Arctic Insects Research Articles

Testing of arctic insect hemolymph as a secondary agent in applied cryopreservation

BACKGROUND: Cold hardiness of insects from extremely cold regions is based on a principle of natural cryoprotection, which is associated with physiological mechanisms provided by cryoprotectants. OBJECTIVE: Since arctic cold-hardy insects are producers of highly effective cryoprotectants, in this study, the hemolymph of Aporia crataegi L. and Upis ceramboides L. from an extremely cold area (Yakutia) was tested as a secondary component of cryoprotective agents (CPA) for cryopreservation (−80°C) of human peripheral blood lymphocytes and skin fibroblasts. MATERIALS AND METHODS: Lymphocytes and skin fibroblasts were treated with various combinations of DMSO and hemolymph extract and step-wise cooled to −80°C. Post-cryopreservation cell viability was assessed by vital staining and morphological appearance. RESULTS: Viability was higher when cells were frozen with a mixture containing DMSO and Upis ceramboides hemolymph compared to the cells frozen in DMSO, while cells frozen with DMSO and Aporia crataegi hemolymph did not survive. The fact that hemolymph of not every cold-resistant insect can be used as a secondary agent along with DMSO indicates that only a unique combination of hemolymph components and its compatibility with cells might result in a positive effect. CONCLUSION: Although the use of insect hemolymph as a complementary agent in applied cryopreservation is a problem in terms of practical application, such studies could initiate new trends in the search for the most successful hemolymph-like cryoprotectant systems.

Temporal regulation of temperature tolerances and gene expression in an arctic insect.

Terrestrial arthropods in the Arctic are exposed to highly variable temperatures that frequently reach cold and warm extremes. Yet, ecophysiological studies on arctic insects typically focus on the ability of species to tolerate low temperatures, whereas studies investigating physiological adaptations of species to periodically warm and variable temperatures are few. In this study, we investigated temporal changes in thermal tolerances and the transcriptome in the Greenlandic seed bug Nysius groenlandicus, collected in the field across different times and temperatures in Southern Greenland. We found that plastic changes in heat and cold tolerances occurred rapidly (within hours) and at a daily scale in the field, and that these changes are correlated with diurnal temperature variation. Using RNA sequencing, we provide molecular underpinnings of the rapid adjustments in thermal tolerance across ambient field temperatures and in the laboratory. We show that transcriptional responses are sensitive to daily temperature changes, and days characterized by high temperature variation induced markedly different expression patterns than thermally stable days. Further, genes associated with laboratory-induced heat responses, including expression of heat shock proteins and vitellogenins, were shared across laboratory and field experiments, but induced at time points associated with lower temperatures in the field. Cold stress responses were not manifested at the transcriptomic level.

ATG8 is conserved between Saccharomyces cerevisiae and psychrophilic, polar-collected fungi.

Autophagy is a conserved catabolic process by which eukaryotic cells respond to stress by targeting damaged or unneeded molecules or organelles for sequestration into specialized vesicles known as autophagosomes. Autophagosomes ultimately facilitate the digestion and recycling of their contents by fusing with the degradative organelle of the cell. Studies of the budding yeast Saccharomyces cerevisiae have revealed various types of stress that can regulate autophagy, including starvation and extreme temperatures. While autophagy has not yet been directly shown to confer the ability to survive extreme cold or freeze-thaw stress in yeast, upregulation of autophagy has been directly implicated in the ability of arctic insects to survive cold temperatures. We are interested in investigating the potential role of autophagy in polar habitat survival by cold-loving (psychrophilic) yeast like Mrakia blollopsis. To begin to examine the conservation of Atg machinery in polar-collected yeast, we focused on Atg8, a small, ubiquitin-like protein that plays an important role in autophagy. We report that Atg8 is conserved between S. cerevisiae and polar-collected yeast, using Atg8 from Mrakia blollopsis (strain TGK1-2) as an example. This study represents the first direct examination of autophagy machinery conservation across mesophilic and psychrophilic species of yeast.

Arctic insect emergence timing and composition differs across thaw ponds of varying morphology

ABSTRACT Freshwater ponds provide habitats for aquatic insects that emerge and subsidize consumers in terrestrial ecosystems. In the Arctic, insects provide an important seasonal source of energy to birds that breed and rear young on the tundra. The abundance and timing of insect emergence from arctic thaw ponds is poorly understood, but understanding these fluxes is important, given the role of insects in food webs and current rates of environmental change at high latitudes. We aimed to evaluate emerging insect communities from thaw ponds with different morphologies, identify environmental covariates influencing insect composition, and describe temporal changes in insect abundance. We collected environmental information and insects that emerged over two growing seasons and examined the phenology and taxonomic composition of insects arising from different pond classes: low centered polygon, small coalescent, large coalescent, and trough ponds. Our findings indicated no differences in the timing of total emergence across ponds of varying morphology. Community dissimilarity was primarily associated with center or margin habitat and variables that differed strongly among pond classes. These insects, which provide important provisions for various species of birds, are likely to experience changes in emergence phenology and composition due to ongoing, rapid warming in the region.

