH concentrations and pulsatile LH patterns provided an index of neuroendocrine reproductive activity. Mothers were transferred from natural photoperiod to artificial long days (16 h light:8 h dark) at the summer solstice so that no decrease in photoperiod would be experienced. These ewes began reproductive activity synchronously at the expected time in the autumn. One of each pair of twin lambs was treated exactly as the mothers; to determine the normal timing of puberty the remaining twin was maintained in a photoperiod simulating the natural decrease in daylength. In all 6 control lambs experiencing the simulated natural photoperiod, reproductive activity occurred synchronously at 28 +/- 1 weeks of age (2 October +/- 7 days). However, in their twin sisters which did not experience a decrease in photoperiod, only 2 of 6 lambs had begun reproductive activity by the end of the experiment at 52 weeks of age (March), and these were both delayed relative to their twin control lambs exposed to decreasing daylength. Therefore, a decrease in photoperiod is necessary for the normal timing of puberty in the spring-born, female sheep, whereas seasonally anoestrous, mature sheep can enter the breeding season at a normal time in the absence of decreasing photoperiod. We suggest that the requirement for a decreasing photoperiod by the spring-born lamb reflects its limited photoperiodic history as compared to the adult.     

Rhythms of reproduction,metabolism and coat growth in deer: a model for all non-domesticated seasonal ungulates?

Seasonally-breeding deer living in cool temperate environments exhibit pronounced seasonal rhythms of voluntary food intake, growth rate and fattening and the growth and development of the coat. All of these rhythms are considered to be entrained by photoperiod and melatonin, although so far this has only been demonstrated to be the case in one species, the red deer. This paper reviewed current data on seasonal rhythms in several species of deer. A comparison of two species with different breeding seasons, the red deer and the Pere David's deer, indicated that seasonal reproduction, appetite and coat growth rhythms are linked and may be controlled by a single circannual rhythm generator. All of these seasonal rhythms should be considered as adaptive for species living in cool temperate environments with marked fluctuations in food supply and climatic conditions.     

Activity patterns and resource use by sheep and red deer grazing across a grass/heather boundary

Diet selection and activities of sheep and red deer grazing alone and together across a grass (primarily Lolium perenne/Agrostis capillaris ) and heather ( Calluna vulgaris ) boundary, between July and October, are described.
Throughout the experiment, both animal species had a much higher proportion of grass than heather in their diet. Time spent grazing on each type of vegetation was similarly greater on grass, and the bite rates of the animals were also more rapid on the grass than on the heather.
Red deer spent more time grazing per 24 hours than did sheep, with sheep grazing less during darkness hours than red deer. Total grazing time of both species did not decrease with decreasing daylength from July to October.
There were no consistent differences in the diurnal patterns of activity of sheep and red deer, with both species showing 2-3 large grazing peaks per day.
There were no consistent effects of mixing sheep and deer on their grazing behaviour and diet composition.     

Seasonality of reproduction in sheep and its control by photoperiod

Seasonality of the reproductive cycle in sheep is a general phenomenon for mid-latitude breeds. The proximal part (breeding season) and also partially distal part (end of gestation and beginning of lactation) of this cycle is controlled by photoperiod, whatever the form of light regimens. Data are presented which indicate that male and female do not necessarily have the same photoperiodic sensitivity. Gonadal stimulation in the ram starts 1.5-2 months earlier than in the ewe under annual variations. Photoperiod controls the reproductive cycle by the intermediary of the hypothalamo-pituitary axis. There are both a steroid-independent and a steroid-dependent effect of light, depending on both decreasing and increasing daylength in mid-latitudes. Data are also presented which support Bunning's hypothesis on photoperiodic time measurement in mammals. Sheep measure photoperiodic time by using a circadian rhythm of photosensitivity. Daylength is not measured by the total duration of exposure to light but by the illumination of two special set points during the day, one of them entraining the circadian rhythm of photosensitivity and the other inducing or not inducing a physiological response if it is coincident, or not coincident, with photoinducible phase of that rhythm. A photoinducible phase has been found for prolactin secretion, and perhaps also for LH secretion. Melatonin secretion is used by sheep for measuring daylength. However, that secretion disappears during two set points during the day, thus raising the possibility of using alternatively melatonin and light pulse for controlling the reproductive cycle in sheep.     

Control of annual endocrine rhythms in the edible dormouse: nonprimary effect of photoperiod.

