Phytoplankton photosynthesis is a critical flux in the carbon cycle, accounting for approximately 40% of the carbon dioxide fixed globally on an annual basis and fuelling the productivity of aquatic food webs. However, rapid evolutionary responses of phytoplankton to warming remain largely unexplored, particularly outside the laboratory, where multiple selection pressures can modify adaptation to environmental change. Here, we use a decade-long experiment in outdoor mesocosms to investigate mechanisms of adaptation to warming (+4 °C above ambient temperature) in the green alga Chlamydomonas reinhardtii, in naturally assembled communities. Isolates from warmed mesocosms had higher optimal growth temperatures than their counterparts from ambient treatments. Consequently, warm-adapted isolates were stronger competitors at elevated temperature and experienced a decline in competitive fitness in ambient conditions, indicating adaptation to local thermal regimes. Higher competitive fitness in the warmed isolates was linked to greater photosynthetic capacity and reduced susceptibility to photoinhibition. These findings suggest that adaptive responses to warming in phytoplankton could help to mitigate projected declines in aquatic net primary production by increasing rates of cellular net photosynthesis.

The efficiency of carbon sequestration by the biological pump could decline in the coming decades because respiration tends to increase more with temperature than photosynthesis. Despite these differences in the short‐term temperature sensitivities of photosynthesis and respiration, it remains unknown whether the long‐term impacts of global warming on metabolic rates of phytoplankton can be modulated by evolutionary adaptation. We found that respiration was consistently more temperature dependent than photosynthesis across 18 diverse marine phytoplankton, resulting in universal declines in the rate of carbon fixation with short‐term increases in temperature. Long‐term experimental evolution under high temperature reversed the short‐term stimulation of metabolic rates, resulting in increased rates of carbon fixation. Our findings suggest that thermal adaptation may therefore have an ameliorating impact on the efficiency of phytoplankton as primary mediators of the biological carbon pump.

Models of marine biogeochemistry capture the effects of temperature on phytoplankton growth via the monotonic, exponential Eppley coefficient, without considering the physiological or evolutionary processes that underpin this emergent, across‐species temperature response. Here, we investigated both the within‐ and across‐species temperature dependence of growth rate for 18 species of marine phytoplankton. We found that the temperature dependence of growth rate derived across species was lower than the average temperature response within species. This finding supports a “partial compensation” model of thermal adaptation and suggests that adaptation can partially compensate for the underlying thermodynamic effects of temperature on physiological rates observed within species. We also found that thermal tolerance traits (e.g. the optimum temperature for growth) systematically covaried with a host of key functional traits (e.g. cell size, elemental composition). Consequently, turnover in species composition in a warmer ocean, linked to interspecific variability in thermal tolerance traits, could be associated with major shifts in the functional trait composition of marine phytoplankton communities with far reaching implications for ecosystem functioning.

A mathematical gas exchange model, using sinusoidal forcing functions of inert inspired gas (A. Zwart, R. C. Seagrave, and A. Van Dieren. J. Appl. Physiol. 41: 419-424, 1976), has been extended by us to include dead space (VD), a single alveolar compartment (VA) perfused with blood flow (Qp), and a shunt (Qs). In this new work we use N2O as the indicator gas in the mathematical model and in the experimental studies, in low enough concentrations [<6% (vol/vol)] to avoid anesthetic effects. Mathematical relationships between the inspired and expired N2O gas partial pressures, the blood gas N2O partial pressures, and their variation with forcing frequency are derived for a continuous ventilation uptake and a conventional anesthetic gas distribution model. We show that these gas and blood gas N2O relationships give direct derivation of cardiorespiratory parameters such as VA, Qp, the dead space-to-total ventilation ratio (VD/VT), and the shunt-to-total blood flow ratio (Qs/QT) without altering the subject's oxygenation and that they are essentially free from recirculation effects at high forcing frequencies > or = 2 min-1. Theoretical results from the model are presented for a wide range of forcing frequencies between 2 x 10(-2) and 10 min-1 (sinusoid periods 30-0.1 min), and these show that VA, Qp, and VD/VT can all be measured by N2O forcing frequencies > or = 1 min-1. We also present results from five animal studies, with an experimental inspired gas forcing frequency range of 0.125 to 2 min-1, which show qualitative agreement with the predictions of the continuous ventilation model. During these animal studies both mass spectrometric N2O respiratory gas measurements and intravascular polarographic arterial and mixed venous blood N2O partial pressure measurements were made, and examples of these in vivo measurements are presented, together with examples of the calculations derived from them.

Rising sea surface temperatures are expected to lead to the loss of phytoplankton biodiversity. However, we currently understand very little about the interactions between warming, loss of phytoplankton diversity and its impact on the oceans' primary production. We experimentally manipulated the species richness of marine phytoplankton communities under a range of warming scenarios, and found that ecosystem production declined more abruptly with species loss in communities exposed to higher temperatures. Species contributing positively to ecosystem production in the warmed treatments were those that had the highest optimal temperatures for photosynthesis, implying that the synergistic impacts of warming and biodiversity loss on ecosystem functioning were mediated by thermal trait variability. As species were lost from the communities, the probability of taxa remaining that could tolerate warming diminished, resulting in abrupt declines in ecosystem production. Our results highlight the potential for synergistic effects of warming and biodiversity loss on marine primary production.

We report single-entity measurements of the degree of calcification of individual phytoplankton cells. Electrogenerated acid is used to dissolve the calcium carbonate (CaCO3) shell (coccosphere) of individual coccolithophores and the...

Coccoliths are plates of biogenic calcium carbonate secreted by calcifying marine phytoplankton; annually these phytoplankton are responsible for exporting >1 billion tonnes (1015 g) of calcite to the deep ocean. Rapid and reliable methods for assessing the degree of calcification are technically challenging because the coccoliths are micron sized and contain picograms (pg) of calcite. Here we pioneer an opto‐eletrochemical acid titration of individual coccoliths which allows 3D reconstruction of each individual coccolith via in situ optical imaging enabling direct inference of the coccolith mass. Coccolith mass ranging from 2 to 400 pg are reported herein, evidencing both inter‐ and intra‐species variation over four different species. We foresee this scientific breakthrough, which is independent of knowledge regarding the species and calibration‐free, will allow continuous monitoring and reporting of the degree of coccolith calcification in the changing marine environment.

The 21st century has seen an acceleration of anthropogenic climate change and biodiversity loss, with both stressors deemed to affect ecosystem functioning. However, we know little about the interactive effects of both stressors and in particular about the interaction of increased climatic variability and biodiversity loss on ecosystem functioning. This should be remedied because larger climatic variability is one of the main features of climate change. Here, we demonstrated that temperature fluctuations led to changes in the importance of biodiversity for ecosystem functioning. We used microcosm communities of different phytoplankton species richness and exposed them to a constant, mild, and severe temperature-fluctuating environment. Wider temperature fluctuations led to steeper biodiversity–ecosystem functioning slopes, meaning that species loss had a stronger negative effect on ecosystem functioning in more fluctuating environments. For severe temperature fluctuations, the slope increased through time due to a decrease of the productivity of species-poor communities over time. We developed a theoretical competition model to better understand our experimental results and showed that larger differences in thermal tolerances across species led to steeper biodiversity–ecosystem functioning slopes. Species-rich communities maintained their ecosystem functioning with increased fluctuation as they contained species able to resist the thermally fluctuating environments, while this was on average not the case in species-poor communities. Our results highlight the importance of biodiversity for maintaining ecosystem functions and services in the context of increased climatic variability under climate change.