Physiology of diapause in the adult bark beetle, Ips acuminatus Gyll., studied in relation to cold hardiness

The ovarioles of the bark beetle, Ips acuminatus are telotrophic. Ovarian development is suppressed at an immature stage with primary germ cells present in the germaria. Lower oxygen consumption is found in beetles during autumn and early winter, and a substantial rise in mean respiration rates occurs in the beginning of January paralleled by a resumption of pre-vitellogenesis in all females maintained either at 3°C or out of doors after return to 21°C for 2 weeks. It is concluded that I. acuminatus enters faculative diapause soon after enclosure to the adult, and that diapause is terminated by mid-winter in beetles kept for 18 weeks at either 3°C or out of doors. The specimens remain thereafter in reproductive quiescence until ovarian development can proceed.Photic cues are neither involved in the elevation of mean respiration rates, nor needed to abolish the inhibition of ovarian maturation in beetles kept at 3°C or in those returned to 21°C. However, follicle formation in ovarioles is only seen in positive phototactic females reared during “long-day” conditions, suggesting a photoperiodic regulation of the later stages of vitellogenesis.Detectable amounts of ethylene glycol are found at the beginning of November in freezing-susceptible I. acuminatus hibernating in its galleries underneath bark of Scots pine (Pinus silvestris) at 3°C. The gradual catabolism of the cryoprotective solute at 3°C through December occurred at a time when individuals achieved the competence to resume ovarian maturation during 2 weeks at 21°C, but prior to the substantial rise in their mean respiration rates. However, resumption of ovarian development in spring had no effect on the capability of outdoor beetles to enhance their supercooling capacity when subjected to sub-zero temperatures. Since the ability to respond to temperature changes occurred in post-diapause I. acuminatus as well, the maintenance of prolonged cold hardiness in specimens could not be related to diapause itself. Apparently, the ability of beetles to resynthesize ethylene glycol when a detectable level is present in the organism remains unaltered during overwintering.     

Moulting reduces freeze susceptibility in the Antarctic mite Alaskozetes antarcticus (Michael)

Abstract.  The effect of moulting on the cold hardiness of the oribatid mite Alaskozetes antarcticus (Michael) is investigated. Non moulting animals show clear seasonal patterns of cold hardiness with high supercooling points (SCPs) at the peak of summer and an increasing proportion of low SCPs with declining environmental temperatures. By contrast, both field-fresh and laboratory acclimated (5 °C) mites in the moult state are consistently found to have low SCPs regardless of environmental temperature.     

Seasonal changes in supercooling capacity and major cryoprotectants of overwintering Asian longhorned beetle (Anoplophora glabripennis) larvae

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Change in overwintering mechanism of the Cucujid beetle, Cucujus clavipes

Overwintering larvae of the Cucujid beetle, Cucujus clavipes, were freeze tolerant, able to survive the freezing of their extracellular body fluids, during the winter of 1978–1979. These larvae had high levels of polyols (glycerol and sorbitol), thermal hysteresis proteins and haemolymph ice nucleators that prevented extensive supercooling (the supercooling points of the larvae were ? 10°C), thus preventing lethal intracellular ice formation. In contrast, C. clavipes larvae were freeze suspectible, died if frozen, during the winter of 1982–1983, but supercooled to ～ ? 30°C. The absence of the ice nucleators in the 1982–1983 larvae, obviously essential in the now freeze-susceptible insects, was the major detected difference in the larvae from the 2 years. However, experiments in which the larvae were artifically seeded at ? 10°C (the temperature at which the natural haemolymph ice nucleators produced spontaneous nucleation in the 1978–1979 freeze tolerant larvae) demonstrated that the absence of the ice nucleators was not the critical factor, or at least not the only critical factor, responsible for the loss of freeze tolerance in the 1982–1983 larvae. The lower lethal temperatures for the larvae were approximately the same during the 2 winters in spite of the change in overwintering strategy.     

Geographic variation in cold hardiness of eggs and neonate larvae of the yellow-spotted longicorn beetle Psacothea hilaris

Cold hardiness of eggs and neonate larvae of the yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe) was examined using six geographical populations in Japan. Particular attention was paid to cold hardiness of eggs and neonate larvae of the subtropical population (Ishigaki), because the east Japan populations are considered to have been introduced from a subtropical area, and the overwintering stage in the east Japan populations is incidentally shifted from the original mature larval stage to the egg or neonate larval stages. When the eggs were exposed to low temperatures for 1 h, the decrease in hatchability became significant at –12°C in the southernmost two populations (Ishigaki and Naze), and at –16°C in the northern populations. After 1 h exposure to –20°C, few eggs could hatch in the Ishigaki population, whereas 27–55% of the eggs survived in the northern populations. Pre-chilling of the eggs at 10°C for 10 days enhanced the cold hardiness in all populations. This effect was particularly distinct in the subtropical population; the eggs of the Ishigaki population became as cold hardy as those of the northern populations after acclimation. These results suggest that the subtropical population is capable of establishing itself in east Japan, where the winter is cold.     

