Settlement decisions by the two-spotted spider mite Tetranychus urticae

In silk-spinning arthropods, silk can be used for web building, protection, and communication. Silk is an informative material about the presence of conspecifics. It can therefore inform on habitat suitability and hence assist in habitat choice. In this context, we investigated the influence of silk on microhabitat choice by the two-spotted spider mite, Tetranychus urticae. Three factors that could potentially influence habitat choice were manipulated: the strain, number, and the stage of mites. Our study showed that these factors all influence the choice of microhabitat. The tendency of whether to settle on a silk-covered area was influenced by the origin of mites (strain effect). Adult females showed a higher tendency to settle on an area covered with the silk laid by numerous congeners (number effect). Moreover, larvae seemed to be more responsive to the presence of silk than adults (stage effect). This suggests that individuals use silk as a social cue in selecting their microhabitat and that the spatial organization and group behaviour seem to be shaped by the individuals’ response to social cues, such as the amount of silk already present.     

UV tolerance in the two-spotted spider mite, Tetranychus urticae

The two-spotted spider mite, Tetranychus urticae was exposed to UV-C (250 nm), UV-B (300 nm), and UV-A (350 nm). In non-diapausing females, the median effective doses for 50% mortality plus escape incidence (ED₅₀) were 21 (UV-C) and 104 kJ m⁻² (UV-B); those for 50% oviposition rate in continuous darkness-treated mites were 6.2 (UV-C) and 41 kJ m⁻² (UV-B). No significant effects of UV-A on mortality and oviposition rate were observed. The ED₅₀ values for UV-B were similar to the natural UV-B observed for 2-5 days in summer when T. urticae inhabits the undersides of leaves. Therefore, T. urticae possibly uses leaves as a filter to avoid the deleterious effects of UV-B. In diapausing females, low mortality was observed even at high doses of UV radiation, but more than half escaped even at low doses. The orange body color of diapausing females results from accumulation of carotenoids, a scavenger for UV-induced reactive oxygen species; this may explain the low mortality of diapausing females. Diapausing females may overcome the deleterious effects of UV-B during winter in the absence of leaves by emigrating to UV-free environments and by accumulating carotenoids.     

Effects of acclimation and diapause on the thermal tolerance of the two-spotted spider mite Tetranychus urticae

The two-spotted spider mite, Tetranychus urticae, is a worldwide pest species that overwinters as diapausing females. Cold hardening is presumed to start during diapause development to ensure the successful overwintering of this species. To address this hypothesis, we compared cold tolerance between non-diapausing and diapausing females. We measured supercooling point (SCP) and survival to acute cold stress by exposing the mites at a range of sub-zero temperatures (from −4 to −28 °C for 2 h). The mean SCPs of non-diapausing and diapausing females were −19.6±0.5 and −24.7±0.3 °C respectively, and freezing killed the mites. Diapausing females were significantly more cold tolerant than non-diapausing ones, with LT₅₀ of −19.7 and −13.3 °C, respectively. Further, we also examined the effects of cold acclimation (10 d at 0 or 5 °C) in non-diapausing and diapausing females. Our findings indicated that diapause decreased SCP significantly, while cold acclimation had no effect on the SCP except for non-diapausing females that were acclimated at 5 °C. Acclimation at 5 °C enhanced survival to acute cold stress in diapausing and non-diapausing females, with LT₅₀ of −22.0 and −17.1 °C, respectively. Altogether, our results indicate that T. urticae is a chill tolerant species, and that diapause and cold acclimation elevate cold hardiness in this species.     

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 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.     

Characterization of vitellin protein in the twospotted spider mite, Tetranychus urticae (Acari: Tetranychidae)

In mites, vitellogenin synthesis, regulation and uptake by the oocytes as vitellin remain practically unknown. Although a partial sequence of the gene is now available, no previous studies have been conducted that describe the native vitellin protein in mites. The objective of this study was to characterize vitellin in the twospotted spider mite, Tetranychus urticae. The native twospotted spider mite vitellin migrated as a single major band with a molecular weight of 476 ± 14.5 kDa as compared to 590 ± 25.5 kDa for vitellin from the American dog tick, Dermacentor variabilis. However, isoelectric focusing analysis of native spider mite vitellin showed five bands with pI values slightly acidic to neutral (pH 5.8, 6.2, 6.7, 7.0 and 7.2), as is the case for insect and tick vitellins. Reducing conditions (SDS-PAGE) also revealed multiple subunits ranging from 290.9 to 3.6 kDa and was similar to that found in D. variabilis. Spider mite vitellin weakly bound lipids and carbohydrates compared to the tick. Unlike D. variabilis, the spider mite egg yolk protein does not bind heme. The significance of non-heme binding in mites is discussed.     