We investigated the extent to which photoperiodic fluctuations synchronize annual thyroid and gonadal rhythmicity in edible dormice. The effects of different daylength manipulations (LD 4:20, LD 6:18, LD 18:6, LD 20:4) were examined during the two critical ascending and regressive phases of the annual plasma testosterone and thyroxine cycles that correspond to naturally increasing or decreasing photoperiod variations. The data failed to demonstrate any essential photoperiodic contribution to control systems that generate these two annual biological rhythms in dormice.     

The influence of birth date on the development of seasonal cycles in red deer hinds (Cervus elaphus)

The relative importance of age and photoperiod in the timing of seasonal cycles was studied in red deer during the first 18 months of life. Seasonal cycles of appetite, live-weight change, coat growth, oestrus and metabolic rate were examined in two groups of seven hind calves born in early May (group E) or nine weeks later in July (group L).
There was no difference between groups in the timing or amplitude of the seasonal voluntary food intake (VFI) cycle, or in the timing of seasonal changes in rate of live-weight gain, although mean live-weight gain was significantly greater for group L than group E during the first winter of life.
Four animals in group L retained their calf coats throughout the first winter. In group L, the first summer coat fibres appeared approximately four weeks earlier, and mean fibre length for the subsequent winter coat was longer in group L than in group E. There was no difference between groups in the timing or amplitude of the seasonal cycle of plasma prolactin concentration, or in the incidence and mean time of onset of puberty.
There was no difference between groups in mean energy expenditure per unit of live weight at any time of the year. Mean daily energy expenditure was significantly higher in summer than in winter, but values measured when the animals were resting and corrected for differences in VFI were similar throughout the year.
It is concluded that the seasonal cycles of appetite, live-weight change and oestrus in young red deer hinds are not modified by age, although the normal cycle of coat replacement can be disturbed in animals born towards the end of the calving season.     

Short photoperiod inhibits winter growth of subtropical grasses

Grass development is influenced by length of photoperiod, but no direct measurements under natural conditions exist on mass accumulation in response to photoperiod by subtropical grass species. Grasslands of the subtropics are a major resource, but their growth is inhibited substantially during the short-photoperiod months. This research was designed to examine the consequences on grass production under field conditions when the limitation of short photoperiod is artificially removed. Lights, which extended the daylength to 15 h, were placed over plots of four subtropical forage grasses representing three species (Paspalum notatum Flugge; Cynodon dactylon L.; Cynodon nlemfuensis Vanderyst) to measure their mass accumulation in response to extended photoperiod in a 2-year experiment. Forage yields in all grasses at 5-week harvests during the time of shortest daylength were increased up to 6.2-fold more than the yield under the natural daylength. For the 4.5-month period of shortest daylength in each year, forage yields were increased for all grasses with one grass having a yield increase of 3.6-fold under the extended photoperiod as compared to natural daylength. These results demonstrated that selection of grasses that are insensitive to photoperiod could substantially increase forage yield of subtropical grasslands to benefit animal production and enhance carbon sequestration.     

Circadian rhythm of sex pheromone-releasing behaviour in females of the dermestid beetle,Trogoderma glabrum: Regulation by photoperiod

The effects on sex pheromone-releasing, or calling behaviour, of diel photoperiods of varying daylength, of light cycle phase shifts, and of continuous illumination were investigated in Trogoderma glabrum females. On light régimes with 8 to 20 hr daylengths, calling maxima tended to centre close to photophase midpoints. Although influencing the time of day at which calling occurred, daylength had little effect on the amount of activity or the length of the calling period. When 16 : 8 LD light cycles were advanced or delayed by 4 hr, the time of day at which calling peaks were observed shifted within 2 to 4 cycles so that a constant phase relationship with photoperiod was maintained. Daily calling peaks were evident in groups of females exposed to between 1 and 5 days of continuous illumination, but mean calling time occurred earlier in the day as light exposures were lengthened. It was concluded that a circadian rhythm of calling behaviour exists in T. glabrum females. and that the rhythm can be entrained to 24 hr periodicity by photoperiod.     