Seasonal changes of glycogen/trehalose contents,supercooling points and survival rate in mature larvae of the overwintering soybean pod borer Leguminivora glycinivorella

The mature larvae of the soybean pod borer Leguminivora glycinivorella, spend over 9 months (October-next August) in the inactive state until pupation down to 3 cm below the surface in soil. Trehalose content of inactive larvae increases in early winter, attaining a maximum (ca 30 mg/g), and decreases in spring, with a concomitant decrease and increase of glycogen. The median supercooling points seasonally change from ?19.8°C (October) to ?25.0°C (February), and to ?17.0°C (June). The lower supercooling points in winter are in part due to the absence of unusually high values (> ?18°C). The increase in trehalose does not seem to be effective in depressing the supercooling points. The larvae are freeze-intolerant, but ambient temperatures in outdoor conditions are always above the supercooling points. The survival rates are very high throughout the inactive period.     

Metabolism and cold tolerance of Chinese white pine beetle Dendroctonus armandi (Coleoptera: Curculionidae: Scolytinae) during the overwintering period

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Cold-hardiness of the arctic beetle, Pytho americanus Kirby Coleoptera,Pythidae (Salpingidae)

The arctic beetle, Pytho americanus Kirby, is frost tolerant in both larval and adult stages. This is the first demonstration that an insect can tolerate freezing in more than one life stage, a situation which would be congruous with its northern distribution and allow it to spread its life cycle over a number of growing seasons. The main biochemical correlates during the cold hardening process of low temperature acclimation are increasing glycerol and decreasing glycogen concentrations. Glycerol is the only polyol to be synthesized during acclimation, and it accumulates to a maximum of 8.2 and 12.2% of the fresh body weight in larvae and adults respectively. This coincides with the peak of frost tolerance. In addition to its normally assumed roles in cryoprotection it is suggested that glycerol may further serve to minimize dehydration in the overwintering insect by increasing the level of ‘bound’ water. Evidence is presented that indicates that glycerol is synthesized mainly from carbohydrate reserves, especially glycogen, but it does not rule out the possibility that a proportion of free glycerol comes from glyceride sources.P. americanus larvae and adults have low supercooling potential and maintain their supercooling points in the region of ?4° to ?8°C. It is hypothesized that these elevated supercooling points are a result of the presence in the haemolymph of nucleating agents which ensure ice formation at high sub-zero temperatures. It is believed that this beetle overwinters in a frozen state within its microhabitat, which is under bark of fallen spruce which is, in turn, covered by an insulating blanket of snow. The advantages of this overwintering strategy are discussed.     

外源海藻糖对花绒寄甲成虫存活和耐寒性的影响

【目的】探讨取食不同浓度外源海藻糖对室内饲养的花绒寄甲 Dastarcus helophoroides 成虫存活和耐寒性的影响。【方法】在室内分别用含3%, 6%和9%海藻糖的半人工饲料饲养花绒寄甲成虫,以取食不含海藻糖的半人工饲料的花绒寄甲成虫为对照组,统计饲养10周后的存活率,测定未经低温处理和10℃低温处理3 d的成虫过冷却点和含水量。【结果】取食含6%海藻糖的半人工饲料的花绒寄甲成虫存活率最高,为86.67%。不管是否经低温处理,取食含9%海藻糖的半人工饲料的成虫与取食含3%和6%海藻糖的半人工饲料的成虫以及不含海藻糖的半人工饲料的成虫(对照)相比,其过冷却点均最低,其中未经低温处理的成虫过冷却点为-19.30℃,而经低温处理的成虫过冷却点为-21.60℃。低温处理对取食不含海藻糖的半人工饲料的成虫的含水量有显著影响,而对取食含海藻糖的半人工饲料的成虫含水量无显著影响。【结论】外源海藻糖对花绒寄甲成虫的存活和过冷却点有显著影响,可以利用外源海藻糖提高室内饲养花绒寄甲成虫的存活率和耐寒性。     

Thermal stability properties of an antifreeze protein from the desert beetle Microdera punctipennis

An insect antifreeze protein gene Mpafp698 was cloned by the RT-PCR approach from the desert beetle Microdera punctipennis. The gene was constructed and heterogeneously expressed in Escherichia coli as fusion proteins, His-MpAFP698, glutathione S-transferase (GST)-MpAFP698, and maltose-binding protein (MBP)-MpAFP698. The thermostability and thermal hysteresis activity of these proteins were determined, with the aim of elucidating the biological characteristics of this protein. The approximate thermal hysteresis (TH) value of the purified His-MpAFP698 was 0.37 °C at 0.84 mg/ml, and maintained approximately 95.7% of the TH activity at 100 °C for 5 min. Furthermore, heat incubation showed that MBP-MpAFP698 was 10 °C more thermostable than MBP protein, indicating that MpAFP698 could, to some extent, improve the thermal stability of the fused partner MBP protein. This study suggests that MpAFP698 has a high thermal stability and could be used to improve the thermal stability of the less stable proteins by producing fusion proteins, which could be used for biotechnological purposes.     

