External coincidence and photoperiodic time measurement in the spider mite Tetranychus urticae

The incidence of diapause in the spider mite Tetranychus urticae was predicted for various photoperiodic regimes, according to the external coincidence model of photoperiodic time measurement. A phase response curve was constructed for the hypothetical photoperiodic oscillator in these mites: entrainment of this photoperiodic oscillator to a variety of ‘complete’ and ‘skeleton’ photoperiods was calculated using a transformation method for circadian rhythms. The external coincidence model proved adequate to describe experimental results with T. urticae in ‘complete’ photoperiods (T = 24 hr), symmetrical ‘skeleton’ photoperiods (T = 24 hr), asymmetrical ‘skeleton’ photoperiods (T = 24 hr) (night-interruption experiments), and ‘resonance’ experiments, in which the light component of a light/dark cycle was held constant at 8 hr and the dark component was varied over a wide range in successive experiments, providing cycles with period lengths up to 92 hr. The external coincidence model proved inadequate to explain results obtained in a ‘T-experiment’ with T. urticae comprising 1 hr pulses of light in a cycle of LD1:17.5 (T = 18.5 hr) with the first pulse of the train starting at different circadian phases. The validity and limitations of the external coincidence model as an explanation of photoperiodic time measurement in T. urticae are discussed in view of the above results.     

Light-break experiments and photoperiodic time measurement in the spider mite Tetranychus urticae

To explain photoperiodic induction of diapause in the spider mite Tetranychus urticae (Acarina: Tetranychidae) a theoretical model was developed, consisting of two components, viz. a “clock” and a photoperiodic “counter” mechanism. The clock executes photoperiodic time measurement according to hourglass kinetics; the counter accumulates the photoperiodic information contained in a number of successive $\text{light}\text{dark}$ cycles by adding up the number of “long” and “short” nights experienced by the developmental stages of the mites sensitive to the photoperiod. The influence of the circadian system on photoperiodic induction is interpreted as an inhibitory effect exerted on the expression of the photoperiodic response; this effect is encountered only in certain photoperiodic regimes, where the circadian system and the photoperiod are out of “resonance” with each other. This “hourglass timer oscillator counter model”, devised to give a theoretical explanation of photoperiodic time measurement, the summation of photoperiodic information, and the influence of the circadian system on photoperiodic induction, proved to be consistent with experimental results obtained with T. urticae in both symmetrical and asymmetrical “skeleton” photoperiods, the latter based on diel as well as non-diel $\text{light}\text{dark}$ cycles.     

Photoperiodic induction of diapause in the spider mite Tetranychus urticae: qualitative or quantitative time measurement?

Abstract. Insects and mites may measure photoperiods eitfier by classifying them as long or short relative to a critical value (qualitative time measurement) or by using the absolute value (quantitative time measurement). The spider mite Tetranychus urticae is thought to use a qualitative mechanism of time measurement. In this paper we present the results of experiments with an inbred line of the spider mite (to keep genetic variation in photoperiodic responses small), to test whether quantitative aspects also play a role. Differences in diapause incidence in different long-night photoperiods at different temperatures may be an indication of quantitative responses to photoperiod. The effect of temperature on the photoperiodic response curve was studied at 16^oC, 19^oC and 22^oC. The response curves appeared to be similar at 16^oC and 19^oC, with a critical nightlength between 10 and 11 h. At 22^oC, diapause induction was less than 100% in all long-night regimens and die critical nightlength had shifted to 12 h. Maximum diapause induction (93%) occurred in a light-dark cycle with a 16 h dark phase (LD 8:16 h). Diapause induction was lowest in long-night photoperiods with dark phases of 20 h and longer. The number of light-dark cycles needed for 50% diapause induction at 19^oC varied. between 12.1 and 14.7 for LD 6:18 h, between 10.9 and 12.5 for LD 8:16 h, between 10.6 and 11.6 for LD 10:14 h, and between 10.1 and 10.7 for LD 12:12 h. Independent of die light-dark regimen, diapause induction took place in some individuals after receiving 8 cycles and virtually all individuals entered diapause after 16 cycles. No effect was found of the photoperiodic treatment during prediapause development (LD 6:18 h, LD 8:16 h, LD 10:14 h, LD 12:12 h) on diapause duration. The average diapause duration at LD 10:14 h and 19^oC was 61 days over all four treatments. We explained the results by hypothesising that nightlengths are assessed qualitatively and mat the photoperiodic clock operates more accurately near the critical nightlength.     