Effects of photoperiod and low temperature on diapause termination in the yellow-spotted longicorn beetle (Psacothea hilaris)

Abstract. .The effects of photoperiod and low temperature on diapause termination in the yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe) (Coleoptera: Cerambycidae), were examined using a population from Ino, Japan. Diapausing insects obtained by rearing larvae under short daylength (12 or 13 h) at 25^oC were subjected to various treatments. When the photoperiod was changed at the same temperature, diapausing larvae showed a long-day response with a critical daylength between 13.5 and 14h. The diapause was terminated and consequently pupation occurred if the daylength was longer than 13.5 h. Chilling the diapausing larvae at 10^oC for 30 or more days also terminated diapause in most larvae irrespective of the photoperiods during and after chilling treatment. In contrast, the post-chilling photoperiod had a critical effect on development of diapausing larvae chilled for only 15 days.     

Experimental infection of some North American wild ruminants and domestic sheep with Mycobacterium paratuberculosis: clinical and bacteriological findings

Mycobacterium paratuberculosis originally isolated from bighorn sheep (Ovis canadensis) with spontaneous paratuberculosis was used to orally inoculate Rocky Mountain elk (Cervus elaphus nelsoni) calves, mule deer (Odocoileus hemionus) fawns, white-tailed deer (Odocoileus virginianus) fawns, bighorn X mouflon (Ovis musimon) hybrid lambs, and domestic lambs. All experimentally exposed animals became infected. During the first year of infection, hybrid and domestic sheep were able to control the infection but infection was progressive in elk and deer. Clinical paratuberculosis occurred only in mule deer.     

Aspects of the induction of diapause in a laboratory strain of the mite Tetranychus urticae

Some diapause characteristics were studied in a strain of the spider mite. Tetranychus urticae. which had been reared on bean plants in the laboratory for over 15 yr. The diapause induction response curve was of the long-day type, showing a sharply defined critical daylength of 13 hr 50 min. In constant darkness no diapause induction occurred, but with a photoperiod of 1L:23D diapause incidence was already complete. A thermoperiod with a 5°C amplitude induced diapause in combination with a short-day photoperiod only when the low phase of the thermoperiod coincided with the scotophase. The same thermoperiod did not induce any diapause in constant darkness. The photoperiodic reaction of the laboratory strain used in these experiments appeared to remain constant over a very long period of time and to be independent of the diapause history of previous generations of mites.Although photoperiodic sensitivity was demonstrated during the whole postembryonic development, sensitivity was maximal at the end of the protonymphal instar and declined rapidly during the deutonymphal instar. Only 2 inductive cycles of 10L:14D were required to induce up to 62% diapause if the mites were kept in continuous darkness during the remainder of their development. Long days or continuous light could reverse the inductive effect of a sequence of short-day cycles previously applied to the mites.Light breaks of 1 hr duration applied at different times during the dark period of a 10L:14D photoperiod generated a sharp bimodal response curve with two discrete points of sensitivity to the light breaks at 10 hr after ‘dusk’ and 10 hr before ‘dawn’, thus showing a remarkable similarity with the results obtained in light break experiments with some species of insects.     

Development and seed number in indeterminate soybean as affected by timing and duration of exposure to long photoperiods after flowering

BACKGROUND AND AIMS: Long photoperiods from flowering to maturity have been found to delay reproductive development in soybean (Glycine max) and to increase the number of seeds per unit land area. This study was aimed to evaluate whether sensitivity to photoperiod after flowering (a) is quantitatively related to the length of exposure to long days and (b) persists throughout the whole pod-setting period. It was also evaluated whether seed number was related to changes in the duration of post-flowering phenophases. METHODS: Two field experiments were conducted with an indeterminate cultivar of soybean of maturity group V. In expt 1, photoperiods 2 h longer than natural daylength were applied during different numbers of days from the beginning pod stage (R3) onwards, while in expt 2 these photoperiod extensions were imposed during 9 consecutive days starting at different times between R3 and R6 (full seed) stages. KEY RESULTS: There was a quantitative response of development to the number of cycles with a long photoperiod. The exposure to long photoperiods from R3 to R5 (beginning of seed growth) increased the duration of R3-R6 regardless of the timing of exposure. The stages of development comprised in the R3-R6 phase were delayed by current as well as by previous exposure to long days. A positive relationship was found between seed number and the duration of R3-R6, irrespective of the timing and length of exposure to the long photoperiod. CONCLUSIONS: Sensitivity to photoperiod remained high during the reproductive period and was highly and positively coupled with the processes of generation of yield.     