Life on the Edge: Ecological Genetics of a High Arctic Insect Species and Its Circumpolar Counterpart.

Arctic ecosystems are subjected to strong environmental constraints that prevent both the colonization and development of many organisms. In Svalbard, few aphid species have established permanent populations. These high arctic aphid species have developed peculiar life-history traits such as shortened life cycles and reduced dispersal capacities. Here, we present data on the distribution and population genetics of Acyrthosiphon svalbardicum in Spitsbergen, the main island of the Svalbard archipelago, and compared its genetic structure with that of its close relative Acyrthosiphon brevicorne, sampled in the top of Scandinavian mainland. We found that A. svalbardicum is common but heterogeneously distributed along the west coast of Spitsbergen. We recorded this species up to 79°12’, which constitutes the northernmost location for any aphid. Genetic structure examined using microsatellite markers showed more pronounced spatial differentiation in A. svalbardicum than in A. brevicorne populations, presumably due to reduced dispersal capacities in the former species. Although populations of A. brevicorne and A. svalbardicum were well-delineated at nuclear loci, they shared similar cytoplasmic DNA haplotypes as revealed by sequence analysis of two DNA barcodes. These results raise questions about whether these two taxa are different species, and the colonization sources and history of the Svalbard archipelago by A. svalbardicum.

Rapid induction of the heat hardening response in an Arctic insect.

The ability to cope with increasing and more variable temperatures, due to predicted climate changes, through plastic and/or evolutionary responses will be crucial for the persistence of Arctic species. Here, we investigate plasticity of heat tolerance of the Greenlandic seed bug Nysius groenlandicus, which inhabits areas with widely fluctuating temperatures. We test the heat tolerance and hardening capacity (plasticity) of N. groenlandicus using both static (heat knock down time, HKDT) and dynamic (critical thermal maximum, CTmax) assays. We find that N. groenlandicus is able to tolerate short-term exposure to temperatures up to almost 50°C and that it can quickly increase heat resistance following heat hardening. Furthermore, we find that this hardening response is reversible within hours after hardening. These findings contrast with common observations from temperate and tropical insects and suggest high thermal plasticity in some Arctic insects which enables them to cope with extreme temperature variability in their habitats.

Use of arctic insect hemolymph as a complimentary agent in applied cryopreservation

Use of arctic insect hemolymph as a complimentary agent in applied cryopreservation

Declining diversity and abundance of High Arctic fly assemblages over two decades of rapid climate warming

Insects are particularly vulnerable to rapid environmental changes, which are disproportionally affecting high latitudes. Increased temperature could influence insect species differentially and reshape assemblages over time. We quantified temporal assemblage turnover of Arctic Diptera (flies) in the Muscidae, one of the most diverse and abundant families of Arctic insects, using time series data from Zackenberg, north‐east Greenland. We measured temporal patterns of abundance, diversity, and composition of muscid assemblages in wet fen, mesic and arid heath habitats from yearly collections spanning 1996–2014 and tested their relationship to climate. A total of 18 385 individuals representing 16 species of muscid flies were identified. A significant decrease of 80% of total muscid abundance was observed during the study period. Species richness declined in each habitat type but this trend was not significant across habitats. The number of common and abundant species also decreased significantly over time across habitats revealing a temporal modification of species evenness. Significant temporal changes in composition observed in the wet fen and across habitats were mainly driven by a change in relative abundance of certain species rather than by species replacement. Shift in composition in each habitat and decline in muscid abundance across habitats were associated with summer temperature, which has significantly increased over the study period. However, relationships between temperature and muscid abundance at the species level were noticeable for a few species only. Significant directional change in composition was documented in the wet fen but no biotic homogenization across habitats was observed. As one of the few studies of species‐level changes in abundance, diversity and composition of an insect taxon in the Arctic over the past two decades, our study shows that habitat types may modulate insect species responses to recent climate change and that contrasting species responses can alter species assemblages within a few decades.

