Repeated cycles of chilling and warming effectively terminate prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis (Coleoptera: Curculionidae) requires one or more years to complete its life cycle, owing to prolonged larval diapause. To compare the effects of temperature cycles and total periods of chilling on the termination of prolonged diapause, larvae were subjected to different chilling (5 degrees C) and warming (20 degrees C) cycles ranging from 30 to 720 days, and all cycles were repeated until the sum of chilling and warming periods reached 720 days. The prolonged diapause of C. sikkimensis was more effectively terminated by repeated cycles of chilling and warming than by prolonging the continuous chilling period. However, extremely short temperature cycles were not highly effective in enhancing diapause termination, even when such cycles were repeated many times. To examine the role of warming periods on diapause termination, diapause larvae were subjected to a sequence of chilling (120 days at 5 degrees C) and warming (240 days at 20 degrees C) with a warming period (0-120 days at 20 degrees C) inserted in the chilling period. Diapause larvae that were not reactivated in the first chilling period required exposure to a certain period of warming before they were able to complete diapause development in the subsequent chilling. Thus, C. sikkimensis appears to spread its reactivation times over several years in response to seasonal temperature cycles.     

槲栎超寄生橡子内象甲幼虫的适应对策

栎属植物的橡子常常受到象甲的侵害, 对橡子存活产生影响, 但有关橡子对象甲幼虫适合度影响的研究尚未见报道。本研究旨在通过对槲栎Quercus aliena种子雨进程以及象甲幼虫逃逸过程的调查, 研究二者之间的进化适应关系。结果表明： 当年槲栎的种子产量为51.92±29.26粒/m², 虫蛀率达到42.4%。虫蛀的橡子中65%以上被象甲超寄生, 且显著大于完好的橡子。在种子下落过程中, 早期从橡子中逃逸的象甲幼虫要比后期逃逸的大。对某一超寄生的橡子而言, 首先逃逸出的象甲幼虫要比随后逃逸的幼虫大。另外, 从早期脱落的橡子中逃逸的幼虫明显小于后期脱落的橡子。象甲幼虫的干重随着橡子内象甲幼虫数量的增加而显著下降。象甲幼虫寄生数量与橡子的长径和短径呈现明显的正相关关系。尽管超寄生降低了象甲虫的适合度, 但雌性成体象甲产卵过程中偏好选择大橡子在一定程度上可以减少这种不利影响。因此, 小橡子可能更容易逃脱象甲的超寄生而获得较大的存活机会。     

Thermal response and reversibility of prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis undergoes prolonged larval diapause that is terminated by chilling and warming cycles. To examine the effects of warming temperatures and their duration on diapause termination, we exposed diapause larvae that had not been reactivated after chilling at 5 °C to 20 or 25 °C and chilled them again before incubation at 20 °C. With increasing warming duration at 20 °C, diapause termination after chilling increased and shorter chilling durations became effective. In contrast, few or no larvae warmed at 25 °C terminated diapause after chilling, irrespective of the warming duration. To investigate the effect of warming temperature on diapause intensity, larvae with diapause weakened by initial incubation at 20 °C after the first chilling were subsequently incubated at 15, 20, or 25 °C, then chilled at 5 °C before incubation at 20 °C. Diapause termination increased significantly after the larvae were treated at 15 or 20 °C but decreased significantly after they were treated at 25 °C. The intensification of prolonged diapause at 25 °C was reversed when the larvae were transferred to 20 °C. Diapause intensity in C. sikkimensis therefore decreases at 20 °C, increases at 25 °C, and can be reversed by alternately exposing diapause larvae to 20 and 25 °C. In C. sikkimensis, prolonged diapause does not always proceed in one direction, and its intensity fluctuates in response to ambient temperature conditions.     

