Geographic and parental influences on diapause by a polyphagous insect herbivore

1 Facultative diapause permits insects to initiate an additional generation when conditions are favourable, or to enter diapause when they are not. Although the principle environmental cues for diapause induction are often temperature and photoperiod, we demonstrated recently that the quality of host plant consumed by the larvae of Choristoneura rosaceana (Harris) (Lepidoptera: Tortricidae) influences the proportion of larvae that enter diapause and therefore voltinism. 2 Previous work has also suggested that non‐Mendelian parental effects influence diapause by C. rosaceana larvae, but the mechanism was not established. Here, we report further evidence for parental effects on diapause and examine whether parental diet influences diapause induction. 3 We collected larvae of C. rosaceana from six sites at three latitudes in the Province of Québec. There was an increase in the proportion of larvae entering diapause with increasing latitude. Rearing larvae under identical conditions on artificial diet for two generations significantly reduced variation in diapause among populations within a given latitude, and supported the existence of parental effects that influence offspring diapause. 4 Experiments with different host plants and high‐ vs. low‐quality artificial diets provided no evidence that the parental effects on offspring diapause were based on the quality of food consumed by parents. Diapause in C. rosaceana appears influenced by a hierarchy of cues, including temperature, photoperiod, current larval diet and parental environment.     

Historical and projected interactions between climate change and insect voltinism in a multivoltine species

Climate change can cause major changes to the dynamics of individual species and to those communities in which they interact. One effect of increasing temperatures is on insect voltinism, with the logical assumption that increases in surface temperatures would permit multivoltine species to increase the number of generations per year. Though insect development is primarily driven by temperature, most multivoltine insect species rely on photoperiodic cues, which do not change from year‐to‐year or in response to climate warming, to initiate diapause. Thus, the relationship between climate change and voltinism could be complex. We use a phenology model for grape berry moth, Paralobesia viteana (Clemens), which incorporates temperature‐dependent development and diapause termination, and photoperiod‐dependent diapause induction, to explore historical patterns in year‐to‐year voltinism fluctuations. We then extend this model to predict voltinism under varying scenarios of climate change to show the importance of both the quality and quantity of accumulated heat units. We also illustrate that increases in mean surface temperatures > 2 °C can have dramatic effects on insect voltinism by causing a shift in the ovipositional period that currently is subject to diapause‐inducing photoperiods.     

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 variation of reproductive diapause in the Drosophila auraria complex (Diptera: Drosophilidae)

ABSTRACT. In the Drosophila auraria complex (D. auraria Peng, D. biauraria Bock & Wheeler, D. triauraria Bock & Wheeler, D. quadraria Bock & Wheeler and D. subauraria Kimura), D. triauraria from the island of Yaku (30.3N in latitude) and D. quadraria from Taiwan (24.1N) had no or weak diapause, while strains of all the four species from the main islands of Japan had a firm reproductive diapause. These strains entered diapause at short daylengths and the critical daylength was longer and the incidence of diapause higher at a low rather than high temperature. In each species, the critical daylength was longer in strains from higher latitudes. Among sympatric populations of different species, D. subauraria always had the longest critical daylength, while D. auraria had the shortest one. The populations of D. biauraria and D. triauraria had longer critical daylengths than the sympatric populations of D. auraria , but this difference became smaller northwards. Since a longer critical daylength induces diapause earlier in autumn, D. subauraria would enter diapause earliest among these species, and D. auraria would do so latest. The late start of diapause in D. auraria may be related to its adaptation to domestic environments. On the other hand, the adaptive significance of the late start of diapause in northern populations of D. biauraria and D. triauraria is not clear.     

