Using solar energy suboptimally: How efficient are phytoplankton at converting sunlight into the products of photosynthesis? The two other pathways that that absorbed energy can take are emission back to the environment by fluorescence or conversion to heat. Lin et al. measured phytoplankton fluorescence lifetimes in the laboratory and combined them with satellite measurements of variable chlorophyll fluorescence. Combined, they determined the quantum yields of photochemistry and fluorescence in four ocean basins. Approximately 60% of absorbed solar energy is converted to heat, a figure 50% higher than has been determined for conditions of optimal growth. Science , this issue p. 264

Approximately 45% of the photosynthetically fixed carbon on Earth occurs in the oceans in phytoplankton, which account for less than 1% of the world's photosynthetic biomass. This amazing empirical observation implies a very high photosynthetic energy conversion efficiency, but how efficiently is the solar energy actually used? The photon energy budget of photosynthesis can be divided into three terms: the quantum yields of photochemistry, fluorescence, and heat. Measuring two of these three processes closes the energy budget. The development of ultrasensitive, seagoing chlorophyll variable fluorescence and picosecond fluorescence lifetime instruments has allowed independent closure on the first two terms. With this closure, we can understand how phytoplankton respond to nutrient supplies on timescales of hours to months and, over longer timescales, to changes in climate.

Variable fluorescence techniques are increasingly used to assess phytoplankton photosynthesis. All fluorescence techniques and models for photosynthetic electron transport rates (ETRs) are amplitude‐based and are subject to errors, especially when phytoplankton growth is nutrient‐limited. Here we develop a new, kinetic‐based approach to measure, directly and in absolute units, ETRs and to estimate growth rates in phytoplankton. We applied this approach to investigate the effects of nitrogen limitation on phytoplankton photophysiology and growth rates. Nutrient stress leads to a decrease in the quantum yield of photochemistry in Photosystem II (F_v/F_m); however, the relationship between F_v/F_m and growth rates is highly nonlinear, which makes it impossible to quantify the reduction in phytoplankton growth rates from F_v/F_m alone. In contrast, the decline in growth rates under nitrogen stress was proportional to the decrease in kinetic‐based photosynthetic rates. Our analysis suggests the kinetic fluorescence measurements markedly improve the accuracy of ETR measurements, as compared to classical amplitude‐based measurements. Fluorescence‐based methods for primary production rely on measurements of ETRs and then conversion to carbon fixation rates by using the electron yields of carbon fixation. The electron yields exhibit 10‐fold variability in natural phytoplankton communities and are strongly affected by nutrient limitation. Our results reveal that a decrease in the growth rates and the electron yields of carbon fixation is driven by, and can be quantified from, a decrease in photosynthetic turnover rates. We propose an algorithm to deduce the electron yields of carbon fixation, which greatly improve fluorescence‐based measurements of primary production and growth rates.

Amundsen Sea polynyas are among the most productive, yet climate‐sensitive ecosystems in the Southern Ocean and host massive annual phytoplankton blooms. These blooms are believed to be controlled by iron fluxes from melting ice and icebergs and by intrusion of nutrient‐rich Circumpolar Deep Water, however the interplay between iron effects and other controls, such as light availability, has not yet been quantified. Here, we examine phytoplankton photophysiology in relation to Fe stress and physical forcing in two largest polynyas, Amundsen Sea Polynya (ASP) and Pine Island Polynya (PIP), using the combination of high‐resolution variable fluorescence measurements, fluorescence lifetime analysis, photosynthetic rates, and Fe‐enrichment incubations. These analyses revealed strong Fe stress in the ASP, whereas the PIP showed virtually no signatures of Fe limitation. In spite of enhanced iron availability in the PIP, chlorophyll biomass remained ∼ 30–50% lower than in the Fe‐stressed ASP. This apparent paradox would not have been observed if iron were the main control of phytoplankton bloom in the Amundsen Sea. Long‐term satellite‐based climatology records revealed that the ASP is exposed to significantly higher solar irradiance levels throughout the summer season, as compared to the PIP region, suggesting that light availability controls the magnitude of phytoplankton blooms in the Amundsen Sea. Our data suggests that higher Fe availability (e.g., due to higher melting rates of ice sheets) would not necessarily increase primary productivity in this region. Furthermore, stronger wind‐driven vertical mixing in expanding ice‐free areas may lead to reduction in light availability and productivity in the future.

Phytoplankton in the Arctic Ocean are subject to nitrogen limitation in the summer, however, how severely the nitrogen stress affects phytoplankton physiology remains largely unknown. In the summers of 2015–2018, we examined the distribution of phytoplankton photophysiological properties across two contrasting regions of the Arctic Ocean with distinctly different levels of nitrogen availability in the upper water column. We quantified the extent of nitrogen stress using a highly sensitive fluorescence induction and relaxation system to obtain continuous underway measurements and via discrete sample analyses of phytoplankton physiology, as well as nutrient enrichment incubations. The results revealed vast regions in the Chukchi Sea where phytoplankton photosynthesis was severely nitrogen‐stressed. Thereby, the maximum quantum yield of photochemistry in photosystem II showed only a small decrease (12 ± 9%) relative to its nutrient‐replete values, while the maximum photosynthetic electron transport rates under saturating irradiance were impaired to a greater extent (40 ± 17%). This phytoplankton photosynthesis response is indicative of a severe nitrogen limitation, which results in dramatic reduction in growth and net primary production rates. Nutrient enrichment incubations also revealed a marked increase in large‐size phytoplankton growth (>10 μm) after the nitrogen stress was alleviated, suggesting that the larger cells were more susceptible to nitrogen stress. These results are important for understanding how regional nitrogen fluxes control variability in the primary production and phytoplankton community structure and how these processes might change with rapid climate changes in the Arctic Ocean.