Photoperiodic time measurement in spider mites

Summary A model has been developed which describes in mathematical terms the incidence of diapause in the spider miteTetranychus urticae under different photoperiodic regimens. The model has been derived from Beck's (1974a, b, 1975, 1976a) Dual System Theory of photoperiodic time measurement, by means of a number of essential alterations and modifications. The spider mite's model is composed of two hour-glass timers, one of which starts at lights-off and measures the length of the night, whereas the other is initiated both by the onset of the first hour-glass timer and by lights-on. This second hour-glass defines the time at which the first hour-glass is read off, the state of the first hour-glass at this particular time being decisive for the developmental alternative (diapause or nondiapause) to be determined. The model may be classified as a form of internal coincidence according to the terminology of Pittendrigh (1972), since it is based on the interaction of two internal systems rather than on the coincidence of light with a particular light-sensitive phase of the timing mechanism, as in the case of external coincidence (cf. Saunders, 1978). Good agreement is attained between diapause incidences predicted by this model and incidences observed in spider mites, both in the diapause induction response curve and in asymmetrical skeleton photoperiods.The authors are grateful to Dr. S.D. Beck for the program printout of the Dual System Theory put at their disposal. It is also a pleasure to thank Miss C. van Ruth and Mr. G. van de Berg for technical assistance, and Mr. H. Bos for drawing the figures. The investigations were supported by a grant to the first author from the Netherlands Organization for the Advancement of Pure Research (ZWO).     

Photoperiodic time measurement in the male lizardAnolis carolinensis

Summary Photoperiod plays an important role in controlling the annual reproductive cycle of the male lizardAnolis carolinensis. Groups of anoles were exposed to various experimental lighting regimens to determine how the lizards were measuring the length of the day. The experimental regimens were designed to discriminate between the following two general classes of hypotheses: (1) Photoperiodic time measurement is based on an hourglass or interval timer which measures the length of the light or dark or (2) Photoperiodic time measurement is based on an endogenous circadian rhythm of photoperiodic photosensitivity. The experiments demonstrated thatAnolis uses an hourglass mechanism which measures the absolute length of the light period. This is in contrast to the higher vertebrates (birds and mammals) which measure photoperiodic time by means of a circadian oscillation of responsiveness to light.     

Photoperiodic time measurement in the aphid Megoura viciae

Megoura produces parthenogenetic virginoparae in long day conditions, gamic oviparae in short days. The nature of this photoperiodic response has been analysed by rearing parent apterae in a wide range of circadian and non-circadian light cycles. By varying the light and dark components independently in a two-component cycle it has been established that the time measuring function is associated primarily with the dark period. There is no evidence that an endogenous circadian oscillation is implicated: thus (a) the ‘short day’ response is abolished by ‘night interruptions’ positioned in the early or late night. But this bimodal response pattern remains unchanged when the duration of the ‘main’ photoperiod is varied from ca. 6 hr to at least 25·5 hr. The stability of the maxima within the scotophase is inconsistent with the ‘coincidence’ models of photoperiodic timing that have been proposed. It is suggested that the essential timing process operates on the hour-glass principle, beginning anew with the onset of each period of darkness; (b) night interruption experiments employing very long (up to 72 hr) scanned dark periods yielded response maxima explicable in terms of the hour-glass hypothesis but did not reveal any circadian relationship between the maxima.The ‘dark reaction’ comprises a sequence of four stages, definable by the effects of light. Stage 1, extending from dark hr 0 to ca. 2·5, is fully photoreversible: at the next dark period the entire timing sequence is repeated up to the 9·5 hr critical night length. Towards the end of stage 1 reversibility is gradually lost and after a light interruption the reaction is resumed from a later time equivalent than dark hr 0; the subsequent critical night length is therefore reduced. The extent of the photoreversal is related to light duration. The period of maximum light insensitivity (stage 2) is attained at the end of the fourth hour. From ca. dark hr 5 to just short of the critical night length light exerts an increasingly promotive action which favours the production of virginoparae. This dark process is not photoreversible. Stage 4, which begins at hr 9·5, marks the end of the timing sequence. Light will not then annul the non-promotive action of the previous long night.Light has three effects which are determined by its duration and position within the cycle. The two terminal effects, mentioned above, are associated with the interception of dark stages 1 and 3 by either short (1 hr) or longer photoperiods. Light also prepares or primes the dark period timer. Thus the critical length is increased, and timing accuracy lost, if the preceding photoperiod is less than ca. 6 hr. Light during stage 4 has a priming action but no terminal function. Repeated cycles are ‘read’ in various ways, depending on the cycle structure. For example, if light intercepts stage 3, a two-component cycle is interpreted as the overlapping sequence light/dark/light. One and the same photoperiod then acts terminally in respect of the preceding dark period and as a primer for the next dark period.There is also a mechanism for summing the promotive effects produced by repeated interruption of dark stage 3. With complex (four-component) cycles both halves of the same cycle may contribute. ‘Product accumulation’ falls below threshold if the frequency of presentation of a given promotive cycle is too low. This occurs if there are very long, relatively non-promotive dark components. Such cycles are accepted as ‘continuous darkness’.     

