scholarly journals A year in the life of a central California kelp forest: physical and biological insights into biogeochemical variability

2016 ◽  
Author(s):  
David A. Koweek ◽  
Kerry J. Nickols ◽  
Paul R. Leary ◽  
Steve Y. Litvin ◽  
Tom W. Bell ◽  
...  
Keyword(s):  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Kelp Forest ◽  
Total Alkalinity ◽  
Kelp Forests ◽  
Saturation State ◽  
Central California ◽  
High Co2 ◽  
Spatial Gradients ◽  
Spatially Distributed

Abstract. Kelp forests are among the world's most productive marine ecosystems, yet little is known about their biogeochemistry. This study presents a fourteen-month time series (July 2013–August 2014) of surface and benthic dissolved inorganic carbon and total alkalinity measurements, along with accompanying hydrographic measurements, from six locations within a central California kelp forest. We present ranges and patterns of variability in carbonate chemistry, including pH (7.70–8.33), pCO2 (172–952 µatm), and the aragonite saturation state, ΩAr (0.94–3.91). Surface-to-bottom gradients in CO2 system chemistry were as large as the spatial gradients throughout the bottom of the kelp forest. Dissolved inorganic carbon variability was the main driver of the observed CO2 system variability. The majority of spatial variability in the kelp forest can be explained by advection of cold, dense high CO2 waters into the bottom of the kelp forest, with deeper sites experiencing high CO2 conditions more frequently. Despite the strong imprint of advection on the biogeochemical variability of the kelp forest, surface waters were undersaturated with CO2 in the spring through fall, indicative of the strong role of photosynthesis on biogeochemical variability. We emphasize the importance of spatially distributed measurements for developing a process-based understanding of kelp forest ecosystem function in a changing climate.

scholarly journals A year in the life of a central California kelp forest: physical and biological insights into biogeochemical variability

2017 ◽  
Vol 14 (1) ◽  
pp. 31-44 ◽  
Author(s):  
David A. Koweek ◽  
Kerry J. Nickols ◽  
Paul R. Leary ◽  
Steve Y. Litvin ◽  
Tom W. Bell ◽  
...  
Keyword(s):  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Kelp Forest ◽  
Total Alkalinity ◽  
Kelp Forests ◽  
Saturation State ◽  
Central California ◽  
High Co2 ◽  
Spatial Gradients ◽  
Spatially Distributed

Abstract. Kelp forests are among the world's most productive marine ecosystems, yet little is known about their biogeochemistry. This study presents a 14-month time series (July 2013–August 2014) of surface and benthic dissolved inorganic carbon and total alkalinity measurements, along with accompanying hydrographic measurements, from six locations within a central California kelp forest. We present ranges and patterns of variability in carbonate chemistry, including pH (7.70–8.33), pCO2 (172–952 µatm), and the aragonite saturation state, ΩAr (0.94–3.91). Surface-to-bottom gradients in CO2 system chemistry were as large as the spatial gradients throughout the bottom of the kelp forest. Dissolved inorganic carbon variability was the main driver of the observed CO2 system variability. The majority of spatial variability in the kelp forest can be explained by advection of cold, dense high-CO2 waters into the bottom of the kelp forest, with deeper sites experiencing high-CO2 conditions more frequently. Despite the strong imprint of advection on the biogeochemical variability of the kelp forest, surface waters were undersaturated with CO2 in the spring through fall, indicative of the strong role of photosynthesis on biogeochemical variability. We emphasize the importance of spatially distributed measurements for developing a process-based understanding of kelp forest ecosystem function in a changing climate.

scholarly journals Inorganic carbon fluxes on the Mackenzie Shelf of the Beaufort Sea

2017 ◽  
Author(s):  
Jacoba Mol ◽  
Helmuth Thomas ◽  
Paul G. Myers ◽  
Xianmin Hu ◽  
Alfonso Mucci
Keyword(s):  
Sea Ice ◽  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Beaufort Sea ◽  
Total Alkalinity ◽  
The Arctic ◽  
Surface Mixed Layer ◽  
Saturation State ◽  
Carbon Transfer ◽  
Mackenzie Shelf

