Seasonal adaptations and the role of lipids in oceanic zooplankton.

Oceanic zooplankton species exhibit quite diverse life history traits. A major driving force determining their life strategies is the seasonal variability in food supply, which is most pronounced in polar oceans where fluctuations in primary production are extreme. Seasonal adaptations are closely related to the trophic level of zooplankters, with strongest pressures occurring on herbivorous organisms. The dominant grazers, calanoid copepods and krill (Euphausiacea), have developed fascinating solutions for successful overwintering at higher latitudes. They usually exhibit a very efficient storage and utilization of energy reserves to reduce the effect of a highly seasonal primary production. The predominant larger Calanus species from the Arctic and Calanoides acutus from the Antarctic biosynthesize large amounts of high-energy wax esters with long-chain monounsaturated fatty acids and alcohols (20:1 and 22:1 isomers) as major components. They survive the dark season at depth in a stage of dormancy called diapause. In contrast, the Antarctic Calanus propinquus, a winter-active species, synthesizes primarily triacylglycerols, which are dominated by long-chain monounsaturated fatty acids with 22 carbon atoms (2 isomers) and yield even higher calorific contents. The omnivorous and carnivorous species, which are less subjected to seasonal food shortage, usually do not exhibit such an elaborate lipid biosynthesis. Herbivores usually do not utilize much of their enormous lipid reserves for overwintering, but channel this energy towards reproductive processes in late winter/early spring. Timing of reproduction is critical especially at high latitudes due to the short production period, and lipid reserves ensure early spawning independent of external resources. These energetic adaptations (dormancy, lipid storage) are supplemented by other life strategies such as extensive vertical migrations, change in the mode of life, and trophic flexibility.