Late Tertiary origins of the Arctic beetle fauna

During the past 30 years, Quaternary insect paleontologists working in the Arctic have studied rare deposits of fossil insects that date back millions of years. Some of these fossils may be as old as the Late Miocene. The fossils have been preserved mostly in permafrost environments, and their state of preservation is often exceptional. The vast majority of identifiable beetle specimens appear to match modern, extant species. This morphological constancy through time appears to reflect constancy of physiological adaptations, as well. The latter aspect is demonstrated indirectly through the ecological compatibility of species found in the ancient fossil assemblages. While most of the species themselves appear to have remained constant, the biological communities in which they lived have shifted dramatically through time. Based on both insect and plant fossil data, Late Tertiary environments of the Arctic were substantially warmer than they are today, supporting the growth of coniferous forests, up to the shores of the ancient Arctic Ocean. By about 2 million years ago (my) or shortly thereafter, Arctic tundra communities came into existence in parts of Beringia. Arctic insect faunal diversity declined markedly with the onset of Quaternary cooling.

Overwintering adaptations in Arctic sawflies (Hymenoptera: Tenthredinidae) and their parasitoids: cold tolerance

AbstractAlthough the extreme winter conditions of Arctic habitats are considered to be important determinants of the faunal composition of Arctic regions, the overwintering biology of most Arctic insects is unknown. A significant proportion of the insect fauna at high latitudes are hymenopterans, yet little information is available on their overwintering strategies. In this study I examined the overwintering strategies of willow gall-forming and catkin-feeding nematine sawflies and their parasitoids. All sawfly species overwintered as prepupae, were freezing-tolerant, and survived exposure to −50 °C. Freezing at high subzero temperatures was initiated by ice nucleators associated with the posterior hind gut wall. Heterogeneity of overwintering habitats with respect to temperature was not a determinant of the overwintering success of these Arctic sawflies. Divergent overwintering mechanisms were evident in the sawfly parasitoid taxa. Endoparasitoid larvae, like their sawfly hosts, were freezing-tolerant. Freezing of immature, feeding endoparasitoid larvae occurred at the freezing point of the host prepupa and was a consequence of the inoculation of the endoparasitoids' fluid compartments by ice crystals growing in the host hemolymph. Peculiarities in the structure of the endoparasitoid larval gut suggest that the site of nucleation is across the gut wall. Outside their hosts, however, endoparasitoid larvae can supercool extensively, and their tolerance of extremely low temperatures is similar to that of their hosts. Overwintering strategies adopted by the ectoparasitoids were also diverse, with both freezing-tolerant and freezing-intolerant species present in the parasitoid community. Freezing-intolerant species could not survive winter temperatures in the field in the absence of an insulating layer of snow.

Seasonal Adaptations in Arctic Insects

Many insect species live in the arctic and show a wide range of adaptations to its extreme severity and seasonality. Long, cold winters are met, for example, by cold hardiness and choice of protected sites. Cold hardiness includes both widespread tolerance to freezing and extreme supercooling ability, as well as unusual responses in a few species, such as lack of typical cryoprotectants. Adaptations to short, cool summers include activity at low temperatures, selection of warm habitats and microhabitats, melanism and hairiness coupled with basking behaviour, and prolonged or abbreviated life cycles. Diapause ensures that many species emerge early in summer, with brief synchronized reproduction that maximizes the time for offspring development before winter returns. Some species overwinter in sites that thaw earliest in spring, even if they are relatively exposed in winter. Other adaptations respond to year-to-year variability: for example, prolonged diapause can provide insurance against unsuitable summers. All of these adaptations are co-ordinated. For example, cold hardiness relies on physiological and biochemical adaptations but also on habitat choice and timing. Because the adaptations are complex, predicted climatic warming probably will have unexpected effects. In particular, an increase in temperature that increases summer cloud when sea ice melts would likely reduce temperatures for insect development and activity, because sunshine provides critical warmth to insects and their microhabitats. Changes in moisture will also be important. Moreover, responses differ among species, depending especially on their microhabitats. The complexity of the responses of insects to arctic conditions reinforces the need for research that is sufficiently detailed.

Life on the edge: insect ecology in arctic environments.

The restricted Arctic insect fauna is usually explained by a lack of recolonization since the last glacial period, inadequate supply of suitable resources, or insufficient adaptation to such a harsh environment. These hypotheses and others that attempt to explain the latitudinal gradient of species distributions and abundance are reviewed. Arctic habitats available to insects are strongly heterogeneous, requiring a similarly diverse array of adaptive responses, characteristic of those species that have colonized and survived in such a stressful climate. Important adaptations in morphology (size, wings), behavior (activity patterns, thermoregulation), life cycles, and ecophysiology (cold hardiness, anaerobiosis, desiccation resistance) are discussed. The current focus of global climate change research on polar regions is identified, particularly the opportunity to study fundamental ecological processes and spatial dynamics in the relatively simple Arctic ecosystems.