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.     

Effect of host plant nitrogen fertilization on the biology of the two-spotted spider mite, Tetranychus urticae

Females of Tetranychus urticae Koch were reared on leaf discs of apple trees and bush beans grown at different N concentrations (0.6–75 mM NO _inf3 ^sup- ). N-deficiency increased pre-imaginal development time, preoviposition period, and decreased female weight, fecundity and oviposition rate of the mites. N, water, amino acid, and sugar content of the apple leaves were positively correlated with weight and egg production and negatively correlated with development time and pre-oviposition period. The reverse correlations were found with total phenol content of the leaves and above mite parameters. A reduction of leaf N by 50% was related with a tenfold decline in fecundity on apple leaves. The stress mainly affected the oviposition rate and to a lesser extent the oviposition period. On apple leaves the net reproductive rate (av. no. offspring per , R_o), mean length of a generation (T), and innate capacity for increase (r_m) were R_o=40.3, T=17.1, and r_m=0.22 for the standard N concentration, and 4.7, 25.0, and 0.06 for strong N-deficiency, respectively.

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Zusammenfassung Die polyphage Gemeine Spinnmilbe ist ein wirtschaftlich bedeutender Schädling. Die Rolle einzelner Nährstoffe bei der Milbenvermehrung wurde schon von einigen Autoren mit unterschiedlichen Ergebnissen untersucht. Der Stickstoff scheint aber einer der Hauptfaktoren zu sein.Ueber Gewebekulturen klonierte Apfelbäume sowie aus Samen gezogene Buschbohnen wurden in Nährlösungen mit verschiedenen N-Angeboten kultiviert. Aus solchen Pflanzen wurden Blattscheiben ausgestanzt und mit Weibchen von T. urticae besetzt. Gemessen wurden die tägliche Eiablage bis zum natürlichen Tod, die Entwicklungsdauer und das Weibchengewicht. Mit dem bei N-Mangel abnehmenden N, Aminosäuren- und Zuckergehalt (v.a. Sorbit) im Blatt korrelierend nahmen auch Gewicht, Ablagerate und Fekundität ab, bzw. die Entwicklungsdauer und Praeovipositionsperiode zu. Die umgekehrte Wirkung auf diese Milbenparameter hatte der Gesamtphenolgehalt. Eine Reduktion des N-Gehalts der Blätter auf die Hälfte (1.5% N) bewirkte auf Apfelblättern eine Abnahme von Fekundität und Ovipositionsrate um das zehnfache, bzw. eine Zunahme der Präovipositionsperiode um mehr als das dreifache. Der Stress beeinflusste v.a. die Maxima der Ablageverläufe und weniger die Lebensdauer. Der Wassergehalt war ebenfalls mit der Fekundität positiv korreliert.Auf Bohnenblättern legten die Tiere mehr Eier und erreichten ein höheres Gewicht als auf Apfel, dies bei gleichem N-Gehalt beider Pflanzen. Es wirken somit noch andere Faktoren auf die Reproduktion der Spinnmilben.Die Populationsparameter wurden ebenfalls sehr stark beeinflusst. Auf Apfelblättern ergab sich bei der Kontrolle eine Nettoreproduktionsrate R_o von 40.3 und bei starker N-Defizienz 4.7. Die mittlere Generationsdauer T sowie die spezifische natürliche Wachstumsrate r_m betrugen für die Kontrolle 17.1 Tage, bzw. 0.22 und für starken N-Mangel 25.0 Tage, bzw. 0.06.Die Ergebnisse zeigen, dass sich bereits kleinere Unterschiede im Stickstoffgehalt und damit zusammenhängend im Zucker- und Phenolgehalt stark auf die Populationsdynamik der Gemeinen Spinnmilbe auswirken können.