Delayed puberty in lambs chronically treated with oestradiol

Intact female lambs were chronically treated with low levels of oestradiol by Silastic implant from 20 weeks of age. Reproductive cycles were initiated in only 33% of these lambs (3 of 9) compared to 80% of untreated females (11 of 14) by 45 weeks when the study was terminated. Moreover, in the 3 oestradiol-treated lambs which began cycles, the age at first oestrus was delayed 3 weeks (37 +/- 1 weeks of age vs 34 +/- 1 weeks of age for untreated controls). Retardation of the pubertal process was not due to absence of the pubertal rise in circulating LH. At about 32 weeks of age, chronic oestradiol treatment was no longer able to suppress tonic LH secretion and serum LH increased in intact, oestradiol-treated lambs. These results indicate that a maturational decrease in responsiveness to oestradiol inhibition of tonic LH secretion can be demonstrated in the intact female, as in the ovariectomized female. However, chronic oestradiol suppression of prepubertal LH secretion also delays onset of reproductive cycles. This finding raises the possibility that low tonic LH secretion, presumably in the form of slow pulses, is necessary for development or maintenance of ovarian function before puberty. In the absence of LH during the last part of sexual maturation, the ability of the ovary to respond to the high frequency LH pulses during the pubertal gonadotrophin rise may be delayed.     

Photoperiodic time measurement in the male deer mouse, Peromyscus maniculatus

Weanling male deer mice, Peromyscus maniculatus, were exposed for three weeks either to light-dark (LD) cycles with periods (T=L+D) ranging from T=23 (1L:22D) to T=25.16 (1L:24.16D) or to 24-h LD cycles with photoperiods ranging from 1 (1L:23D) to 19 (19L:5D) h. Both the circadian locomotor activity rhythms and the response of the reproductive system to these LD cycles were assessed. The results demonstrate that the photoperiodic effectiveness of light depends on the phase of the light relative to the animal's circadian system, as marked by the circadian activity rhythm. Light falling during the animal's subjective night, from activity onset to at least 11.8 h after activity onset, stimulates growth and maturation of the reproductive system, whereas light falling during the rest of the circadian cycle is nonstimulatory.     

Photoperiodic time measurement in insects: a review of clock models

Based on analyses of responses of insects and mites to a wide range of diel and nondiel experimental light-dark schedules, a variety of models have been developed for the photoperiodic clocks in these species by nearly as many investigators. According to some of these models, the photoperiodic clock is based on a mechanism separate from the circadian system, that is, a so-called "hourglass." According to other models, the clock is based on one or more circadian oscillators that may be coupled to each other and that may or may not show a certain degree of damping. In this context, a rapidly damping oscillator could be regarded as an hourglass. The present article gives an overview of the many different clock models and their philosophies, and it makes comparisons among them to provide a better understanding about how these models are related, if at all, and why the double circadian oscillator model is the most favored model at present.     

Pinealectomy delays puberty in ewe lambs

Fifteen pinealectomized and 15 unoperated ewes were exposed to constant light for 3 weeks before and 10 weeks after lambing. Fourteen pinealectomized and 15 unoperated ewes were allowed to lamb outdoors. Five ewe lambs born in constant light to the 2 groups of dams were pinealectomized at 10 weeks of age. Ewes and lambs were then returned to the field. Puberty (determined by weekly progesterone analysis) was significantly delayed (P less than 0.05) in the pinealectomized ewe lambs. Median pubertal age in pineal-intact ewe lambs was 37 weeks compared to 49 weeks in pinealectomized lambs. Constant light during the first 10 weeks of life had no effect upon puberty onset nor did the pineal status of the dam. Control lambs entered seasonal anoestrus at the time pinealectomized ewe lambs were entering puberty. Pinealectomized lambs entered anoestrus at the same time as control lambs were beginning their second breeding season. These results confirm a key role of pineal-mediated hormonal signals in the control of puberty in the sheep.     