Abstract. The Mackenzie Shelf in the southeastern Beaufort Sea is a region that has experienced large changes in the past several decades as warming, sea-ice loss, and increased river discharge have altered carbon cycling. Upwelling and downwelling events are common on the shelf, caused by strong, fluctuating along-shore winds, resulting in cross-shelf Ekman transport, and an alternating estuarine and anti-estuarine circulation. Downwelling carries inorganic carbon and other remineralization products off the shelf and into the deep basin for possible long-term storage in the world oceans. Upwelling carries dissolved inorganic carbon (DIC) and nutrient-rich waters from the Pacific-origin upper halocline layer (UHL) onto the shelf. Profiles of DIC and total alkalinity (TA) taken in August and September of 2014 are used to investigate the cycling of inorganic carbon on the Mackenzie Shelf. The along-shore transport of water and the cross-shelf transport of inorganic carbon are quantified using velocity field output from a simulation of the Arctic and Northern Hemisphere Atlantic (ANHA4) configuration of the Nucleus of European Modelling of the Ocean (NEMO) framework. A strong upwelling event prior to sampling on the Mackenzie Shelf is analyzed and the resulting influence on the carbonate system, including the saturation state of waters with respect to aragonite and pH, is investigated. TA and the oxygen isotope ratio of water (δ18O) are used to examine water-mass distributions in the study area and to investigate the influence of Pacific Water, Mackenzie River freshwater, and sea-ice melt on carbon dynamics and air-sea fluxes of carbon dioxide (CO2) in the surface mixed layer. Understanding carbon transfer in this seasonally dynamic environment is key to quantify the importance of Arctic shelf regions to the global carbon cycle and provide a basis for understanding how it will respond to the aforementioned climate-induced changes.

scholarly journals CO<sub>2</sub> perturbation experiments: similarities and differences between dissolved inorganic carbon and total alkalinity manipulations

2009 ◽  
Vol 6 (10) ◽  
pp. 2145-2153 ◽  
Author(s):  
K. G. Schulz ◽  
J. Barcelos e Ramos ◽  
R. E. Zeebe ◽  
U. Riebesell
Keyword(s):  
Ocean Acidification ◽  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Total Alkalinity ◽  
Calcium Carbonate Precipitation ◽  
Carbonate Chemistry ◽  
Carbonate System ◽  
Experimental Conditions ◽  
Saturation State ◽  
Basic Approaches

Abstract. Increasing atmospheric carbon dioxide (CO2) through human activities and invasion of anthropogenic CO2 into the surface ocean alters the seawater carbonate chemistry, increasing CO2 and bicarbonate (HCO3−) at the expense of carbonate ion (CO32−) concentrations. This redistribution in the dissolved inorganic carbon (DIC) pool decreases pH and carbonate saturation state (Ω). Several components of the carbonate system are considered potential key variables influencing for instance calcium carbonate precipitation in marine calcifiers such as coccolithophores, foraminifera, corals, mollusks and echinoderms. Unravelling the sensitivities of marine organisms and ecosystems to CO2 induced ocean acidification (OA) requires well-controlled experimental setups and accurate carbonate system manipulations. Here we describe and analyse the chemical changes involved in the two basic approaches for carbonate chemistry manipulation, i.e. changing DIC at constant total alkalinity (TA) and changing TA at constant DIC. Furthermore, we briefly introduce several methods to experimentally manipulate DIC and TA. Finally, we examine responses obtained with both approaches using published results for the coccolithophore Emiliania huxleyi. We conclude that under most experimental conditions in the context of ocean acidification DIC and TA manipulations yield similar changes in all parameters of the carbonate system, which implies direct comparability of data obtained with the two basic approaches for CO2 perturbation.

scholarly journals Estimates of ikaite export from sea ice to the underlying seawater in a sea ice–seawater mesocosm

2016 ◽  
Vol 10 (5) ◽  
pp. 2173-2189 ◽  
Author(s):  
Nicolas-Xavier Geilfus ◽  
Ryan J. Galley ◽  
Brent G. T. Else ◽  
Karley Campbell ◽  
Tim Papakyriakou ◽  
...  
Keyword(s):  
Sea Ice ◽  
Water Column ◽  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Carbon Dynamics ◽  
Total Alkalinity ◽  
Main Process ◽  
Saturation State ◽  
Ice Growth ◽  
Ice Melt