Freeze tolerance in larvae of the winter-active Diamesa mendotae Muttkowski (Diptera: Chironomidae): a contrast to adult strategy for survival at low temperatures

The winter-active Diamesa mendotae Muttkowski (Diptera: Chironomidae) is freeze intolerant in the adult stage with a low mean supercooling point (SCP) of ~−20 °C. However, cold-hardiness strategies for immatures of this species are unknown. In this study, we measured SCP values for D. mendotae larvae, pupae and adults using surface-contact thermometry. In addition, the lower lethal temperature (LLT) was determined for the larval stage. The mean SCPs for larvae (−7.4 °C) and pupae (−9.1 °C) were relatively high compared to adults (−19.7 °C). Our results indicate that the larvae of D. mendotae are freeze tolerant with a LLT₉₉ (−25.4 °C), ~−10 °C lower than their minimum SCP (−15.6 °C). Freeze tolerance in these larvae may be a strategy to provide protection from short-term exposures to ice crystals or to permit diapause within frozen substrates. The change in cold-hardiness strategy from freeze tolerant to freeze intolerant between the larval and adult stages of this species is likely a result of the different habitats occupied by these two life stages.     

Experimental studies on the cold tolerance of Alaskozetes antarcticus

The cold tolerance mechanism of the Antarctic terrestrial mite Alaskozetes antarcticus (Michael) was investigated in cultured animals. Freezing is fatal in this species and winter survival occurs by means of supercooling, which is enhanced by the presence of glycerol in the body. There is an inverse, linear relationship between the concentration of glycerol and the supercooling point, which may be as low as ?30°C. Feeding detracts from supercooling ability by providing ice nucleators in the gut which initiate freezing at relatively high sub-zero temperatures. Experiments on the effects of various environmental factors showed that low temperature acclimation gave rise to increased glycerol concentrations and suppressed feeding, while desiccation also stimulated glycerol production. Photoperiod had no effect on cold tolerance in this species. The juvenile instars of A. antarcticus were found to possess a greater degree of low temperature tolerance than adults.     

Photoperiodic and thermal regulation of antifreeze protein levels in the beetle Dendroides canadensis

The importance of photoperiod, temperature and their interaction in controlling the seasonal pattern of haemolymph antifreeze protein levels in larvae of the beetle Dendroides canadensis was investigated. A complete photoperiodic response curve for antifreeze protein production was generated at 20°C with larvae collected in early fall. Individuals exposed to a 10-h photoperiod or less, including constant darkness, had significantly elevated antifreeze levels over those maintained in an 11-h photoperiod or more, including constant light. The critical daylength resulting in 50% population response lies between LD 11:13 and LD 10:14. This photoperiodic response was masked at sufficiently low (threshold between 15 and 10°C) and high (threshold between 25 and 30°C) temperatures. Partial photoperiodic response curves (at 17 and 25°C) obtained within this specified temperature range indicate that the position of the critical photoperiod (between 10 and 11 h) is stable while the amplitude of the response curve is temperature dependent.Experiments investigating the mechanisms controlling the spring depletion of protein antifreeze levels suggest that both photoperiod and temperature are important.The dominant response of photoperiod in the fall along with the modifying effects of temperature are considered to provide the necessary precision to assure adequate cold tolerance early in the fall and the flexibility to protect the species from yearly variation in weather conditions.     

Influence of sex,maturity and host substances on pheromones in the guts of the bark beetles, Ips paraconfusus and Dendroctonus brevicomis

Pheromones and metabolites of host (ponderosa pine) compounds were found in association with the hindgut of both naturally fed and of non-fed, host vapour-exposed bark beetles, Ips paraconfusus and Dendroctonus brevicomis. Much smaller amounts were found in the corresponding heads and mid guts. Sex-specific differences in content of pheromones were observed as in earlier studies. Exposure of I. paraconfusus to vapours of a pheromone component, ipsenol and other monoterpene alcohols resulted in their accumulation in the hindgut but relatively very low amounts in the head. The possible sites of pheromone biosynthesis are discussed. Exposure of male I. paraconfusus to vapours of host compounds, myrcene and α-pinene, revealed that immature adults do not produce the pheromone components, ipsenol and ipsdienol, as mature adults do while both immature and mature sexes produced another pheromone component, cis-verbenol, as well as trans-verbenol and myrtenol. Immature D. brevicomis adults did not contain pheromones until their exposure to vapours of (?)-α-pinene which caused production of trans-verbenol but only about 10% that of mature adults treated similarly. Verbenone, a male-produced inhibitory pheromone of D. brevicomis, apparently was not synthesized from (?)-α-pinene in females nor was its synthesis in males enhanced by exposure to this host compound.     