     

Intermittent character of adult diapause in Aelia acuminata (Heteroptera)

The response to photoperiod of field-collected Aelia acuminata was studied by exposing insects to three photoperiodic regimes. After the completion of diapause the insects do not respond to photoperiod: in mid-February the length of the pre-oviposition period is the same under both long and short photoperiods. After the onset of oviposition, insects again become responsive to photoperiod regardless of the time of year: under long day, oviposition lasts until death, whereas under short day the females cease oviposition after about 3 weeks. After a further 2 or 3 months without ovipositing, these females which are still experiencing short-day conditions resume reproduction for a further 4–7 weeks and long-lived individuals have three oviposition periods. A regulatory mechanism is suggested for the alternation between oviposition and diapause.     

Structural and expression analysis of hepatic vitellogenin gene during ovarian maturation in Anguilla japonica

Vitellogenin (Vtg), the precursor molecule for yolk, is synthesized in the liver under estrogenic control. In all oviparous species, including fish, the process of vitellogenesis is crucial to subsequent embryonic development. This study attempted to obtain the cDNA encoding for Vtg from female Japanese eels, Anguilla japonica. Rapid amplification of cDNA ends (RACE) and polymerase chain reaction (PCR) were used to amplify Vtg cDNA prepared from liver extracts. Obtained PCR products were subcloned and sequenced. The overall sequence of eel Vtg cDNA isolated in this study contained 5395 bp nucleotides. This Vtg sequence encodes 1743 amino acids of the precursor molecule, and is entirely composed of the characteristic N-terminal lipovitellin-I region, an internal polyserine domain region, and a c-terminal lipovitellin-II region. The deduced amino acid sequence from these clones shares 34–61% identity with other teleost Vtgs. Northern blot assays of Vtg gene expression following hormonal treatment demonstrated that this Vtg is synthesized in the liver under stimulation by estradiol injection. However, Vtg synthesis may not be enhanced by salmon pituitary homogenate (SPH) induction for the developing ovarian follicles. Notably, the effect of methyltestosterone, following SPH injection, may be more appropriate for the uptake of Vtg by ovarian follicle maturation during the artificial maturation of Japanese female eels.     

Effects of inbreeding on variation in diapause duration and early fecundity in the Kanzawa spider mite

In arthropods, both diapause duration and ability to produce eggs in early adult life (early fecundity) are important life‐history traits for successful settling in a new habitat. In herbivorous spider mites (Acari: Tetranychidae), inbreeding frequently occurs because new colonies are established by only one or a few females. In the present study, we investigated the impact of inbreeding on the phenotypic variance in diapause duration and early fecundity in the Kanzawa spider mite, Tetranychus kanzawai Kishida. Diapause duration was compared between the inbreeding treatment (strongly inbred strains) and the control (strains immediately taken from the stock culture) under winter‐mimicking laboratory conditions. The variance in diapause duration was smaller in the inbreeding treatment than in the control, though the magnitude of decrease of variance was less than expected. On the other hand, early fecundity did not show any reduction of variance. The present results revealed that inbreeding reduces phenotypic variation, as expected in theory.     

Effects of host-plant species on diapause induction of the Kanzawa spider mite, Tetranychus kanzawai

In many herbivorous arthropods, incidence of diapause, which is considered to reflect the timing of diapause, changes depending on the host plants they utilize. Several theoretical studies suggest that the optimal timing of diapause induction depends on life‐history traits; if the development time of the arthropod is short, fecundity is high, or survival rate remains high throughout the season, the optimal timing of diapause induction would be shifted toward the end of the season. For herbivorous arthropods, these life history traits may change among their host plants. Here we examined whether a population of the Kanzawa spider mite, Tetranychus kanzawai Kishida (Acari: Tetranychidae), shows the predicted pattern of diapause induction on two host plants, the kidney bean [Phaseolus vulgaris (Leguminosae)] and Japanese Orixa [Orixa japonica (Rutaceae)], on which the mites show different performances. Rearing conditions were controlled in two ways. In the first experiment, day length and temperature were kept constant throughout the mite lifetime at either of three conditions from mid October to early November. In the second experiment, the conditions were changed from 20 °C and L11.5:D12.5 at immature stages to 18 °C and L11:D13 at adult stage in order to better approximate field conditions. In the first experiment, diapause incidence on P. vulgaris was lower than on O. japonica. This tendency became stronger in the second experiment, suggesting a difference in the timing of diapause induction among host plants. On the other hand, P. vulgaris was proven to allow high performance, i.e., greater lifetime fecundity and shorter development times, although it had no effect on the survival rate. The relationship between diapause incidence and performance is consistent with the prediction of theoretical studies that a short development time or high fecundity delays the timing of diapause induction.     