Fighting behaviour in territorial male roe deer <Emphasis Type="Italic">Capreolus capreolus</Emphasis>: the effects of antler size and residence

In all areas where they have been studied, male roe deer are believed to have a territorial mating system, although few quantitative studies have been conducted and there remains considerable debate about the function of male roe deer territories. We observed 139 aggressive interactions between male roe deer in Storfosna Island (Norway) during one territorial season (March–August). We recognised seven rank levels of escalation according to the potential danger of the behaviour. On the basis of the number of escalation levels included in the interactions, the complexity of the fights was also scored. We recorded the presence of other individuals during the interaction, the age, the antler size, the territorial status and the residency status of the two contestants and tested how these variables affected escalation, complexity and outcome of the fights. Most of the interactions ended with low levels of escalation, and physical contact occurred only in fights between two territorial bucks. The escalation was also affected by the difference in antler size index (the bucks escalated more when the difference in antler size was smaller) and increased with an increasing number of female deer present during the interaction. The resident buck won in 81% of the fights. When it drew or lost, it was generally both inferior in age and antler size, and the duration and escalation of the interactions were higher. However, even when a fight was lost, no territory loss occurred. These results are consistent with the evolutionary game theory and the proposed low risk–low gain strategy of roe deer bucks.     

An epizootic of hemorrhagic disease in white-tailed deer in Missouri

As part of a white-tailed deer (Odocoileus virginianus) survival study in Missouri (USA) we were actively monitoring 97 radio-collared deer when 8 (8%) died. This mortality, which occurred from 20 August to 23 September 1996, consisted of five adult females, two yearling females and one yearling male. Based on the seasonality of this mortality and the isolation of epizootic hemorrhagic disease virus (EHDV) serotype 2 from one of these animals, we believe that these losses resulted from an epizootic of hemorrhagic disease. The remains of five unmarked deer that may have died from HD also were found on the study area during this same period. During the fall following this mortality, we tested serum from 96 deer taken by hunters in the immediate area. Fifteen (16%) were positive for EHDV or bluetongue virus (BTV) antibodies as determined by agar gel immunodiffusion tests. Serum neutralization test results indicated that previous infections were caused by EHDV virus serotype 2. Based on these data, and assuming that there was no prior exposure to EHDV serotype 2 in this population, the exposure rate for this epizootic was 24% of which 8% died. We noted hoof interruptions in only two of the 96 deer sampled. During this mortality event, the Missouri Department of Conservation received no reports of dead deer, and without the radio-monitored animals the event would have been undetected.     

Photoperiod gene control over partitioning between reproductive and vegetative growth

Summary The hypothesis tested was that lack of photoperiod gene activity allows inherent partitioning of photosynthate to continued growth of the earliest potential buds, flowers, pods, and seeds (the organs that give rise to the yield). Alternatively, and competitively, photoperiod gene activity causes the photosynthate to be partitioned predominantly toward continued growth of new vegetative organs plus later initiation of more reproductive (yield) organs. This hypothesis was tested by comparing an insensitive and a photoperiod-sensitive bean (Phaseolus vulgaris L.) cultivar and their F₁ with F₂ segregates of undetermined genotype. Randomly derived homozygous F₈ segregates were also compared. The F₈ generation included one photoperiod-insensitive and one photoperiod-sensitive genotype in a 1:1 ratio, which verified control by one photoperiod gene. Under long daylength (LD), in addition to early versus late flowering and maturity, the two genotypes expressed opposite levels of 23 other traits that would be changed by competitive partitioning of the photosynthate. In contrast, under short daylength (SD), both genotypes flowered and matured early, and both expressed the levels for all 25 traits that the photoperiod-insensitive genotype expressed in both SD and LD. The photoperiod gene interacted with daylength to control the levels of all three major physiological components of yield: the aerial biomass, harvest index, and days to maturity. Included among the other traits with levels altered by daylength-modulated photoperiod gene activity were: the number of branches, nodes, leaves and leaf area, the rate of yield accumulation, and sink activity.Department of Plant Breeding and Biometry paper no. 758     