Progressive restriction of host plant exploitation along a climatic gradient: the willow psyllid Cacopsylla groenlandica in Greenland

1. Host plant selection by the endemic willow psyllid Cacopsylla groenlandica was studied at eight sites in three locations along a N–S climatic gradient in west Greenland.2. C. groenlandica oviposited and developed on four willow species: Salix glauca, S. arctophila, S. uva‐ursi and S. herbacea. Development took place on female catkins of all species and on leaf shoots of S. glauca.3. At sites in southern Greenland, C. groenlandica fully utilized all four willow species. However, there was progressive reduction of host plant species and tissues exploited as latitude increased, such that towards the northern limit of its range the psyllid became highly specialized, feeding only on female catkins of S. glauca, despite the presence of alternative hosts. Where plant growth was tightly constrained within a restricted growing season, the demands of synchronizing with more than one species may have become too great, resulting in specialization on a single host.4. The probable mechanisms producing specialization were between‐species differences in host plant phenology, linked to variation in the growing season, set against the partially non‐overlapping geographical distribution patterns of the host willows.5. A small degree of phenological asynchrony between host plants can be advantageous for the insect. Under favourable conditions the reproductive season can be extended by using a series of host plants with staggered phenologies. Some ‘early’ hosts also create predator‐free space. However, as asynchrony increases, the advantages disappear as the insect becomes unable to exploit the full range of host options.6. The importance of host plant synchrony in the life histories of arctic insects is discussed.

Insect Cold-Hardiness: Insights from the Arctic

Cold-hardiness and related adaptations of insects in the Arctic correspond to characteristic climatic constraints. Some species are long-lived and are cold-hardy in several stages. In the Arctic, diapause and cold-hardiness are less likely to be linked than in temperate regions, because life-cycle timing depends as much on the need to coincide development with the short summer as on the need to resist winter cold. Winter habitats of many species are exposed rather than sheltered from cold so that development in spring can start earlier. Several features of cold-hardiness in arctic species differ from the characteristics of temperate species: these include very cold-hardy insects with low supercooling points that are not freezing tolerant; freezing-tolerant species that supercool considerably rather than freezing at relatively high subfreezing temperatures; mitochondrial degradation linked with the accumulation of cryoprotectants; and the possibly limited occurrence of thermal hysteresis proteins in winter. Several interesting relationships between cold-hardiness and water have been observed, including different types of dehydration. Winter mortality in arctic insects appear to be relatively low. Adaptations to cold in summer include retention of cold-hardiness, even freezing tolerance; selection of warm sites; and behaviour such as basking that allows elevated body temperatures. Studies especially on the high-arctic moth Gynaephora groenlandica show that various factors including cold-hardiness and other summer and winter constraints dictate the structure of energy budgets and the timing of life cycles. Future work should focus on the biological and climatic differences between arctic and other areas by addressing habitat conditions, life-cycle dynamics, and various aspects of cryoprotectant production at different times of year. Even in the Arctic cold-hardiness is complex and involves many simultaneous adaptations.Key words: cold-hardiness, insects, supercooling, freezing-tolerance, cryoprotectants, metabolism, energy budgets, life cycles, habitat selection

Corrigendum: A balanced life table for Gynaephora groenlandica (Lepidoptera: Lymantriidae), a long-lived high arctic insect, and implications for the stability of its populations

Corrigendum: A balanced life table for <i>Gynaephora groenlandica</i> (Lepidoptera: Lymantriidae), a long-lived high arctic insect, and implications for the stability of its populations

Respiratory Metabolism of sub-Antarctic insects from different habitats on Marion Island

Oxygen uptake of the foliage-dwelling larvae ofEmbryonopsis halticella Eaton (Lepidoptera: Yponomeutidae) and adults ofEctemnorhinus marioni Jeannel (Coleoptera: Curculionidae), the litter-dwelling larvae ofPringleophaga marioni (Lepidoptera: Tineidae) and the wrack-dwellingParactora dreuxi Seguy (Diptera: Helcomyzidae) was examined over the range of temperatures experienced by these insects in their microhabitats. With the exception of the kelp fly,P. dreuxi, Q10s and activation energies were generally lower than those found in temperate and Arctic insects, but were similar to values found in beetles from sub-Antarctic South Georgia Island. Q10 and activation energy of each species reflected the temperature regime found in its microhabitat. Activation energies of the Marion Island species were intermediate between those found in temperate and polar arthropods, but towards the polar end of the range. The hypothesis that insects are capable of showing respiratory adaptation to temperature is supported.