Inter- and intra-population variations in diapause attribute of the two-spotted spider mite,Tetranychus urticae Koch,in Japan

Populations of the two-spotted spider mite, Tetranychus urticae Koch collected from various localities and from various host plants in Japan showed wide variations in diapause attribute. Diapause percentages at 18°C/9L15D varied from nearly 100% in the north to 0% in the south-west. At intermediate latitudes the mites showed wide inter-population variations. Populations on herbaceous hosts in vinyl- or glass-houses gave significantly lower incidence of diapause than those on roses and deciduous fruit trees. Presence of winter hosts and better host quality under protected environments seemed to favour non-diapausing mites. The temperature threshold for diapause expression also varied widely among local populations. Northern populations consistently had higher and less variable thresholds than populations at intermediate latitudes with thresholds between 15 and 18°C. Inbred lines derived from a population in Kyoto exhibited a wide variation in diapause percentage at 18°C. These results show that diapause in T. urticae is a quantitative threshold trait and that populations in central Japan consist of a variety of genotypes with different diapause traits. This might provide a genetic source for adaptation to local and temporal variations in environmental conditions.     

Geographic variations and combining abilities in the two-spotted spider mite, Tetranychus urticae

Geographic variations and combining abilities of three fitness traits: the mean number of offspring, and the mean number and proportion of female offspring, in the two-spotted spider mite were investigated using four widely separated strains of Tetranychus urticae Koch. Analyses of data obtained from a complete set of diallel crosses, involving the four strains, revealed large and highly significant specific combining abilities and reciprocal effects in all the fitness characteristics examined. It was concluded that the whole chain of reproductive events, from egg production of parental females to the fertility of hybrid daughters could be severely affected by incompatibility between parental populations.

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Résumé L'étude a porté sur les différences géographiques et les combinaisons de trois caractères adaptatifs—nombre moyen de descendants, nombre moyen et proportion de femelles dans la descendance—sur quatre souches différentes de Tetranychus urticae. L'analyse des données obtenues à partir d'un lot complet de croisements dialléliques, sur les quatre souches, révèle pour tous les caractères adaptatifs examinés une forte aptitude à l'association et aux effets réciproques. On peut en conclure que la chaîne totale des événements intervenant dans la reproduction—de la production d'oeufs par les mères jusqu'à la fécondité des filles hybrides—pourrait être fortement modifiée par l'incompatibilité entre populations parentales.

     

Cold hardiness in the two-spotted spider mite (Tetranychus urticae Koch)

Summary The effect of temperatures below 0° C on different stages of the two-spotted spider mite, and in particular on active adult females, has been investigated.Survival of active females at-5° was higher at approximately 100% relative humidity than at a lower degree of humidity. Higher rates of survival were also caused by acclimation of active females and larvae for one day at +5° before storage at-15°. No differences in cold-hardiness were found in active females from an organophosphorous resistant strain and a susceptible strain, and the ability to survive depended on the time-temperature exposures (see Fig. 3).Eggs were killed very rapidly at-15° C, while larvae survived longer exposures than active females. Compared at the LT₅₀ level females in diapause survived three times the exposure endured by acclimated active females.Supercooling points measured at a rate of cooling of 2° C per minute were highest in diapausing females and lowest in eggs. Acclimation had no effect on the supercooling points of active females. All stages were killed by freezing. It seems likely that the difference in survival between acclimated and not acclimated active females at-15° is caused by different ability to live in a supercooled state.