Celestial moderation of tropical seabird behavior

Most animals, including birds, have cyclic life histories and numerous studies generally conducted on captive animals have shown that photoperiod is the main factor influencing this periodicity. Moon cycles can also affect periodic behavior of birds. Few studies have investigated the influence of these environmental cues in natural settings, and particularly in tropical areas where the change in photoperiod is slight and some bird species keep cyclic behaviors. Using miniaturized light sensors, we simultaneously investigated under natural conditions the influence of photoperiod and moon phases on the migration dates and at-sea activity of a tropical seabird species, the Barau's petrel, throughout its annual cycle. Firstly, we found that birds consistently started their pre- and post-breeding migrations at precise dates corresponding in both cases to a day-duration of 12.5 hours, suggesting a strong influence of the photoperiod in the regulation of migration behavior. We also found that mean population arrival dates to the colony changed from year to year and they were influenced by moon phases. Returns at their colonies occurred around the last full moon of the austral winter, suggesting that moon cycle is used by birds to synchronize their arrival. Secondly, variations of day-time activity were sinusoidal and correlated to seasonal changes of daylength. We thus hypothesize that the photoperiod could directly affect the behavior of the birds at sea. Night-time at-sea activity exhibited a clear cycle of 29.2 days, suggesting that nocturnal foraging was highly regulated by moon phase, particularly during the non-breeding season. To our knowledge, this is the first study to document a mixed regulation of the behavior of a wild bird by photoperiod and moon phases throughout its annual cycle.     

Reproductive ecology of female Oligocottus snyderi Greeley: a North American intertidal sculpin

The reproductive condition of female Oligocottus snyderi Greeley during 18+ months at Dillon Beach, California was examined. Seasonal differences in reproductive condition were identified by diameter measurements of ovarian follicles in pre-vitellogenic, cortical alveoli and vitellogenic stages of development which showed that female O. snyderi appeared capable of reproduction during 6 to 8 months of the year. Reproduction occurred from late autumn to early spring and it is likely that females reproduce more than once a year. Follicular development within the population was asynchronous; development within individual females was group synchronous. The percentage of females with vitellogenic follicles in each collection was negatively correlated with photoperiod and productivity and positively correlated with wave action. Conversely, the percentage of newly recruited individuals in each sample was positively correlated with photoperiod and productivity. It appears that reproduction and recruitment in O. snyderi are significantly influenced by annual productivity cycles; daylength changes may provide an anticipatory cue for the productivity cycle.     

Seasonal changes in moult,body mass and reproductive condition in siskins Carduelis spinus exposed to daylength regimes simulating different latitudes

Birds use change in daylength during the year to time events during their annual cycles. Individual Eurasian siskins Carduelis spinus can breed and winter in widely separated areas in different years. Birds at different latitudes will experience different changes in photoperiod. So how does latitude affect photoperiodic control? Our aim in this study was to find whether Siskins caught from the wild in Britain and exposed to different photoperiodic regimes, typical of widely separated latitudes, would differ in the subsequent timing and duration of their moults and associated processes. Siskins were caught in late February and early March, and initially kept outside on natural photoperiods. From the spring equinox (21 March), they were divided into three groups kept under photoperiodic regimes that simulated latitudes 40°, 55° and 70°N respectively. All three groups showed highly significant subsequent changes in body mass, fat scores and cloacal protuberance size. Moult of the primary feathers started during June – August (mean 9 July), and lasted 61–99 days (mean 75 days). Birds that started to moult late in the season had shorter moult durations. All individuals showed lower mass and fat levels during moult than before or after moult. Crucially, there were no significant differences in the timing of these events between the three photoperiodic groups. Apparently these birds did not use prevailing absolute photoperiod or the prevailing rate of change in photoperiod to time moult‐related seasonal events, but used instead some other feature of the annual photoperiod cycle or some form of interval timer linked to photoperiod.     

Extreme Seasonal Growth in Arctic Deer: Comparisons and Control Mechanisms

Arctic ungulates have the capacity to voluntarily restrict growthto times of the year of predictable high quality seasonal herbageabundance. The constraints of the arctic environment mean thatnot only must the timing of this growth seasonality be accuratebut growth rate must be maximised for a short period of time.It is known that daylength is a critical cue for the timingof seasonal rhythms of growth; a manipulated photoperiod of16L:8D stimulates out of season growth in reindeer during winterand this growth is associated with an increase in the plasmalevels of insulin like growth factor 1 (IGF 1). In contrasta manipulated photoperiod of 8L: 16D delays the spring risein growth. There is good evidence from a boreal cervid, thered deer, that the number of hours of day influences growthrate also and this is associated with IGF 1. Thus daylengthmay have a dual role in growth seasonality, timing and amplitude.Although the mechanisms underlying the timing of rhythms bydaylength are partly understood a putative role for daylengthin controlling the rate of an event is a new concept.