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Oxygen-isotope values of invertebrate cuticle preserved in lake sediments have been used in palaeoenvironmental reconstructions, generally with the assumption that fractionation of oxygen isotopes between cuticle and water (\(\upalpha_\textcuticle-\textH_2\textO\)) is independent of temperature. We cultured chironomid larvae in the laboratory with labelled oxygen-isotope water and across a range of closely controlled temperatures from 5 to 25 C in order to test the hypothesis that fractionation of oxygen isotopes between chironomid head capsules and water (\(\upalpha_\textchironomid-\textH_2\textO\)) is independent of temperature. Results indicate that the hypothesis can be rejected, and that \(\upalpha_\textchironomid-\textH_2\textO\) decreases with increasing temperature. The scatter in the data suggests that further experiments are needed to verify the relationship. However, these results indicate that temperature-dependence of \(\upalpha_\textchironomid-\textH_2\textO\) should be considered when chironomid δ18O is used as a paleoenvironmental proxy, especially in cases where data from chironomids are combined with oxygen-isotope values from other materials for which fractionation is temperature dependent, such as calcite, in order to derive reconstructions of past water temperature.
If \(\upalpha_\textchironomid-\textH_2\textO\) is indeed independent of temperature, it should be possible to reconstruct past values of \(\updelta^18 \textO_\textH_2\textO\) directly from measurements of δ18Ochironomid in fossil material, with obvious applications in palaeoclimate reconstruction. Wooller et al. (2004) noted similarities between chitin and aquatic cellulose in respect of oxygen-isotope fractionation from host water. Both have broadly similar fractionation factors (previous studies quote α values between about 1.023 and 1.028), suggesting similarity in the biochemical reactions by which oxygen derived from water is incorporated in the two polymers, despite the fact that cellulose is produced by plants and chitin by animals and fungi. Moreover, \(\upalpha_\textcellulose-\textH_2\textO\) is also generally regarded as temperature independent (Beuning et al. 1997). However, independence from temperature remains unproven for either biopolymer and results from culture experiments and field collections suggest there may in fact be temperature dependence of δ18O for Cladocera (Verbruggen et al. 2011; Schilder et al. 2015), and cellulose (Aucour et al. 1993). Clearly, temperature dependence needs either to be disproven or, if shown to be true, to be quantified in order for accurate reconstructions of \(\updelta^18 \textO_\textH_2\textO\) to be made.
Here, we report the results of a laboratory culture study of the influence of temperature on oxygen-isotope fractionation between host water and the chitinous head capsules of chironomid larvae using culture experiments under conditions of constant \(\updelta^18 \textO_\textH_2\textO\) and shared diet, leaving temperature as the variable to be investigated for its influence on δ18Ochironomid.
We reared larvae of the chironomid Chironomus riparius (Meigen 1804) from eggs (supplied by Huntington Life Sciences Ltd). This species was chosen based on its eggs being readily available. Eggs were reared in glass Erlenmeyer flasks, containing 2 L of bottled mineral water and 500 g of sand that was first combusted at 550 C for six hrs to eliminate any organic matter that might have provided extraneous food sources. The flasks were placed inside isothermal cabinets set at nominal temperatures of 5, 10, 15, 20 and 25 C. Duplicate experiments at each temperature (triplicate at 15 C) were conducted concurrently in the same cabinet to minimize temperature differences between the replicates. The flasks were kept in complete darkness to inhibit photosynthetic activity and loosely sealed with aluminium foil to minimize evaporation. Typically, every other day, each flask was provided with 1.5 mL of a suspension of finely ground Tetramin fish food flakes. This was prepared weekly by blending 4 g of fish food flakes with 1 L of water and kept refrigerated. The isotopic composition of food was not measured but was assumed to be constant. Rationing was adjusted according to water quality and larval behaviour, because decomposition of uneaten food can lead to increased microbial activity and reduced dissolved oxygen concentration, which may subsequently hinder larval development. Water quality was maintained through regular partial water replacements by siphoning off one litre of water from each flask weekly and replacing it with stock mineral water stored at the relevant temperature. This ensured satisfactory dissolved oxygen concentration and optimal environmental conditions for growth and development. Experiment duration varied depending on larval growth rates but in all cases, experiments were terminated once the majority of larvae had reached their final instar.