Abstract. The precipitation of ikaite and its fate within sea ice is still poorly understood. We quantify temporal inorganic carbon dynamics in sea ice from initial formation to its melt in a sea ice–seawater mesocosm pool from 11 to 29 January 2013. Based on measurements of total alkalinity (TA) and total dissolved inorganic carbon (TCO2), the main processes affecting inorganic carbon dynamics within sea ice were ikaite precipitation and CO2 exchange with the atmosphere. In the underlying seawater, the dissolution of ikaite was the main process affecting inorganic carbon dynamics. Sea ice acted as an active layer, releasing CO2 to the atmosphere during the growth phase, taking up CO2 as it melted and exporting both ikaite and TCO2 into the underlying seawater during the whole experiment. Ikaite precipitation of up to 167 µmolkg−1 within sea ice was estimated, while its export and dissolution into the underlying seawater was responsible for a TA increase of 64–66 µmolkg−1 in the water column. The export of TCO2 from sea ice to the water column increased the underlying seawater TCO2 by 43.5 µmolkg−1, suggesting that almost all of the TCO2 that left the sea ice was exported to the underlying seawater. The export of ikaite from the ice to the underlying seawater was associated with brine rejection during sea ice growth, increased vertical connectivity in sea ice due to the upward percolation of seawater and meltwater flushing during sea ice melt. Based on the change in TA in the water column around the onset of sea ice melt, more than half of the total ikaite precipitated in the ice during sea ice growth was still contained in the ice when the sea ice began to melt. Ikaite crystal dissolution in the water column kept the seawater pCO2 undersaturated with respect to the atmosphere in spite of increased salinity, TA and TCO2 associated with sea ice growth. Results indicate that ikaite export from sea ice and its dissolution in the underlying seawater can potentially hamper the effect of oceanic acidification on the aragonite saturation state (Ωaragonite) in fall and in winter in ice-covered areas, at the time when Ωaragonite is smallest.

scholarly journals Macroalgae contribute to nested mosaics of pH variability in a sub-Arctic fjord

2015 ◽  
Vol 12 (6) ◽  
pp. 4907-4945 ◽  
Author(s):  
D. Krause-Jensen ◽  
C. M. Duarte ◽  
I. E. Hendriks ◽  
L. Meire ◽  
M. E. Blicher ◽  
...  
Keyword(s):  
Arctic Ocean ◽  
Boundary Layers ◽  
Large Scale ◽  
Dissolved Inorganic Carbon ◽  
Spatial Scales ◽  
Total Alkalinity ◽  
The Arctic ◽  
Kelp Forests ◽  
Coastal Habitats ◽  
Saturation State

Abstract. The Arctic Ocean is considered the most vulnerable ecosystem to ocean acidification (OA) and large-scale assessments of pH and the saturation state for aragonite (Ωarag) indicate that it is already close to corrosive states (Ωarag < 1). In high-latitude coastal waters the regulation of pH and Ωarag is far more complex than offshore because increased biological activity and input of glacial meltwater affect pH. As most calcifiers occupy coastal habitats, the assessment of risks from OA to these vulnerable organisms cannot be derived from extrapolation of current and forecasted offshore conditions, but requires an understanding of the regimes of pH and Ωarag in their coastal habitats. To increase knowledge of the natural variability of pH in the Arctic coastal zone and specifically to test the influence of benthic vegetated habitats, we quantified pH-variability in a Greenland fjord in a nested scale approach. A sensor array logging pH, O2, PAR, temperature and salinity was applied on spatial scales ranging from km-scale across the horizontal extension of the fjord, over 100 m scale vertically in the fjord, 10–100 m scale between subtidal habitats with and without kelp forests and between vegetated tidal pools and adjacent vegetated shores, to cm-m scale within kelp forests and mm-scale across boundary layers of macrophyte tissue. In addition, we assessed the temporal variability in pH on diurnal and seasonal scales. Based on pH-measurements combined with relationships between salinity, total alkalinity and dissolved inorganic carbon we also estimated variability of Ωarag. Results show variability in pH and Ωarag of up to 0.2–0.3 units at several scales, i.e. along the horizontal and vertical extension of the fjord, between seasons and on a diel basis in benthic habitats and within 1 m3 of kelp forest. Vegetated intertidal pools exhibited extreme diel pH variability of > 1.5 units and macrophyte boundary layers a pH-range of up to 0.8 units. Overall, Ωarag was favorable to calcification, and pelagic and benthic metabolism was an important driver of pH and Ωarag producing mosaics of variability from low levels in the dark to peak levels at high irradiance. We suggest that productive coastal environments may form niches of high pH in a future acidified Arctic Ocean.