Photoperiodic time measurement in insects and mites: a critical evaluation of the oscillator-clock hypothesis

The validity of the oscillator-clock hypothesis for photoperiodic time measurement in insects and mites is questioned on the basis of a re-interpretation of available experimental evidence. The possible role of the circadian system in photoperiodism in arthropods is critically reviewed. Apart from the outcome of kinetic experiments, based on diel and non-diel light/dark cycles, evidence from various genetic and physiological experiments is discussed in relation to the oscillator-clock hypothesis. The conclusion is that photoperiodic time measurement in insects and mites is performed by a non-circadian 'hourglass' clock. Experimental evidence suggests a non-clock role for the circadian system in the photoperiodic mechanism of insects and mites.     

Photoperiodic time measurement in the spider mite Tetranychus urticae: A novel concept

To explain photoperiodic induction of diapause in the spider mite Tetranychus urticae a new theoretical model was developed which took into account both the hourglass and rhythmic elements shown to be present in the photoperiodic reaction of these mites. It is emphasized that photoperiodic induction is the result of time measurement as well as the summation and integration of a number of successive photoperiodic cycles: the model, therefore, consists of separate ‘clock’ and ‘counter’ mechanisms. In current views involvement of the circadian system in photoperiodism is interpreted in terms of the hypothesis that the photoperiodic clock itself is based on one or more circadian oscillators. Here a different approach has been chosen as regards the role of the circadian system in photoperiodism: the possibility, previously put forward by other authors, that some aspect of the photoperiodic induction mechanism other than the clock is controlled by the circadian system was investigated by assuming a circadian influence on the photoperiodic counter mechanism. The derivation of this ‘hourglass timer oscillator counter’ model of photoperiodic induction in T. urticae is described and its operation demonstrated on the basis of a number of diel and nondiel photoperiods, with and without light interruptions.     

Genome‐wide association with delayed puberty in swine

An improvement in the proportion of gilts entering the herd that farrow a litter would increase overall herd performance and profitability. A significant proportion (10–30%) of gilts that enter the herd never farrow a litter; reproductive reasons account for approximately a third of gilt removals, with anestrous and failure to conceive the most common reasons for culling. Tools to select gilts for reproductive longevity through genomics or alternative phenotypes would be of great benefit to the producer. Ninety‐one gilts that failed to display behavioral estrus by 240 days (cases) and 127 pubertal littermates (controls) were genotyped with the Illumina Porcine SNP60 Beadchip. One hundred and seventy‐four SNPs with the most significant associations were genotyped in an additional 86 cases and 103 controls. Twelve of these associations were significant in the final analysis. The most significant (P < 1.5 × 10^?14) region associated with failure to attain puberty was on chromosome 4 surrounding the NHLH2 gene. Delayed pubertal development and age at first estrus have been associated with NHLH2 in mice. Because attainment of puberty is a complex trait, identifying genes that affect pubertal age would greatly contribute to our knowledge of reproductive development as well as overall fertility.     

Delayed puberty caused by hyperthyroidism in ram lambs is not a result of suppression in body growth

Over a period of 8 weeks ram lambs (16 weeks old) were made hyperthyroidal (serum thyroxine approximately equal to 150 ng/ml, compared with control approximately equal to 48 ng/ml) by daily subcutaneous injections of thyroxine or maintained at a constant body weight by restriction of the feed intake. Hyperthyroidal and restricted-intake lambs remained at a constant body weight during the period of treatment whilst control rams gained body weight. Testicular growth was normal in restricted-intake lambs but was suppressed in hyperthyroidal animals. Hyperthyroidism, but not feed restriction, was also associated with decrease in LH pulse frequency (1.3 +/- 0.3/12 h compared with controls 4.8 +/- 0.9/12 h. Hyperthyroidal lambs showed normal LH responses to exogenous LHRH. After cessation of treatment testicular growth continued to be suppressed for up to 16 weeks in previously hyperthyroidic rams; thereafter testes began to increase in size but at 30 weeks after treatment were still smaller than those of control rams. It is concluded that elevated thyroxine concentrations directly influence sexual maturation in ram lambs through actions at hypothalamic and/or higher brain centres which control LH secretion. Transient hyperthyroidism during sexual maturation may cause permanent impairment of sexual development.     