Winter survival of Harmonia axyridis in The Netherlands

Since the establishment of Harmonia axyridis in Europe, populations of native ladybirds have decreased. Overwintering survival is one of the aspects of the biology of H. axyridis that may contribute to its firm establishment in and invasion of a new area. In this study winter survival of five wild H. axyridis populations was assessed under natural and semi-natural conditions, with a focus on the potential influence of location and orientation on winter survival.Overwintering survival of H. axyridis in the Netherlands is high: 70.8-88.2%. When overwintering at one central site, populations sampled at five locations showed statistically significant different mortality rates. Furthermore, winter survival of H. axyridis at the sample sites was higher when beetles were hibernating at the southwestern sides of buildings, where most aggregations of ladybirds were found. Survival was higher at sheltered sites compared to exposed sites.Harmonia axyridis has a comparable or higher overwintering survival than most common native ladybird species. A high overwintering survival results in a large post-hibernation population in spring, leading to a rapid population build-up. Thus, the high winter survival probably contributes to the success of the exotic H. axyridis.     

Sterols and fatty acids of an antarctic sea ice diatom, Stauroneis amphioxys

Unialgal clonal cultures of the diatom Stauroneis amphioxys Gregory, isolated from sea ice of the Indian Ocean sector ofthe Southern Ocean, were grown at 3° and 20°. The relative abundances offatty acids, sterols and phytol for the two cultures are comparable. The two sterols observed [24-methylcholesta-5,22E-dien-3β-ol (79%) and cholesterol (21%)] did not vary with culture temperature. The-major fatty acid composition is typical ofmost diatoms. A pronounced change of ratio with temperature occurred with the pair 16:4 Δ6, 9, 12, 15 and 16:3 Δ6, 9, 12 followed by 18:4 Δ6, 9, 12, 15:18:3 Δ9,12,15 and 20:5 Δ5,8,11,14,17:20:4 Δ8,11,14,17; thus the relative abundances of 16:4, 18:4, 20:5 and 22:6 increase at the lower growth temperature. The total amounts of unsaturated acids do not change with temperature suggesting an effect on the final desaturase step. No cryoprotective role for such changes in lipid composition was inferred.     

Seasonal patterns of the thermal response in relation to sugar and polyol accumulation in overwintering adults of elm leaf beetle, Xanthogaleruca luteola (Coleoptera: Chrysomelidae)

Elm leaf beetle, Xanthogaleruca luteola (Muller), is one of the key pests of elm trees that survive winter in reproductive diapause in sheltered locations. Overwintering adults of the elm leaf beetle showed a complex sugar/polyol cryoprotectant system. The major components of the multiple systems were glucose, myo-inositol and trehalose. In this study, we investigated the seasonal profile of low molecular weight compounds and glycogen in natural population and also in response to thermal constant regimes (5 and 15 °C). Among these components, a remarkable seasonal pattern of accumulation/depletion was observed in myo-inositol over the course of hibernation with the development of diapause progress. Incubating at 5 °C only elicited a strong response in myo-inositol synthesis during diapause. It suggests that the elm leaf beetle accumulates myo-inositol not only in relation to entering diapause but also in response to low temperatures and their interactions. The laboratory acclimation experiments showed that adults exposed to 15 °C had no chance for accumulation of low molecular weight carbohydrate even during diapause. The results of this study illustrated that overwintering adults of elm leaf beetle produce myo-inositol as the primary substance which plays a specific role in some biochemical adjustments in overwintering adults of X. luteola.     

Cold tolerance of first-instar nymphs of the Australian plague locust, Chortoicetes terminifera

The cold tolerance of first-instar nymphs of the Australian plague locust, Chortoicetes terminifera, was examined using measures of total body water content, supercooling point and mortality for a range of sub-zero temperature exposure regimes. The supercooling points for starved and fed nymphs were −13.1 ± 0.9 and −12.6 ± 1.6 °C, and freezing caused complete mortality. Above these temperatures, nymphs were cold tolerant to different degrees based on whether they were starved or given access to food and water for 24 h prior to exposure. The rate of cooling also had a significant effect on mortality. Very rapid cooling to −7 °C caused 84 and 87% mortality for starved and fed nymphs respectively, but this significantly decreased for starved nymphs if temperature declined by more ecologically realistic rates of 0.5 and 0.1 °C min⁻¹. These results are indicative of a rapid cold hardening response and are discussed in terms of the likely effects of cold nights and frost on first-instar nymphal survival in the field.