scholarly journals Impacts of ikaite export from sea ice to the underlying seawater in a sea ice-seawater mesocosm

2016 ◽  
Author(s):  
N.-X Geilfus ◽  
R. J. Galley ◽  
B. G. T. Else ◽  
T. Papakyriakou ◽  
O. Crabeck ◽  
...  
Keyword(s):  
Sea Ice ◽  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Open Water ◽  
Carbon Dynamics ◽  
Co2 Exchange ◽  
Total Alkalinity ◽  
Main Process ◽  
Saturation State ◽  
Ice Growth

Abstract. Ikaite precipitation within sea ice could act as a significant sink for atmospheric CO2. However, the fate of these ikaite crystals is still poorly understood. We quantify temporal inorganic carbon dynamics from initial sea ice formation from open water to its melt during a month-long experiment in a sea ice-seawater mesocosm pool. Within sea ice, ikaite precipitation and CO2 exchange with the atmosphere were the main processes affecting inorganic carbon dynamics, while the dissolution of ikaite was the main process affecting inorganic carbon dynamics in the underlying seawater. Based on the total alkalinity (TA) and total dissolved inorganic carbon (TCO2) within sea ice and seawater, we estimated ikaite precipitated up to 167 ± 3 µmol kg-1 within sea ice; up to 57 % of the ikaite precipitated within sea ice was exported to the underlying seawater where it was dissolved. Ikaite export from the ice to the underlying seawater was associated with brine rejection during sea ice growth, increased sea ice vertical connectivity due to the upward percolation of seawater, and meltwater flushing during sea ice melt. The dissolution of the ikaite crystals in the water column kept the seawater pCO2 undersaturated compared to the atmosphere in spite of increased salinity, TA, and TCO2 associated with sea ice growth. Results indicate that ikaite export from sea ice and its dissolution in the underlying seawater can potentially hamper the effect of oceanic acidification on the aragonite saturation state (Ωaragonite) in fall and winter in ice-covered areas, at the time when Ωaragonite is smallest.

Seasonal variation in inorganic carbon parameters in the southwestern Yellow Sea

2020 ◽  
Author(s):  
Minwoo Seok ◽  
Ahra Mo ◽  
Seunghee Park ◽  
Young Ho Ko ◽  
Seongtae Yun ◽  
...  
Keyword(s):  
Yellow Sea ◽  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Ph Sensor ◽  
Vertical Mixing ◽  
Sampling Interval ◽  
Total Alkalinity ◽  
The Yellow Sea ◽  
Sudden Increase ◽  
Saturation State

&lt;div&gt; &lt;p&gt;To better understand carbon cycles in coastal and marginal seas, time-series monitoring is essential because of large temporal variabilities. In this regard, we conducted monthly field researches from April 2017 to May 2019 at the Socheongcho (SCC) Ocean Research Site (37&amp;#176;N&amp;#8218;124&amp;#176;E) in the Yellow Sea located between Korea and China. At each survey, we collected surface seawater samples during approximately 7 days with an sampling interval of two hours (except for spring 2017). Total alkalinity (TA) and dissolved inorganic carbon (DIC) were analyzed by using VINDTA 3C system, Apollo SciTech DIC analyzer and Apollo SciTech Alkalinity Titrator. In addition, a pH sensor (SeapHOx) was installed at the surface layer from September 2018 to June 2019 which is also capable of measuring salinity, temperature and oxygen.&amp;#160;Based on the observations, we estimated a partial pressure of carbon dioxide (&lt;em&gt;p&lt;/em&gt;CO&lt;sub&gt;2&lt;/sub&gt;) and aragonite saturation state. As expected, seasonal variations in TA and DIC were strongly associated with those of salinity. We also detected a sudden increase DIC in October when vertical mixing was greatly enhanced. Despite a large outgassing during the fall season, annual mean air--sea influx of CO&lt;sub&gt;2&lt;/sub&gt; was &amp;#8764;0.61mol&amp;#183;m&lt;sup&gt;&amp;#8722;2&lt;/sup&gt;&amp;#183;year&lt;sup&gt;&amp;#8722;1&lt;/sup&gt;, suggesting that the study area was a weak sink for atmospheric CO&lt;sub&gt;2&lt;/sub&gt;. Aragonite was enerally reduced during winter (&amp;#8764;1.5). However, no undersaturation event was found during the whole investigation.&lt;/p&gt; &lt;/div&gt;