Reduced Sertoli cell number and altered pituitary responsiveness in male lambs undernourished in utero

The nutritional status of females during pregnancy can have profound effects on the fetus reproductive system development that could affect the reproductive potential as later as in adulthood (fetal programming). The aim of the present study was to examine the effects of maternal nutrient restriction imposed during different periods of gestation on pituitary responsiveness at different ages postnatal and on Sertoli cell number in male offspring. Pregnant ewes were fed to 100% of Metabolizable Energy and Crude Protein requirements throughout pregnancy (Control) or to 50% from 0 to 30 (R1) or from 31 to 100 days of gestation (R2). Male lambs were selected and fed to appetite throughout the study. At 2, 5.5 and 10 months of age a GnRH challenge was conducted. At slaughter (10 months) testes were removed and examined histologically. Maternal undernutrition did not affect the time of the onset of puberty, defined as the first increase in plasma testosterone concentrations > or =1 ng/ml. The LH and FSH response to GnRH challenge did not differ between groups at 2 and 5.5 months but at 10 months of age a higher (P<0.05) FSH response was found in R2 group. Testes weight did not differ between groups at slaughter. Mean Sertoli cell number was significantly lower in animals of R2 group compared with Control (P<0.01). A smaller seminiferous tubules diameter was detected in R2 group (P<0.05), while the diameter of the lumen was similar in all groups. Collectively, these results provide clear evidence for a direct effect of nutrient restriction during pregnancy on Sertoli cell number in adulthood. The lower number of Sertoli cells is the most candidate factor for the higher pituitary activity through a reduction in the suppressive effect of inhibin.     

Photoperiodic control of diapause in Pseudopidorus fasciata (Lepidoptera: Zygaenidae) based on a qualitative time measurement

In the zygaenid moth, Pseudopidorus fasciata, both larval diapause induction and termination are under photoperiodic control. In this study, we investigated whether photoperiodic time measurement (with a 24-h light-dark cycle) in this moth is qualitative or quantitative. Photoperiodic response curves, at 22, 25, and 28 degrees C indicated that the incidence of diapause depended on whether the scotophases exceeded the critical night length (CNL) or not. All scotophases longer than the CNL-induced diapause; all scotophases shorter than the CNL-inhibited diapause. The CNL was 10.5h at 25 and 28 degrees C, and 10h at 22 degrees C. By transferring from various short photoperiods (LD 8:16, LD 9:15, LD 10:14, LD 11:13, LD 12:12, and LD 13:11) to a long photoperiod (LD 16:8) at different times, the number of light-dark cycles required for 50% diapause induction at 25 degrees C was 7.14 at LD 8:16, 7.2 at LD 9:15, 7.19 at LD 10:14, 7.16 at LD 11:13, and 7.13 at LD 12:12, without showing a significant difference between the treatments. Only at LD 13:11 (near the CNL), the number of light-dark cycles was significantly increased to 7.64. The intensity of diapause induced under different short photoperiods (LD 8:16, LD 9:15, LD 10:14, LD 11:13, and LD 12:12) at 25 degrees C was not significantly different with an average diapause duration of 36 days. The duration of diapause induced under LD 13:11 was significantly reduced to 32 days. All results indicate that the night-lengths are measured as either "long" or "short" compared with some critical value and suggest that photoperiodic time measurement for diapause induction in this moth is based on a qualitative principle.     

Treatment of delayed puberty and hypogonadism in girls.

The therapeutic management of female delayed puberty depends more on the objectives than on the underlying cause. We will have to consider the development of sex characteristics, the occurrence of menarche and the promotion of growth. In this paper, we will review how girls with delayed puberty of different etiologies can benefit from the following therapeutic alternatives: follow-up without hormonal therapy; administration of growth hormone, anabolic steroids (e.g. oxandrolone) or estrogens and progestogens, and psychological support.     

Ovarian activity and uterus organometry in delayed puberty gilts

About 30% of the total number of gilts selected for reproduction at the large breeding farm units in Vojvodina (Republic of Serbia) are culled due to prolonged pre-insemination anoestrus (estrus not detected until 8 mo of age). The aim of this study was to provide the answer to the following question: do the culling gilts reach cyclic ovarian activity at all? One hundred seventy five culled gilts in which external estrus manifestations were not detected by 8 mo of age were sacrificed and their reproductive organs were examined for determination of sexual maturity (ovaries exhibiting pre-ovulatory follicles 8 to 11 mm in diameter, corpora hemorrhagica, corpora lutea and corpora albicantia). Uterine weights and horn length were also determined. Functional ovaries were observed in 107 (61.1%) examined gilts, with 62 animals having one and 45 having two puberty ovarian cycles (57.9% and 42.1%, respectively). Pathomorphological changes which could result in prolonged pre-insemination anoestrus were not observed on the reproductive organs of sexually mature gilts. Our results indicate that most of the culling gilts have reached cyclic ovarian activity. The main reason for culling due to the absence of external estrus manifestations in sexually mature gilts could be inadequate estrus detection technology.