The quantity and quality of the haemoglobin (Hb) of Daphnia magna is related to oxygen partial pressure in the water. Both the dynamics of hypoxia-induced Hb gene transcription, as well as Hb properties in animals incubated long-term at hyperoxia, normoxia and hypoxia, were investigated. Examination of Hb gene (dhb1-dhb3) transcription showed the expression of dhb2 and especially dhb3 to increase markedly approximately one hour after the onset of hypoxia, whereas dhb1 was expressed more or less constitutively. At an incubation close to anoxia, an onset of dhb3 transcription was found already after two minutes. In long-term incubated animals, concentration and oxygen affinity of Hb were lower at higher oxygen partial pressures. With decreasing oxygen availability, the subunit composition of Hb macromolecules changed. The share of the dhb2-encoded subunit, DHbF, increased already during moderate hypoxia. The increase of dhb3 mRNA (encoding DHbC) may be related to a transient increase of DHbC in the first days of hypoxia and/or to an additional coding of dhb3 for DHbD. The rise of DHbD, and particularly DHbA, only at severe hypoxia coincided with the increase of Hb oxygen affinity. The dhb1-encoded subunits DHbB and DHbE showed either a relatively moderate increase or even a decrease in concentration at hypoxia. In small animals with restricted homeostasis capabilities such as Daphnia, adaptation of the protein equipment seems to be a more effective strategy than allosteric modulator control.
1. In a combined field and laboratory study, seasonal relationships between water temperature and oxygen content, genetic structure (composition of MultiLocus Genotypes, MLGs) of a Daphnia assemblage (D. galeata-hyalina hybrid species complex), and the physiological properties of clones of frequent MLGs were studied. In accordance with the oxygen-limited thermal tolerance hypothesis, essential physiological variables of oxygen transport and supply were measured within the tolerable temperature range. 2. A few MLGs (types T1-T4) were frequent during early spring and late autumn at surface temperatures below 10°C. Clones of T1-T4 showed a low tolerance towards higher temperatures (above 20°C) and a high phenotypic plasticity under thermal acclimation in comparison to clones derived from frequent MLGs from later seasons, and stored highmedium quantities of carbohydrates at 12 and 18°C. 3. Another MLG (T6) succeeded the MLGs T1-T4. T6 was frequent over most of the year at temperatures above 10°C and below 20°C. A clone derived from T6 exhibited a high tolerance towards warm temperatures and a more restricted phenotypic plasticity. It stored high-medium quantities of carbohydrates at 12, 18 and 24°C and showed a high capacity for acclimatory adjustments based on haemoglobin expression. 4. During the summer period at temperatures ‡20°C, the MLG T6 was found mainly near to the thermocline, where temperature and oxygen content were distinctly lower, and to a lesser extent in surface water. At the surface, another MLG (T19) was predominant during this period. A clone of this MLG showed a very high tolerance towards warm temperatures, minimal phenotypic plasticity, low carbohydrate stores and a high capacity for circulatory adjustments to improve oxygen transport at higher temperatures. 5. This study provides evidence for connections between the spatio-temporal genetic heterogeneity of a Daphnia assemblage and the seasonal changes of water temperature and oxygen content. The data also suggest that not only the actual temperature but also the dynamics of temperature change may influence the genetic structure of Daphnia populations and assemblages.
Daphnia magna Straus, 1820 is a widespread zooplanktic organism enduring considerable changes in oxygen concentration and temperature within its natural habitat. The thermal tolerance window of D. magna was analyzed using the animals' swimming activity as a test parameter in a photometrical assay. Acclimation to different temperatures (10, 20, 30 °C) resulted in a shift of the thermal optimum corresponding to acclimation conditions. Acclimation to warm temperatures also increased the upper thermal tolerance limit in acute thermal tolerance tests. However, the magnitude of the resulting shift in the acute thermal tolerance (LT50) was much smaller. An increase in acclimation temperature by 10 °C changed the thermal optimum by approximately this value, whereas the LT50 was enhanced only by 1–2 °C. The time course of the acclimation process was followed by surveying temperature-dependent swimming activity upon the transfer of animals raised in a medium at 20 °C to a medium at 30 °C. Maximum swimming intensity at 20 °C was lost within 3 days. The swimming behavior resembled that of animals acclimated to 30 °C after 2 weeks, indicating that acclimation to the elevated temperature was achieved.
A field and laboratory study was carried out over 3 yr to determine relationships between seasonal and interannual changes in temperature (year-specific temperature courses, presence or absence of ice in winter) and the genetic structure (composition of multilocus genotypes [MLGs]) of a Daphnia longispina assemblage. Field studies on temperature and genetic structures were linked with laboratory analyses to evaluate the thermal tolerance of long-term 12uC-, 18uC-, and 24uC-acclimated clonal lineages (CLs) derived from abundant MLGs sampled in the field (surface water and thermocline). The tolerance to warm temperatures (heat tolerance) was lowest in CLs derived from MLGs that were dominant directly after or before winter (winter-CLs), higher in ''spring-autumn-CLs,'' and highest in ''summer-CLs.'' Winter-CLs also showed the highest degree of physiological plasticity. The differences in heat tolerance were mainly related to the different genotypes of the phosphoglucomutase (PGM) locus. Temperature conditions during winter and early spring affected the heat tolerance of all CLs as well as the success of different winter survival strategies (overwintering, resting eggs). Heat tolerance was lowest in CLs derived from MLGs sampled in 2006 (after the coldest winter and spring period), higher in CLs from 2005 (after a less cold winter and spring period), and highest in CLs from 2007 (after a warm, ice-free winter). In addition to other environmental factors (predation, parasitism, food), seasonal and interannual changes in temperature affect Daphnia genetic structure through genetic differences in thermal responses, thermal tolerance, and physiological plasticity.Depending on environmental conditions, daphnids alter between asexual (parthenogenesis) and sexual reproduction. Parthenogenetic reproduction, which occurs under favorable conditions (often from spring until autumn in temperate zones), results in the propagation of a varying number of coexisting genotypes (clonal population structure) (Hebert and Crease 1980;Weider 1985; Pantel et al. 2011) that originated from sexual reproduction. Sexual reproduction is inducible by unfavorable conditions (often in late autumn in temperate zones). Thus, selection acting on the different clones, which alters the clonal population structure, as well as genetic recombination and the recruitment of sexually derived genotypes, which arise from resting eggs, are essential factors for the maintenance of genetic diversity in Daphnia populations (Hembre and Megard 2006). Several studies have reported on selection by the spatiotemporal heterogeneity of the environment including predation (De Meester et al. 1995;Cousyn et al. 2001), parasitism (Mitchell et al. 2004), and food quality (Weider et al. 2005; Brzeziński et al. 2010). The influence of spatiotemporal changes in water temperature on the genetic structure of Daphnia populations has also been studied. Carvalho (1987), for instance, reported for Daphnia magna a higher viability and fecundity of winter clones in cold water...
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