Photoperiodic induction of pupal diapause in Sarcophaga argyrostoma: Temperature effects on circadian resonance

Larval cultures of the flesh-fly Sarcophaga argyrostoma maintained in circadian ‘resonance’ experiments produced a high incidence of pupal diapause when the period of the light cycle was close to (T) 24, 48 or 72 hr, but a low incidence of diapause at T 36, 60 or 84 hr. Cultures pre-programmed for diapause by exposing pregnant females to long nights indicated the induction of non-diapause development at T 36, 60 and 84, whereas cultures pre-programmed for diapause-free development by exposing females to continuous light indicated the induction of diapause at T 24, 48 and 72.Raising the temperature reduced the heights of the diapause peaks whereas lowering the temperature raised them. With progeny from long-night-reared flies the lowest temperature tested (18°C) produced a result indistinguishable from an ‘hour-glass’ response, warning that ‘negative’ resonance experiments may merely indicate non-permissive conditions for demonstrating the involvement of circadian rhythmicity in insect photoperiodism.The results of the ‘resonance’ experiments and the effects of temperature are interpreted in terms of a multioscillator ‘external coincidence-photoperiodic counter’ model for the clock.     

Geographic variation of diapause intensity in the spider mite Tetranychus urticae

Abstract. Eight strains of the spider mite Tetranychus urticae, originating from different localities in western and central Europe, with latitudes ranging from 40.5 to 60^oN, displayed marked differences in the period of chilling at 4^oC required for diapause termination under a diapause-maintaining short-day photoperiodic regime at 19^oC, to which the mites were transferred after the cold period. The higher the latitude from which the strains originated the longer was the period of chilling required for diapause termination, suggesting the presence of a gradient in diapause intensity, diapause being deeper the more northern the origin of the strains. Two strains originating from higher altitudes appeared to have a much deeper diapause than expected from their latitudinal origin. In addition, these two mountain strains showed mutual differences in diapause intensity, notwithstanding the fact that they originated from similar latitudes and altitudes; local climatic conditions probably act as strong selective forces with regard to diapause depth. All strains appeared to be sensitive to photoperiod during the period of diapause development. Diapause was quickly completed by a long-day photoperiod (LD 17:7 h), but was maintained by a short-day photoperiod (LD 10:14h). However, even under the latter regime sensitivity to photoperiod gradually diminished and eventually disappeared, thus leading to ‘spontaneous’ termination of diapause. The length of the period of diapause development, as measured by the sensitivity to photoperiod of diapausing mites, varied between strains; it was shorter in the southern strains and longer in the northern strains. The results indicate great variation in diapause intensity between strains, which is probably genetically determined and may have adaptive significance for this widespread species. When young females which had just entered diapause were kept for ever longer periods of time under the diapause inducing short-day regime at which they had been reared, before being transferred to the cold room, the duration of the period of chilling required for diapause termination was found to decrease proportionally in all three strains tested. These results suggest that intensification of diapause does not occur in T. urticae; diapause intensity seems to be highest at the beginning of diapause and to diminish gradually during diapause development.     

Effects of changing photoperiods in the life cycle regulation of the comma butterfly, Polygonia c-album (Nymphalidae)

Abstract. 1. Regulation of adult reproductive diapause and seasonal polyphenism was studied in two populations of the comma butterfly, Polygonia c-album (L.) (Nymphalidae), from Stockholm, Sweden, and Oxford, England.
2. In the univoltine Stockholm population short and long constant day-lengths (in the range 12–22 h) at 20°C were ineffective in averting the production of the dark, generally diapausing, morph, but the non-diapausing light morph could be produced if daylengths were increased during larval development. The procedure was especially effective with increases from 12 to 22 h, but the tendency was the same with a more realistic increase, from 18 to 20 h.
3. In the partially bivoltine English population a critical constant day-length for morph and diapause induction was found between 12 and 18 h. Decreasing daylengths above the critical daylength early or late in larval development resulted in production of the diapausing morph.
4. These results suggest a system for environmental control where day-lengths which increase throughout the larval period indicate an early date before summer solstice, meaning that there is sufficient time for a second generation of offspring to reach the hibernating adult stage before winter. The ecological significance of the results is discussed.     