Investigating the relationship between climate,stand age,and temporal trends in masting behavior of European forest trees

Masting—temporally variable seed production with high spatial synchrony—is a pervasive strategy in wind‐pollinated trees that is hypothesized to be vulnerable to climate change due to its correlation with variability in abiotic conditions. Recent work suggests that aging may also have strong effects on seed production patterns of trees, but this potential confounding factor has not been considered in previous times series analysis of climate change effects. Using a 54 year dataset for seven dominant species in 17 forests across Poland, we used the proportion of seed‐producing trees (PST) to contrast the predictions of the climate change and aging hypotheses in Abies alba, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Quercus petraea, and Quercus robur. Our results show that in all species, PST increased over time and that this change correlated most strongly with stand age, while the standardized precipitation–evapotranspiration index, a measure of drought, contributed to temporal trends in PST of F. sylvatica and Q. robur. Temporal variability of PST also increased over time in all species except P. sylvestris, while trends in temporal autocorrelation and among‐stand synchrony reflect species‐specific masting strategies. Our results suggest a pivotal role of plant ontogeny in driving not only the extent but also variability and synchrony of reproduction in temperate forest trees. In a time of increasing forest regrowth in Europe, we therefore call for increased attention to demographic effects such as aging on plant reproductive behavior, particularly in studies examining global change effects using long‐term time series data.     

Seasonal development and variation in abundance among four annual flight periods in a butterfly: a 20‐year study of the speckled wood (Pararge aegeria)

Insects typically spend the winter in a species‐specific diapause stage. The speckled wood butterfly, Pararge aegeria, is unique in having two alternative diapause stages, hibernating as larvae or pupae. In southern Sweden this creates a seasonal flight pattern with four annual adult flight periods: the first in May (pupal diapause), the second in June (larval diapause), and the third and fourth directly developing offspring generations in July and August, respectively. We address the raison d'être of the two diapause pathways by (1) outdoor rearing of cohorts, and (2) performing transect censuses throughout the season for 20 years. We contend that an early start of next season provides a benefit accruing to pupal diapause; conversely, a large proportion of the offspring from adults of the fourth flight peak are unable to reach the pupal stage before winter, providing a benefit accruing to larval winter diapause. The results obtained show that the two hibernation pathways are unlikely to be genetically distinct because of a strong overlap between the two offspring generations, and because sibling offspring from the third and fourth flight periods are likely to choose either of the two hibernation pathways, thereby resulting in a genetic mixing of the pathways. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 635–649.     

Effects of larval food shortage on diapause induction and adult traits in Taiwanese Monochamus alternatus alternatus

To confirm the facultative diapause of Monochamus alternatus alternatus Hope (Coleoptera: Cerambycidae) and to determine the relationships between available larval food resources, diapause, and adult traits, newly hatched larvae were inoculated singly on 98 Pinus densiflora Siebold & Zuccarini (Pinaceae) bolts and reared at 25 °C, 100% r.h., and L16:D8. Fifty adults emerged from them, between 70 and 126 days after larval inoculation. The remaining 48 bolts that did not produce adults were divided into two groups. One group was transferred to 10 °C, 100% r.h., and L8:D16, and returned 140–154 days later to the original conditions, resulting in adult emergence. The other group was maintained under the original conditions for a mean of 358 days. These bolts did not produce adults. Dissection revealed that development was arrested at final instar in pine bolts. The larvae developed into adults after being exposed to 10 °C, 100% r.h., and L8:D16 for 146 days. Consequently, this species has facultative diapause. Diapause incidence was estimated to be 0.42. Non‐linear model and one‐way ANOVA showed a positive correlation between adult body size and available food resources under conditions of food shortage, and no effects of diapause or available food resources on the ovariole number, respectively. When larvae were inoculated on 28 pine branch sections, the results were similar to those obtained from pine bolts and led to estimation of a low diapause incidence of 0.045. The combined data showed the inhibitory effect of food shortage on diapause induction. Diapause of M. a. alternatus, especially reduced diapause induction in response to environmental deterioration (food shortage), is discussed in relation to risk‐spreading.     