Arctic Insects as Indicators of Environmental Change

The great diversity of terrestrial arthropods in the Arctic suggests that these organisms are especially useful to monitor environmental change there, where warming as a result of climatic change is expected to be especially pronounced and where current conditions are limiting for many organisms. Based on existing information about arctic faunas and how they differ from temperate ones, this paper suggests several elements, including ratios and other quantitative indexes, that can be used for long-term evaluations of change. These elements include composition indexes, range limits, marker species, interspecific ratios, relationship shifts, phenological and physiological indicators, and key sites. Using such elements in a planned way would exploit the diversity of arctic insects and emphasize their importance in arctic systems.Key words: arctic arthropods, arctic insects, arctic fauna, climatic change, environmental changem, onitoring, indicator species, long-term research

Glycerol metabolism in a freeze-tolerant arctic insect: an in vivo 13C NMR study.

Freeze-tolerance in larvae of Gynaephora groenlandica is enhanced by the accumulation of glycerol in the winter. Since summer larvae remain freeze-tolerant despite the lack of glycerol, we investigated glycerol metabolism as a function of acclimation and body temperature using noninvasive 13C NMR spectroscopy. Major constituents of hemolymph isolated from cold- and warm-acclimated larvae were identified with the aid of standard NMR spectra and confirmed by TLC and GLC. Spectra obtained on live, warm-acclimated larvae showed the presence of lipids, glycogen, glucose, trehalose and amino acids. Similar spectra of cold-acclimated or previously frozen larvae showed the additional presence of glycerol. In vitro time-lapse 13C spectra of D-[1-13C]glucose added separately to hemolymph or extracted fat body tissue showed that glycerol is synthesized from glucose in the fat body tissue and distributed to the peripheral tissue via hemolymph. In vivo time-lapse 13C spectra of cold- and warm-acclimated larvae were obtained after injection with D-[1-13C]glucose to monitor the production of labeled metabolic intermediates and end-products. [13C]Glycerol was produced between -30 degrees C and 30 degrees C but accumulated only below 5 degrees C. Above 5 degrees C glycerol was degraded and the 13C label incorporated mainly into glycogen. The mechanism underlying temperature control of glycerol biosynthesis and degradation may provide a clue to the role of glycerol in enhancing freeze-tolerance in these insects.

NMR probes freeze tolerance in arctic insect

Some insects are able to survive subzero temperatures, either by tolerating freezing or by supercooling. Far north of the Arctic Circle, on Canada's Ellesmere Island and Devon Island, lives one of the small number of insect species that can tolerate freezing, the woolly-bear caterpillar, Gynaephora groenlandica . Scientists attribute its survival at least partially to its production of the cryoprotectant glycerol, which protects its protein and membrane structure and function. Knowledge of the physiological mechanisms underlying freeze tolerance is limited. Progress has been hampered in particular by lack of research methods permitting study of processes in intact, unmodified insects. Now, a team of scientists at the University of Notre Dame has employed noninvasive carbon-13 nuclear magnetic resonance spectroscopy, for the first time, to make in-vivo studies of the effects of temperature on the metabolism of a freeze-tolerant arctic insect. The team consists of graduate student Olga Kukal and pr...

A 7-year life cycle for two Chironomus species in arctic Alaskan tundra ponds (Diptera: Chironomidae)

The life cycles of two sibling Chironomus species inhabiting tundra ponds on the arctic coast of Alaska are interpreted from larval and adult data collected over 3 years. Emergence of adults was highly synchronous within each species, and the two emergence periods were always discrete. Larvae of the two species could not be separated morphologically and were treated as a single population through most of the life cycle. Analysis of larval size and development toward pupation indicated that seven cohorts coexist on nearly all sampling dates. A 7-year developmental period for each cohort is hypothesized and is supported by larval growth rates observed in the habitat and by the rates at which apparent cohorts progressed through the larval stages. Ten cohorts observed during the study period showed very similar schedules of growth and development, but cohort abundances varied considerably.This life cycle is among the longest reported for an arctic insect. It results from slow growth during an annual open-water season of about 90 days, though neither food nor temperature limitation could be definitely implicated in causing such slow growth. Coexistence of up to seven cohorts in each species stabilized Chironomus production and standing stock and may be important to benthic-feeding waterfowl which use these ponds.