scholarly journals Carbonate system distribution, anthropogenic carbon and acidification in the western tropical South Pacific (OUTPACE 2015 transect)

2018 ◽  
Vol 15 (16) ◽  
pp. 5221-5236 ◽  
Author(s):  
Thibaut Wagener ◽  
Nicolas Metzl ◽  
Mathieu Caffin ◽  
Jonathan Fin ◽  
Sandra Helias Nunige ◽  
...  
Keyword(s):  
Inorganic Carbon ◽  
South Pacific ◽  
Total Alkalinity ◽  
Total Inorganic Carbon ◽  
Saturation State ◽  
Anthropogenic Carbon ◽  
Subsurface Waters ◽  
Derived Properties ◽  
Intermediate Waters ◽  
South Pacific Gyre

Abstract. The western tropical South Pacific was sampled along a longitudinal 4000 km transect (OUTPACE cruise, 18 February, 3 April 2015) for the measurement of carbonate parameters (total alkalinity and total inorganic carbon) between the Melanesian Archipelago (MA) and the western part of the South Pacific gyre (WGY). This paper reports this new dataset and derived properties: pH on the total scale (pHT) and the CaCO3 saturation state with respect to aragonite (Ωara). We also estimate anthropogenic carbon (CANT) distribution in the water column using the TrOCA method (Tracer combining Oxygen, inorganic Carbon and total Alkalinity). Along the OUTPACE transect a deeper penetration of CANT in the intermediate waters was observed in the MA, whereas highest CANT concentrations were detected in the subsurface waters of the WGY. By combining our OUTPACE dataset with data available in GLODAPv2 (1974–2009), temporal changes in oceanic inorganic carbon were evaluated. An increase of 1.3 to 1.6 µmol kg−1 a−1 for total inorganic carbon in the upper thermocline waters is estimated, whereas CANT increases by 1.1 to 1.2 µmol kg−1 a−1. In the MA intermediate waters (27 kg m−3 <σθ<27.2 kg m−3) an increase of 0.4 µmol kg−1 a−1 CANT is detected. Our results suggest a clear progression of ocean acidification in the western tropical South Pacific with a decrease in the oceanic pHT of up to −0.0027 a−1 and a shoaling of the saturation depth for aragonite of up to 200 m since the pre-industrial period.

Small-Scale Gradients in Big River: Implications for a State-of-the-Art Research Platform in the Elbe

2021 ◽  
Author(s):  
Sina Bold ◽  
Justus E.E. van Beusekom ◽  
Yoana G. Voynova ◽  
Marius Cysewski ◽  
Bryce Van Dam ◽  
...  
Keyword(s):  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Ammonium Phosphate ◽  
Total Alkalinity ◽  
Small Scale ◽  
Suspended Particulate ◽  
Research Platform ◽  
Biogeochemical Parameters ◽  
Art Research ◽  
Set Up

