Freeze-tolerant organisms survive internal ice formation; however, the adaptations to repeated freeze-thaw cycles are often not well investigated. Here we report how three geographically different populations of Enchytraeus albidus (Germany, Iceland and Svalbard) respond to three temperature treatments - constant thawed (0°C), constant freezing (-5°C) and fluctuating temperature (0 to -5°C) - over a period of 42 days. Survival varied between treatments and populations such that enchytraeids from arctic locations had a higher survival following prolonged freeze periods compared with temperate populations. However, enchytraeids from temperate locations had the same survival rate as arctic populations when exposed to repeated freeze-thaw events. Across all populations, metabolic rate decreased markedly in frozen animals (-5°C) compared with thawed controls (0°C). This decrease is likely due to the lower temperature of frozen animals, but also to the transition to the frozen state per se. Animals exposed to repeated freeze-thaw events had an intermediate metabolic rate and freeze-thaw events were not associated with pronounced excess energetic costs. Overwintering under either condition was not associated with a decrease in lipid content; however, during exposure to constant freezing and repeated freeze-thaw events there was a noticeable decrease in carbohydrate stores over time. Thus, animals exposed to constant freezing showed a decrease in glycogen stores, while both glucose and glycogen content decreased over time when the organisms were exposed to repeated freezing. The results therefore suggest that carbohydrate resources are important as a fuel for E. albidus during freezing whereas lipid resources are of marginal importance.
As potent ecosystem engineers, non-native earthworms are altering the fundamental structure and function of previously earthworm-free cold-temperate hardwood forests in North America. Discarded earthworms used for fishing bait has been presumed to be an important vector for the continued spread of non-native earthworms because epicentres of invasion often include boat landings, lakeshores and roads. However, controversy has remained about the overall importance of human-mediated spread vs. natural expansion of established earthworm populations. In this issue of Molecular Ecology, Cameron et al. explore the continued role of humans in dispersing non-native earthworms.
Comment On: Mol Ecol. 2008 Mar;17(5):1189-9718179429
Arctic plant growth is predominantly nitrogen (N) limited. This limitation is generally attributed to slow soil microbial processes due to low temperatures. Here, we show that arctic plant-soil N cycling is also substantially constrained by the lack of larger detritivores (earthworms) able to mineralize and physically translocate litter and soil organic matter. These new functions provided by earthworms increased shrub and grass N concentration in our common garden experiment. Earthworm activity also increased either the height or number of floral shoots, while enhancing fine root production and vegetation greenness in heath and meadow communities to a level that exceeded the inherent differences between these two common arctic plant communities. Moreover, these worming effects on plant N and greening exceeded reported effects of warming, herbivory and nutrient addition, suggesting that human spreading of earthworms may lead to substantial changes in the structure and function of arctic ecosystems.
By means of a literature survey, earthworm species of significant relevance for soil functions in different biogeographical regions of Europe (Atlantic, Boreal, Mediterranean) were identified. These focal earthworm species, defined here according to the EFSA Guidance Document on the environmental risk assessment (ERA) of genetically modified plants, are typical for arable soils under crop rotations with maize and/or potatoes within the three regions represented by Ireland, Sweden and Spain, respectively. Focal earthworm species were selected following a matrix of four steps: Identification of functional groups, categorization of non-target species, ranking species on ecological criteria, and final selection of focal species. They are recommended as appropriate non-target organisms to assess environmental risks of genetically modified (GM) crops; in this case maize and potatoes. In total, 44 literature sources on earthworms in arable cropping systems including maize or potato from Ireland, Sweden and Spain were collected, which present information on species diversity, individual density and specific relevance for soil functions. By means of condensed literature data, those species were identified which (i) play an important functional role in respective soil systems, (ii) are well adapted to the biogeographical regions, (iii) are expected to occur in high abundances under cultivation of maize or potato and (iv) fulfill the requirements for an ERA test system based on life-history traits. First, primary and secondary decomposers were identified as functional groups being exposed to the GM crops. In a second step, anecic and endogeic species were categorized as potential species. In step three, eight anecic and endogeic earthworm species belonging to the family Lumbricidae were ranked as relevant species: Aporrectodea caliginosa, Aporrectodea rosea, Aporrectodea longa, Allolobophora chlorotica, Lumbricus terrestris, Lumbricus friendi, Octodrilus complanatus and Octolasion cyaneum. Five out of these eight species are relevant for each biogeographical region with an overlap in the species. Finally, the earthworm species Ap. caliginosa (endogeic, secondary decomposer) and L. terrestris (anecic, primary decomposer) were selected as focal species. In the Mediterranean region L. terrestris may be substituted by the more relevant anecic species L. friendi. The selected focal species are recommended to be included in a standardized laboratory ERA test system based on life-history traits.