Deuterium oxide alters pupal diapause response in the flesh fly, Sarcophaga crassipalpis

ABSTRACT. Deuterium oxide averts pupal diapause in the flesh fly Sarcophaga crassipalpis Macquart when fed to larvae or when applied topically to photosensitive embryos exposed to short daylength. Deuterium oxide was not effective in promoting diapause when presented to embryos or larvae reared at long daylength. The effect of deuterium oxide appears to be cumulative in the larval state: increasing exposure time progressively reduces diapause response. If flies reared on deuterium oxide are exposed to continuous darkness, the diapause response remains high, thus implying that the physiological capacity for diapause is not disrupted. We suggest that deuterium oxide exerts its effect on the circadian rhythm controlling diapause induction.     

Controversial aspects of photoperiodism in insects and mites

This review examines several controversial aspects of photoperiodism in insects and mites including the role of the circadian system in night length measurement, the nature of apparent hourglass-like responses, and whether or not the circadian component in photoperiodism is the same as that in overt behavioural rhythms. These aspects of the phenomenon are discussed in terms of the entrainment of circadian oscillations by cycles of light and temperature. There is considerable variety of photoperiodic response within the insects (and other arthropods) to show, inter alia, circannual rhythms, internal and external coincidence night length timers, and in some species, non-circadian hourglass-like devices. Many apparent hourglass-like responses, however, could be circadian ‘clocks’ of the external coincidence type involving oscillations that dampen below threshold in extended periods of darkness. The review also concludes that there is little evidence in favour of the “Hourglass clock-oscillator counter” model proposed for the mite Tetranychus urticae by Vaz Nunes and Veerman (1982a). The responses of this species to complex light and temperature cycles may also be interpreted in terms of a damped oscillator version of external coincidence.     

Involvement of vitamin A in the photoperiodic induction of diapause in the spider mite Tetranychus urticae is demonstrated by rearing an albino mutant on a semi-synthetic diet with and without p-carotene or vitamin A

Abstract. Data are presented on the development from egg to adult of a wild-type and an albino strain of the spider mite Tetranychus urticae on a semi-synthetic liquid diet.The diet is a modification of the diet developed by Van der Geest et al. (1983).Apart from the standard diet without added carotenoids, diets supplemented with either β-carotene or vitamin A were tested.A high incidence of diapause was found in the wild-type strain under short-day conditions on all three diets.No diapause was found in the albino mutant under short days on the standard diet.Partial restoration of the photoperiodic response was obtained after addition of β-carotene to the diet, and full restoration was found after the addition of vitamin A.Diapause could be terminated in diet-reared diapausing females of both strains after a cold rest of 3 weeks at 4°C.The results lead to the conclusion that vitamin A or a derivative of vitamin A probably functions as the photoreceptor pigment for the photoperiodic induction of diapause in the spider mite.     

Photoperiodic control of adult diapause in the predacious mite, Amblyseius potentillae: repeated diapause induction and termination

ABSTRACT. Photoperiodic control of facultative reproductive diapause was examined in females of the phytoseiid mite, Amblyseius potentillae (Garman). Full diapause was induced by rearing the mites from egg to adult under short-day photoperiods. Egg-laying females, however, which had experienced a long-day photoperiodic regime during pre-imaginal development, could still be induced to enter diapause when they were transferred to a short-day regime. Diapause development proceeded slowly under a short-day photoperiod, but was accelerated under long days. During diapause development under short days the photoperiodic sensitivity of the females diminished gradually and ultimately disappeared completely. However, after completion of diapause, sensitivity to photo-period reappeared. A second diapause could be induced in post-diapause females under short-day photoperiods and completed again under long-day photoperiods. These results show that A. potentillae remains sensitive to diapause-inducing and diapause-averting daylengths during the adult stage and that a second diapause may be induced after completion of the first one.     