&lt;p&gt;Estuaries are crucial in transforming matter fluxes from land to sea. To better understand and quantify these processes and respective fluxes, it is important to determine the input into an estuary accurately. To allow for such studies in the Elbe estuary in Germany, a state-of-the-art research platform is currently being set-up just upstream of the weir in Geesthacht at the entrance of the estuary. Here, we report on small-scale spatial dynamics of organic matter and associated processes from several cross and longitudinal profiles around the planned location and the implications for the set-up of the aforementioned research platform.&lt;/p&gt;&lt;p&gt;Based on preliminary data obtained in August 2020 during a period of relatively low discharge, we present the following results: (1) In three cross profiles along a 10 km transect of the Elbe upstream of the weir, we observed considerable small-scale gradients regarding currents and various biogeochemical parameters. In comparison to the fairway, water from the riverbanks was depleted in suspended particulate matter, chlorophyll a, dissolved oxygen, and nitrate, and enhanced in ammonium, phosphate and silicate, as well as total alkalinity and dissolved inorganic carbon paralleled by decreasing pH. This suggests that in the summer, organic matter is deposited and remineralised at the riverbanks, resulting in the release of ammonium, phosphate and silicate, and in the removal of nitrate, presumably by denitrification. (2) Along the 10 km transect towards the weir, we observed that concentrations of suspended particulate matter, chlorophyll a, dissolved oxygen, nitrate and pH were decreasing. In contrast, we found that ammonium, phosphate and silicate, total alkalinity and dissolved inorganic carbon increased towards the weir. This suggests an increased sedimentation and subsequent remineralisation due to the reduced flow velocities in front of the weir. (3) An analysis of a 10-year time series from the weir supports this by showing higher ammonium concentrations when discharges were relatively low. The implications of these findings for the set-up of the research platform in this area, as well as for optimising estimates of budgets are discussed. The research platform will contribute to understand further such variations in biogeochemical parameters at the entrance of the Elbe estuary over time.&lt;/p&gt;&lt;p&gt;The research platform is set-up in cooperation with the Helmholtz initiative MOSES (&amp;#8220;Modular Observation Solutions for Earth Systems&amp;#8220;) and will be incorporated in the Elbe-North Sea Supersite of DANUBIUS-RI (&amp;#8220;International Centre for Advanced Studies on River-Sea Systems&amp;#8220;). Funding is provided by European Regional Development Funds, the federal state of Schleswig-Holstein, the Helmholtz Association and the Helmholtz-Zentrum Geesthacht. The research platform, planned to be operational in autumn 2021, will also be open for users e.g. to develop and test new methods and technologies. Data will be made available through the &amp;#8220;Helmholtz Coastal Data Centre&amp;#8221; (HCDC).&lt;/p&gt;

Photosynthetic utilisation of inorganic carbon and its regulation in the marine diatom Skeletonema costatum

2004 ◽  
Vol 31 (10) ◽  
pp. 1027 ◽  
Author(s):  
Xiongwen Chen ◽  
Kunshan Gao
Keyword(s):  
Inorganic Carbon ◽  
Dissolved Inorganic Carbon ◽  
Skeletonema Costatum ◽  
Co2 Concentration ◽  
Marine Diatom ◽  
Disulfonic Acid ◽  
High Co2 ◽  
Low Co2 ◽  
Diatom Skeletonema Costatum ◽  
Very High

Photosynthetic uptake of inorganic carbon and regulation of photosynthetic CO2 affinity were investigated in Skeletonema costatum (Grev.) Cleve. The pH independence of K1/2(CO2) values indicated that algae grown at either ambient (12 μmol L–1) or low (3 μmol L–1) CO2 predominantly took up CO2 from the medium. The lower pH compensation point (9.12) and insensitivity of photosynthetic rate to di-isothiocyanatostilbene disulfonic acid (DIDS) indicated that the alga had poor capacity for direct HCO3– utilisation. Photosynthetic CO2 affinity is regulated by the concentration of CO2 rather than HCO3–, CO32– or total dissolved inorganic carbon (DIC) in the medium. The response of photosynthetic CO2 affinity to changes in CO2 concentration was most sensitive within the range 3–48 μmol L–1 CO2. Light was required for the induction of photosynthetic CO2 affinity, but not for its repression, when cells were shifted between high (126 μmol L–1) and ambient (12 μmol L–1) CO2. The time needed for cells grown at high CO2 (126 μmol L–1) to fully develop photosynthetic CO2 affinity at ambient CO2 was approximately 2 h, but acclimation to low or very low CO2 levels (3 and 1.3 μmol L–1, respectively) took more than 10 h. Cells grown at low CO2 (3 μmol L–1) required approximately 10 h for repression of all photosynthetic CO2 affinity when transferred to ambient or high CO2 (12 or 126 μmol L–1, respectively), and more than 10 h at very high CO2 (392 μmol L–1).

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