Expression of alternative developmental pathways in the cabbage butterfly,Pieris melete and their differences in life history traits

The seasonal life cycle of the cabbage butterfly, Pieris melete is complicated because there are three options for pupal development: summer diapause, winter diapause, and nondiapause. In the present study, we tested the influence of temperature, day length, and seasonality on the expression of alternative developmental pathways and compared the differences in life history traits between diapausing and directly developing individuals under laboratory and field conditions. The expression of developmental pathway strongly depended on temperature, day length, and seasonality. Low temperatures induced almost all individuals to enter diapause regardless of day length; relatively high temperatures combined with intermediate and longer day lengths resulted in most individuals developing without diapause in the laboratory. The field data revealed that the degree of phenotypic plasticity in relation to developmental pathway was much higher in autumn than in spring. Directly developing individuals showed shorter development times and higher growth rates than did diapausing individuals. The pupal and adult weights for both diapausing and directly developing individuals gradually decreased as rearing temperature increased, with the diapausing individuals being slightly heavier than the directly developing individuals at each temperature. Female body weight was slightly lower than male body weight. The proportional weight losses from pupa to adult were almost the same in diapausing individuals and in directly developing individuals, suggesting that diapause did not affect weight loss at metamorphosis. Our results highlight the importance of the expression of alternative developmental pathways, which not only synchronizes this butterfly's development and reproduction with the growth seasons of the host plants but also exhibits the bet‐hedging tactic against unpredictable risks due to a dynamic environment.     

Glucosinolate profile and oviposition behavior in relation to the susceptibilities of Brassicaceae to the cabbage seedpod weevil

Understanding how host‐plant characteristics affect behavioral and physiological responses of insect herbivores is of considerable importance in the development of resistant crop germplasm. Feeding, oviposition preference, larval development, and oviposition behavior of the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (= Ceutorhynchus assimilis Payk.) (Coleoptera: Curculionidae), were investigated on eight Brassicaceae species that differed in their glucosinolate profiles. The least preferred host plants for feeding and oviposition were the Sinapis alba L. lines while the Brassica carinata L. line was most preferred. Larval development occurred most rapidly on Brassica rapa L. and slowest on S. alba. Larval weight was highest on B. napus L. and lowest on S. alba. Total glucosinolate levels did not influence C. obstrictus larval growth or development; however high levels of specific glucosinolates such as p‐hydroxybenzyl and 3‐butenyl glucosinolate were associated with increased developmental time or reduced weight. The time required for oviposition behavioral events was measured on different host‐plant species: B. rapa, B. napus, B. napus×S. alba, B. tournefortii Gouan., B. juncea (L.) Czern, B. carinata, B. nigra (L.) Koch., and S. alba. The early steps in the sequence were completed faster on more susceptible host plants (B. carinata, B. napus, and B. rapa) than on relatively resistant ones (B. tournefortii and B. juncea). Females explored pods of B. nigra and S. alba, but oviposition occurred only rarely on these species. There was no significant difference in the location on the pod on which oviposition occurred among the different plant species. Mean eggs laid per female weevil were highest on the B. napus×S. alba hybrid and lowest on B. nigra and S. alba.     

Free amino acids of the haemolymph of the southwestern corn borer and the European corn borer in relation to their diapause

The titres of free amino acids present in the haemolymph of diapausing larvae of the southwestern corn borer, Diatraea grandiosella and the European corn borer, Ostrinia nubilalis, were examined. High titres of serine were found in the haemolymph of both species. Serine may serve as a storage form of compounds that are required for the synthesis of uric acid and other purines. The high titres of proline found in the haemolymph of O. nubilalis during the fall and winter may contribute to the freezing tolerance of this species. Alanine accumulated in the haemolymph of both species during the winter.     