Hourglass and rhythmic components of photoperiodic time measurement in the pitcher plant mosquito, Wyeomyia smithii

The mosquito, Wyeomyia smithii, enters a larval dormancy or diapause that is initiated, maintained, and terminated by photoperiod. The median or critical photoperiod regulating diapause increases from 12 h of light per day along the Gulf of Mexico, USA (30° N), to over 15 h in southern Canada (49° N). Photoperiodic time measurement in W. smithii comprises both rhythmic and hourglass (interval timer) components. Using interrupted-night and resonance experiments, we show that both the rhythmic and hourglass components are prominent in the southern (ancestral) populations and that the influence of the rhythmic component declines with increasing latitude, while the hourglass component remains strong in northern (derived) populations. Previously, it has been shown that the genetic differences in critical photoperiod between northern populations and their southern ancestors involve not only the additive (independent) effects of genes, but also gene-gene interaction (epistasis). We therefore conclude that adaptive evolution of W. smithii has probably involved the progressive epistatic masking of the ancestral rhythmic component resulting in photoperiodic time measurement in northern populations accomplished principally through a day-interval timer. A comparison of W. smithii with previous studies indicates that the decline in critical photoperiod with increasing latitude represents an overall decrease in response to light rather than a shift in the timing of photosensitivity among arthropods in general. We propose that the underlying functional components of photoperiodic time measurement, as well as the overt photoperiodic response, are either homologous or are themselves responding directly to selection over latitudinal gradients in seasonality. Received: 18 May 1998 / Accepted: 14 September 1998     

捕食螨在我国农林害螨生物防治中的应用

捕食螨是农林生产中极具价值的生防作用物。文章对捕食螨在我国农林害螨生物防治中的应用作一综述,内容包括:与释放应用相关的研究(引进与风险评估,饲养、贮存与运输,食性与释放,捕食与繁殖,定殖与扩散,外界影响因子)、几种主要捕食螨(智利小植绥螨Phytoseiulus persimilis、西方盲走螨Typhlodromus occidentalis、伪钝绥螨Amblyseius fallacis、胡瓜钝绥螨A.cucumeris)的应用、释放应用前景与展望等,旨在提高我国的捕食螨研究与应用水平。     

The effects of feedback lighting on the circadian drinking rhythm in the diurnal new world primate Saimiri sciureus

Feedback lighting provides illumination primarily during the subjective night (i.e., the photosensitive portion of the circadian cycle) in response to a given behavior. This technique has previously been used to test the nonparametric model of entrainment in nocturnal rodents. In three species (Rattus norvegicus, Mesocricetus auratus, and Mus musculus), the free-running period of the locomotor activity rhythm was similar whether the animals were exposed to continuous light or discrete light pulses occurring essentially only during the subjective night (i.e., feedback lighting). In the current experiments, feedback lighting was presented to squirrel monkeys so that light fell predominantly during the subjective night. Feedback lighting was linked to the drinking behavior in this diurnal primate so that when the animal drank, the lights went out. Despite the seemingly adverse predicament, the monkeys maintained regular circadian drinking rhythms. Furthermore, just as the period of the free-running activity rhythms of nocturnal rodents exposed to continuous light or feedback lighting were similar, the period of the drinking rhythms of the squirrel monkeys in continuous light and feedback lighting were comparable (25.6 +/- 0.1 and 25.9 +/- 0.1 hours, respectively), despite a substantial decrease in the total amount of light exposure associated with feedback lighting. The free-running period of monkeys exposed to continuous dark (24.5 +/- 0.1 hours) was significantly shorter than either of the two lighting conditions (P < 0.001). The results presented for the drinking rhythm were confirmed by examination of the temperature and activity rhythms. Therefore, discrete light pulses given predominately during the subjective night are capable of simulating the effects of continuous light on the free-running period of the circadian rhythms of a diurnal primate. The response of squirrel monkeys to feedback lighting thus lends further support for the model and suggests that the major entrainment mechanisms are similar in nocturnal rodents and diurnal primates.