Geographic variation and genetic aspects of reproductive diapause in Drosophila triauraria and D. quadraria

ABSTRACT. Geographic variation and genetic aspect of reproductive diapause were studied in Drosophila triauraria Bock & Wheeler and D. quadraria Bock & Wheeler, in relation to their quantitative response to photoperiods. D. quadraria from the subtropical region had no photoperiodic diapause. In D. triauraria strains, diapause was induced under LD 10:14 or 12:12, but not under LD 14:10 or longer photoperiods. Their diapause was induced more effectively and maintained longer by LD 10:14 than by LD 12:12. The duration of diapause was longer in a northern strain, but the diapause inducing process varied little among different geographical strains. Diapause incidence was 50% or lower in F₁ hybrids between D. triauraria and D. quadraria and backcross progenies between F₁ and D. quadraria , and about 70% in backcross progenies between F₁ and D. triauraria under LD 10:14, but very low under LD 12:12. The lower incidence of diapause in these F; and backcross progenies is assumed to be due to the less efficient induction of diapause, since once diapause was induced in them, it was maintained for a long time, especially in the backcross progenies between F₁ and D. triauraria. These experiments suggest that diapause induction and maintenance are different physiological processes controlled by different genetic systems.     

Geographic variation in diapause induction and mode of diapause inheritance in Tetranychus pueraricola

Abstract:  Diapause induction and photoperiodic response curves were determined for 33 strains of Tetranychus pueraricola derived from kudzu vine at three constant temperatures (15, 18 and 20°C) under a short-day condition (10 : 14 h; light : dark). Females of all but one of the strains entered diapause at all three temperatures with little variation in diapause percentages among the strains. The exception was the southernmost strain, which was found to be a non-diapause (ND) strain. The critical photoperiod gradually decreased towards the south at a rate of about 1 h for each 5 degrees of latitude. The diapause strains (D1 and D2) exhibited 100% diapause, whereas the ND strain exhibited 0% diapause. By crossing these strains, we determined that 'non-diapause' was a dominant character over 'diapause' and the character was controlled by simple Mendelian inheritance. To clarify why the female progeny from the crosses between the D1 and ND strains did not segregate into the diapause and non-diapause phenotypes in a 1:1 ratio in the B₁ generation, round-robin crosses were carried out among the three strains. The results showed that the F₁ generation was reproductively compatible and showed high egg hatchability with a female-biased sex ratio. In the B₁ generation, the crosses between the D1 and ND strains and between the D1 and D2 strains exhibited extremely low egg hatchability and produced mostly female progeny, whereas offspring from the crosses between the D2 and ND strains showed more than 50% hatchability for B₁ eggs and a female-biased sex ratio. Thus, the absence of segregation observed in the crosses between the D1 and ND strains appears to be due to the severe hybrid breakdown that occurred in the B₁ generation.     

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.     

Diapause induction in adults of three Rhipicephalus appendiculatus stocks

Newly-moulted adults of three Rhipicephalus appendiculatus stocks, originating from Kenya, Zambia and Zimbabwe, were kept in different photoperiods (12h : 12h, 13h : 11h and 14h : 10h light : dark). The stock originating from Kenya showed almost no behavioural diapause in any of these day lengths, whereas virtually all individuals of the Zimbabwean stock entered a behavioural diapause irrespective of day length. Adults from the Eastern Province of Zambia, a transition zone between the multivoltine phenology in equatorial Africa and the univoltine in southern Africa, exhibited a photoperiod-dependent diapause response. The possible mechanisms of diapause regulation and their origin are discussed.     

Population consequences of diapause in a model system: the European corn borer

The diapause biology of the European corn borer (ECB), Ostrinia nubilalis (Hübn.), is described based on natural and controlled environment studies of feral and lab-reared ECB's in North Carolina (NC). The diapause response is described as a function of photophase (h of light/day) as well as a function of larval age (instar) at onset of diapause-inducing conditions. A critical photophase of 14.4 h and a critical mean larval instar of 3.3 is found in the lab studies and supported by three years of insectary studies. Seven years of black light trapping of ECB moths in Goldsboro, NC, revealed the likelihood of up to four moth flights/year.Information about the diapause biology of this insect is used to explain both the number of flights and the relative magnitude of the final moth flights. On average, the majority of ECB lineages pass through three generations/year with early maturing ECB's producing a significant and predictable fourth generation. The timing and magnitude of the fourth flight can be partly explained on the basis of the critical photophase and the timing and age structure of previous ECB generations. In most years, the fourth flight is smaller than the third due to the majority of the fourth generation's predisposition towards diapause. However, in at least one case (1977), the fourth flight was unusually large and could be predicted by slight temporal shifts in the previous three flights resulting in the majority of the fourth generation larvae averting diapause. The value of the ECB-diapause interaction as a model system for the explanation and prediction of dynamic phenological events is discussed.