Guild structure of ambrosia beetles attacking a deciduous oak tree Quercus serrata in relation to wood oldness and seasonality in three locations in the Central Japan

In order to determine factors influencing ambrosia beetle guilds on Quercus serrata, we investigated ambrosia beetles guilds by using Q. serrata bait logs in three locations in the Central Japan. Timing of cutting trees and timing of exposure were artificially controlled. Influences of location, timing of cutting, timing of exposure and wood oldness on species richness, abundances and guild structure were analyzed. Species richness and abundance peaked on bolts prepared in April–May, on bolts exposed in July, and on 2–3‐month‐old bolts. Eliminating greatest influences of location on abundance, results of hierarchical partitioning showed that timing of cutting trees had a strong influence on both species richness and abundance. LOC‐A (Aichi), in which Japanese oak wilt disease incidence occurred, showed the greatest species richness and the smallest value of Pielou's evenness. Abundance of the most major species was more than twice that of the second major species, which was a likely cause of the smallest evenness in LOC‐A. Trees killed by the Japanese oak wilt disease may have increased the abundance of the major species. On the contrary, in LOC‐C (Chichibu), alpha and beta diversity both given by Shannon index and Pielou's evenness were greatest among the three locations although species richness was smallest. High similarity between guilds in LOC‐A and LOC‐B (Chiba) was probably caused by similarity in vegetation. The location had the greatest effect on determining guild structure. Effect of timing of exposure was greater than timing of cutting. The effect of wood oldness was negligible. A hierarchical structure among the three factors was a likely cause of their relative importance determining guild structure.     

Tillage affects the activity‐density,absolute density,and feeding damage of the pea leaf weevil in spring pea

Conversion from conventional‐tillage (CT) to no‐tillage (NT) agriculture can affect pests and beneficial organisms in various ways. NT has been shown to reduce the relative abundance and feeding damage of pea leaf weevil (PLW), Sitona lineatus L. (Coleoptera: Curculionidae) in spring pea, especially during the early‐season colonization period in the Palouse region of northwest Idaho. Pitfall traps were used to quantify tillage effects on activity‐density of PLW in field experiments conducted during 2001 and 2002. As capture rate of pitfall traps for PLW might be influenced by effects of tillage treatment, two mark‐recapture studies were employed to compare trapping rates in NT and CT spring pea during 2003. Also in 2003, direct sampling was used to estimate PLW densities during the colonization period, and to assess PLW feeding damage on pea. PLW activity‐density was significantly lower in NT relative to CT during the early colonization period (May) of 2001 and 2002, and during the late colonization period (June) of 2002. Activity‐density was not different between treatments during the early emergence (July) or late emergence (August) periods in either year of the study. Trap capture rates did not differ between tillage systems in the mark‐recapture studies, suggesting that pitfall trapping provided unbiased estimates of PLW relative abundances. PLW absolute densities and feeding damage were significantly lower in NT than in CT. These results indicate that NT provides a pest suppression benefit in spring pea.     

Progeny of the maize weevil,Sitophilus zeamais,is affected by parental exposure to clove and cinnamon essential oils

The interest in and utilization of botanical insecticides, particularly essential oils, has become increasingly relevant to the control of insect pests. However, the potential ecotoxicological risks or flaws (including sublethal effects on the targeted pest generation and its subsequent progeny) of this pest control tool have been neglected frequently. Here, we evaluated the effects of sublethal exposure to clove, Syzygium aromaticum (L.) Merrill & Perry (Myrtaceae), and cinnamon, Cinnamomum verum J. Presl (Lauraceae), essential oils on adult (F0) maize weevils, Sitophilus zeamais Motschulsky (Coleoptera: Curculionidae), and the physiology (e.g., body mass, respirometry, and grain consumption) and population dynamics (e.g., daily emergence and sex ratio) of their progeny. Longevities of the parents were negatively affected by the essential oils in a concentration‐dependent manner. Parental sublethal exposure to clove oil (0.17 μl cm^?2) accelerated offspring emergence but delayed the emergence of females compared to males. Parents that were sublethally exposed to clove (0.17 μl cm^?2) or cinnamon (0.35 μl cm^?2) essential oils produced heavier offspring. Parental sublethal exposure to cinnamon essential oil accelerated offspring emergence (at 0.70 μl cm^?2), delayed female emergence (at 0.17 μl cm^?2), and enhanced grain consumption (at 0.35 and 0.70 μl cm^?2) of the progeny. Thus, our findings indicate that sublethal exposure to clove and cinnamon essential oils is capable of promoting transgenerational effects in S. zeamais that can negatively impact the control efficacy of such products.     