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Zusammenfassung Es wurde die Wirkung von Temperaturen unter 0° auf verschiedene Stadien der Bohnenspinnmilbe, insbesondere auf aktive erwachsene Weibchen untersucht.Die Überlebensrate aktiver Weibchen war bei nahezu hundertprozentiger relativer Luftfeuchtigkeit höher als bei niederer Feuchtigkeit. Höhere Überlebensraten wurden auch durch Akklimatisation aktiver Weibchen und Larven für einen Tag bei 5° vor der Behandlung mit-15° erreicht. Es wurden keine Unterschiede in der Kältewiderstandfähigkeit aktiver Weibchen eines phosphorsäureesterresistenten und eines anfälligen Stammes gefunden. Die Überlebensfähigkeit hängt von Dauer und Temperatur der Einwirkung ab.Eier wurden von-15° sehr schnell abgetötet, während Larven längere Einwirkungszeiten überlebten als aktive Weibchen. Bei einem Vergleich der LT₅₀ überlebten diapausierende Weibchen dreimal längere Einwirkungszeiten, als sie von akklimatisierten aktiven Weibchen ertragen wurden.Unterkühlungspunkte—gemessen bei einer Abkühlungsrate von 2° pro Minute—lagen bei diapausierenden Weibchen am höchsten und bei Eiern am niedrigsten.Akklimatisierung hatte keinen Einfluß auf die Unterkühlungspunkte aktiver Weibchen. Durch Gefrieren wurden alle Stadien getötet. Es erscheint möglich, daß der Unterschied im Überleben akklimatisierter und nichtakklimatisierter Weibchen bei-15° durch die verschiedene Fähigkeit bedingt ist, in einem unterkühlten Zustande zu leben.

     

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.     

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.     

Maintenance of diapause variability in the two-spotted spider mite,tetranychus urticae, in a heterogeneous and stochastic environment

A simple nongenetic mathematical model analyzed the processes responsible for the variations in the diapause percentage among populations of the two-spotted spider mite,Tetranychus urticae Koch. This model incorporates the following assumptions. 1) Mites have diapause (DD), non-diapause (NN), and “plastic” (DN) populations (plasticity exists in the phenotypic expression of diapause in response to habitat conditions at the time of diapause induction). 2) A heterogeneous mite habitat consists of microhabitat L, in which all the non-diapausing mites die during the winter due to the lack of winter host plants, and microhabitat O with winter hosts capable of supporting some of the non-diapausing mites overwinter. 3) Temporal fluctuation of winter conditions which affect the survival and reproduction of non-diapausing mites. Using these assumptions, we compared the fitness functions of the three populations and analyzed the conditions under which each population is favoured over the other two, thereby elucidating the processes involved in the maintenance of variability in diapause. Our analysis revealed: 1) frequent mild winters are of primary importance for the non-diapause trait to be maintained, 2) the existence of winter hosts is also important for the nondiapause trait to be favoured, and this importance depends greatly on the degree of the adaptive diapause expression in the DN mites in response to habitat conditions, i.e., the better the phenotypeenvironment matching in DN, the higher the probability that DN will be favoured, 3) The combined effect of the temporal and spatial variation enhances the maintenance of variablity in the diapause trait of the mites.     

Ultrastructural evidence for an osmoregulatory effect of nikkomycin in the mite Tetranychus urticae

Histological effects of the microbial metabolite and chitin synthesis inhibitor complex Nikkomycin (AMS 0896 Bayer Leverkusen) on osmoregulatory organs of all developmental stages of Tetranychus urticae are described. The metabolite, in a concentration of 100 ppm, was applied via the nutritive plant. Mites fed for 2 to 14 days, and then were collected and immediately fixed. Two osmoregulatory organs occur in T. urticae. The Malpighian complex, differentiated only in females, shows an increased number of apical microvilli in the epithelium of the distal regions after metabolite application, thus resulting in an enlargement of the surface area. Changes in the second osmoregulatory organ, the coxal organ, after Nikkomycin application include depositions of membranous bodies in the lumen as well as in cytoplasmic vacuoles of the proximal tubule. Additionally, an increase in the luminal diameter occurs. Numerous vacuoles of different contents are observed in the cytoplasm of the distal tubule. Consequences of histological alterations in osmoregulatory organs after Nikkomycin application are discussed with special reference to the composition of salivary secretions.