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Résumé La diapause d'O. nubilalis Hubn. est décrite d'après des études en conditions naturelles et programmées de souches sauvages ou élevées au laboratoire en Caroline du Nord. La diapause est décrite comme une fonction de la photophase (heures de jour/24 heures) et du stade larvaire au début des conditions inductrices de la diapause. Une photophase critique de 14,4 h et un stade critique larvaire de 3,3 ont été établis au laboratoire après 3 ans d'études en insectarium. 7 ans de piégeage à la lumière noire à Goldsboro, ont montré la vraisemblance de l'existence de 4 vols par an. Les données sur la diapause de cet insecte sont utilisées pour expliquer tant le nombre de vols que l'importance relative des derniers vols. En moyenne, la majorité des lignées ont 3 générations par an, O. nubilalis précoces produisant une quatrième génération conséquente et prédictible. La data et l'importance du 4ème vol peuvent être partiellement expliquées d'après la photophase critique, et la data et la structure en âge des générations précédentes. La plupart des années, le 4ème vol est moins important que le 3ème par suite de la prédisposition à la diapause de la majorité de la 4ème génération. Cependant, dans un cas au moins, en 1977, le 4ème était anormalement important et pouvait être prédit par de faibles changements temporels dans les 3 précédents vols, détournant de la diapause la majorité des chenilles de 4ème génération. La discussion porte sur la valeur du modèle fourni par la diapause de O. nubilalis pour expliquer et prédire la dynamique des évéments phénologiques.

     

Genetic basis of photoperiodic control of summer and winter diapause in geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae Matsumura (Diptera: Chloropidae) is bivoltine in northern Japan but trivoltine in the southern part of the country. Larvae of the bivoltine strain (Akita, 39° N) develop without delay under L14:D10 but enter summer diapause in the mature larval stage under L15:D9. In contrast, larvae of the trivoltine strain (Aichi, 35° N) develop quickly under L15:D9 but enter summer diapause under L14:D10. The genetic basis of their different summer-diapause behavior was analyzed by reciprocal crossing between the two ecotypes and backcrossing to the hybrids. Although a polygenic system seems to be involved, there is a single gene on the X-chromosome exerting a major effect on summer diapause. The northern bivoltine ecotype has a longer critical photoperiod for the induction of winter diapause in the first larval stage and remains in diapause longer than the southern trivoltine ecotype. The differences in the critical photoperiod and the winter diapause duration between the two ecotypes seem to be controlled by a polygenic system but again a sex-linked or closely linked gene (or genes) plays a major role. The two geographic ecotypes differ significantly in summer- and winter-diapause traits, both of which are characterized by sex-linked inheritance. Despite this, no apparent sterility or inviability in hybrids between the ecotypes occurs.     

Maternal temperature has different effects on the photoperiodic response and duration of larval diapause in blow fly ( Calliphora vicina ) strains collected at two latitudes

Abstract .The blow fly Calliphora vicina Robineau-Desvoidy (Diptera: Calliphoridae) has a wide distribution across northern and temperate Europe. It has a facultative, maternally-induced larval diapause in response to short days. The photoperiodic response, measured at 15 and 20°C, of two populations was compared. A southern population (originating at 51° N) was sensitive to temperature at all daylengths; the incidence of diapause was greatly reduced at 20°C compared with 15°C. The photoperiodic response of a northern population (from 65° N) was sensitive to temperature only in long days; in short days (< 14 h of light) the response of this strain was identical at each temperature.
Variation in parental photoperiod and temperature were found to affect the duration of larval diapause, indicating a role for maternal effects in diapause intensity as well as incidence. However, the between-strain variation was greater than that within strains, indicating qualitative differences in diapause response. These differences may arise from the ecological conditions at the points of origin of the two strains. The northern strain from the harsher climate has a more intense diapause that follows a relatively temperature-insensitive photoperiodic response. In contrast, the southern strain has a shallow diapause and its photoperiodic response may be overridden by the experience of concurrent high temperature.