Ammonium excretion and oxygen respiration of tropical copepods and euphausiids exposed to oxygen minimum zone conditions

Abstract. Calanoid copepods and euphausiids are key components of marine zooplankton communities worldwide. Most euphausiids and several copepod species perform diel vertical migrations (DVMs) that contribute to the export of particulate and dissolved matter to midwater depths. In vast areas of the global ocean, and in particular in the eastern tropical Atlantic and Pacific, the daytime distribution depth of many migrating organisms corresponds to the core of the oxygen minimum zone (OMZ). At depth, the animals experience reduced temperature and oxygen partial pressure (pO₂) and an increased carbon dioxide partial pressure (pCO₂) compared to their near-surface nighttime habitat. Although it is well known that low oxygen levels can inhibit respiratory activity, the respiration response of tropical copepods and euphausiids to relevant pCO₂, pO₂, and temperature conditions remains poorly parameterized. Further, the regulation of ammonium excretion at OMZ conditions is generally not well understood. It was recently estimated that DVM-mediated ammonium supply could fuel bacterial anaerobic ammonium oxidation – a major loss process for fixed nitrogen in the ocean considerably. These estimates were based on the implicit assumption that hypoxia or anoxia in combination with hypercapnia (elevated pCO₂) does not result in a down-regulation of ammonium excretion. We exposed calanoid copepods from the Eastern Tropical North Atlantic (ETNA; Undinula vulgaris and Pleuromamma abdominalis) and euphausiids from the Eastern Tropical South Pacific (ETSP; Euphausia mucronata) and the ETNA (Euphausia gibboides) to different temperatures, carbon dioxide and oxygen levels to study their survival, respiration and excretion rates at these conditions. An increase in temperature by 10 °C led to an approximately 2-fold increase of the respiration and excretion rates of U. vulgaris (Q_{10, respiration} = 1.4; Q_{10, NH4-excretion} = 1.6), P. abdominalis (Q_{10, respiration} = 2.0; Q_{10, NH4-excretion} = 2.4) and E. gibboides (Q_{10, respiration} = 2.0; Q_{10, NH4-excretion} = 2.4; E. mucronata not tested). Exposure to differing carbon dioxide levels had no overall significant impact on the respiration or excretion rates. Species from the ETNA were less tolerant to low oxygen levels than E. mucronata from the ETSP, which survived exposure to anoxia at 13 °C. Respiration and excretion rates were reduced upon exposure to low oxygen levels, albeit at different species-specific levels. Reduction of the excretion and respiration rates in ETNA species occurred at a pO₂ of 0.6 (P. abdominalis) and 2.4 kPa (U. vulgaris and E. gibboides) at OMZ temperatures. Such low oxygen levels are normally not encountered by these species in the ETNA. E. mucronata however regularly migrates into the strongly hypoxic to anoxic core of the ETSP OMZ. Exposure to low oxygen levels led to a strong reduction of respiration and ammonium excretion in E. mucronata (p_{crit  respiration} = 0.6, p_{crit NH4-excretion} = 0.73). A drastic reduction of respiratory activity was also observed by other authors for euphausiids, squat lobsters and calanoid copepods, but was not yet accounted for when calculating DVM-mediated active fluxes into the ETSP OMZ. Current estimates of DVM-mediated active export of carbon and nitrogen into the ETSP OMZ are therefore likely too high and future efforts to calculate these export rates should take the physiological responses of migratory species to OMZ conditions into account.