Alternative responses to rare selection events are differentially vulnerable to changes in the frequency,scope, and intensity of environmental extremes

Extreme weather events are becoming more frequent, severe, and/or widespread as a consequence of anthropogenic climate change. While the economic and ecological implications of these changes have received considerable attention, the role of evolutionary processes in determining organismal responses to these critical challenges is currently unknown. Here we develop a novel theoretical framework that explores how alternative pathways for adaptation to rare selection events can influence population‐level vulnerabilities to future changes in the frequency, scope, and intensity of environmental extremes. We begin by showing that different life histories and trait expression profiles can shift the balance between additive and multiplicative properties of fitness accumulation, favoring different evolutionary responses to identical environmental phenomena. We then demonstrate that these different adaptive outcomes lead to predictable differences in population‐level vulnerabilities to rapid increases in the frequency, intensity, or scope of extreme weather events. Specifically, we show that when the primary mode of fitness accumulation is additive, evolution favors ignoring environmental extremes and lineages become highly vulnerable to extinction if the frequency or scope of extreme weather events suddenly increases. Conversely, when fitness accumulates primarily multiplicatively, evolution favors bet‐hedging phenotypes that cope well with historical extremes and are instead vulnerable to sudden increases in extreme event intensity. Our findings address a critical gap in our understanding of the potential consequences of rare selection events and provide a relatively simple rubric for assessing the vulnerabilities of any population of interest to changes in a wide variety of extreme environmental phenomena.     

Activity of gut alkaline phosphatase,proteases and esterase in relation to diapause of pharate first instar larvae of the gypsy moth,Lymantria dispar

Two distinctly different patterns of gut enzyme activity were noted in relation to diapause in pharate first instar larvae of the gypsy moth, Lymantria dispar. Trypsin, chymotrypsin, elastase, aminopeptidase and esterase activities were low at the initiation of diapause and through the period of chilling needed to terminate diapause. At the completion of a 150 day chilling period, activity of each of these enzymes quickly increased when the pharate larvae were transferred to 25°C. By contrast, activity of alkaline phosphatase (ALP) increased rapidly at the onset of diapause, remained elevated throughout diapause, increased again during postdiapause, and then dropped at the time of hatching. In addition, zymogram patterns of ALP activity differed qualitatively in relation to diapause: several bands were detectable during the pre- and postdiapause periods, but only one band, a band of high mobility, was visible during diapause. The ALP isozyme present in diapausing pharate larvae had a pH optimum of 10.6. Diapause in the gypsy moth can be averted by application of an imidazole derivative, KK-42, and pharate larvae treated with KK-42 showed elevated protease and esterase activity, low ALP activity, and expressed ALP isozymes with low mobility. Thus the overall patterns of gut enzyme activity and the ALP zymogram in KK-42 treated individuals were similar to those observed in untreated individuals at the termination of diapause. Our results suggest a unique pattern of enzyme activity in the gut that is regulated by the diapause program. Arch. Insect Biochem. Physiol. 37:197–205, 1998. © 1998 Wiley-Liss, Inc.     

Effects of defoliation on the resistance and tolerance of rice,Oryza sativa,to root injury by the rice water weevil,Lissorhoptrus oryzophilus

Understanding plant‐mediated interactions in agricultural systems may facilitate the development of novel and improved management practices, which is important, as management of these insects is currently heavily reliant on insecticides. The fall armyworm, Spodoptera frugiperda (JE Smith) (Lepidoptera: Noctuidae, Prodeniini), is a sporadic pest of rice fields in the southern USA. In southwestern Louisiana, this defoliating insect typically attacks rice early in the growth season, before fields are flooded. Defoliation by fall armyworm larvae may trigger increased expression of plant defenses, which may result in increased resistance to subsequent herbivores. The rice water weevil (RWW), Lissorhoptrus oryzophilus Kuschel (Coleoptera: Curculionidae, Stenopelmini), enters rice fields as an adult both before and after flooding, but oviposition and larval infestation occur only after fields are flooded. RWW may be affected by changes in plant resistance caused by fall armyworm defoliation before flooding. The objectives of this study were to investigate the plant‐mediated effects of natural and artificial defoliation on population densities of RWW larvae after flooding and on the ability of rice plants to compensate for root injury by RWW larvae. In the 2015 season, fall armyworm defoliation before flooding resulted in reduced RWW densities after flooding. However, in 2016 no significant effects of fall armyworm defoliation on densities of RWW larvae were detected. Similarly, mechanical defoliation of rice before flooding did not affect RWW densities after flooding. Although lowest yields were observed in plots subjected to both root injury and defoliation, there was little evidence of a greater than additive reduction in yields from simultaneous injury. These results suggest a lack of plant‐mediated interactions among these two pests in rice in the southern USA.     

Reproduction and egg diapause of the oriental chinch bug,Cavelerius saccharivorus okajima (Heteroptera: Lygaeidae), in the subtropical winter season in relation to its wing polymorphism

Reproduction and egg diapause of the oriental chinch bug, Cavelerius saccharivorus, in the subtropical winter season were investigated in relation to its wing polymorphism. Macropterous females collected from the southern part of Okinawa Is. in the autumn season delayed their oviposition and were less fecund early in the adult life period, but survived much longer than brachypterous females collected from the same locality. The total fecundity was not significantly different between wing morphs. The diapause of eggs laid by brachypters tended to be terminated more easily at a high temperature than that of eggs laid by macropters. This indicated that the eggs laid by macropters were more intense in their diapause than those laid by brachypters. However, irrespective of the parental wing form, diapause showed considerable variation in its intensity within and between clutches.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in relation to temporal variation in predation risk by white sharks (Carcharodon carcharias) around a seal rookery in False Bay,South Africa

The marked differences in predation risk posed by white sharks (Carcarodon carcarias) at island rookeries of Cape fur seals (Arctocephalus pusillus pusillus) offer a quasi‐experimental design within a natural system for exploring how prey adjust their behavior in response to temporal variation in predation risk. Here we compare movement of juvenile and adult Cape fur seals at a high risk (Seal Island) and low risk (Egg Island) rookery. We further compare juveniles and adults at Seal Island in low and high risk seasons and at low and high risk times of day within those seasons. Adult fur seals at Seal Island avoided traversing the zone of high white shark predation risk during the high risk period (0700–0959) in the season of high risk (winter), but not during the low risk season (summer). By contrast, adult fur seals at Egg Island showed no temporal discretion in either season. Unlike juvenile fur seals at Egg Island, juveniles at Seal Island adjusted their temporal movement patterns to more closely mimic adult seal movement patterns. This suggests that exposure to predators is the primary driver of temporal adjustments to movement by prey species commuting from a central place.     

Demarcation of diapause development by cold and its relation to time-interval activation of TIME-ATPase in eggs of the silkworm, Bombyx mori

We investigated the mode of action of winter cold in the termination of diapause by investigating Time-Interval-Measuring Enzyme (TIME). First, we determined the period of cold required for the completion of diapause development. Synchronously developing egg batches of a pure strain (C108 Bombyx mori silkworm) were used to minimize variations in hatching time. Hatching occurred with only 18 days chilling at 5 degrees C when the incubation at 25 degrees C after the chilling was elongated. The 18-day period was much shorter than we expected; diapause in B. mori is known to terminate completely with about 100 days of chilling. Even in such a short period of chilling, no sporadic hatching occurred. Moreover, we determined that a temperature-insensitive stage, which we called "Neboke", followed the short cold-requiring stage. Thus, the stage of diapause development was demarcated from other stages of diapause. While the length of diapause development was elongated when chilling was delayed after oviposition, the Neboke stage length was invariant. Cold evidently exerts its effect only on diapause development. When TIME was purified from eggs and chilled in test tubes, a transitory burst of its ATPase activity occurred at a time equivalent to shortly before the completion of diapause development; this was an interval-timer activation. The mechanism by which cold activates TIME to measure the time interval may help explain in biochemical terms the insect's adaptation to its seasonal environments.