CO2 modulation of the rates of photosynthesis and light-dependent O2 consumption in Trichodesmium.

As atmospheric CO2 concentrations increase, so too does the dissolved CO2 and HCO3- concentrations in the world's oceans. There are still many uncertainties regarding the biological response of key groups of organisms to these changing conditions, which is crucial for predicting future species distributions, primary productivity rates, and biogeochemical cycling. In this study, we established the relationship between gross photosynthetic O2 evolution and light-dependent O2 consumption in Trichodesmium erythraeum IMS101 acclimated to three targeted pCO2 concentrations (180 µmol mol-1=low-CO2, 380 µmol mol-1=mid-CO2, and 720 µmol mol-1=high-CO2). We found that biomass- (carbon) specific, light-saturated maximum net O2 evolution rates (PnC,max) and acclimated growth rates increased from low- to mid-CO2, but did not differ significantly between mid- and high-CO2. Dark respiration rates were five times higher than required to maintain cellular metabolism, suggesting that respiration provides a substantial proportion of the ATP and reductant for N2 fixation. Oxygen uptake increased linearly with gross O2 evolution across light intensities ranging from darkness to 1100 µmol photons m-2 s-1. The slope of this relationship decreased with increasing CO2, which we attribute to the increased energetic cost of operating the carbon-concentrating mechanism at lower CO2 concentrations. Our results indicate that net photosynthesis and growth of T. erythraeum IMS101 would have been severely CO2 limited at the last glacial maximum, but that the direct effect of future increases of CO2 may only cause marginal increases in growth.

The physiological cost of diazotrophy for Trichodesmium erythraeum IMS101.

Trichodesmium plays a significant role in the oligotrophic oceans, fixing nitrogen in an area corresponding to half of the Earth's surface, representing up to 50% of new production in some oligotrophic tropical and subtropical oceans. Whilst Trichodesmium blooms at the surface exhibit a strong dependence on diazotrophy, colonies at depth or at the surface after a mixing event could be utilising additional N-sources. We conducted experiments to establish how acclimation to varying N-sources affects the growth, elemental composition, light absorption coefficient, N2 fixation, PSII electron transport rate and the relationship between net and gross photosynthetic O2 exchange in T. erythraeum IMS101. To do this, cultures were acclimated to growth medium containing NH4+ and NO3- (replete concentrations) or N2 only (diazotrophic control). The light dependencies of O2 evolution and O2 uptake were measured using membrane inlet mass spectrometry (MIMS), while PSII electron transport rates were measured from fluorescence light curves (FLCs). We found that at a saturating light intensity, Trichodesmium growth was ~ 10% and 13% lower when grown on N2 than with NH4+ and NO3-, respectively. Oxygen uptake increased linearly with net photosynthesis across all light intensities ranging from darkness to 1100 µmol photons m-2 s-1. The maximum rates and initial slopes of light response curves for C-specific gross and net photosynthesis and the slope of the relationship between gross and net photosynthesis increased significantly under non-diazotrophic conditions. We attribute these observations to a reduced expenditure of reductant and ATP for nitrogenase activity under non-diazotrophic conditions which allows NADPH and ATP to be re-directed to CO2 fixation and/or biosynthesis. The energy and reductant conserved through utilising additional N-sources could enhance Trichodesmium's productivity and growth and have major implications for its role in ocean C and N cycles.

Effect of light on the growth of non-nitrogen-fixing and nitrogen-fixing phytoplankton in an aquatic system.

We discuss a mathematical model of growth of two types of phytoplankton, non-nitrogen-fixing and nitrogen-fixing, that both require light in order to grow. We use general functional responses to represent the inhibitory effect their biomass has on the exposure to light. We give conditions for the existence and local stability of all of the possible steady-states (die out, single species survival, and coexistence). We derive conditions for global stability of the die out and single-species steady-states and for persistence of both species when the coexistence steady-state exists. Numerical investigation illustrates the qualitative dynamics demonstrating that even under constant environmental conditions, both stable intrinsic oscillatory behavior and a period doubling route to chaotic dynamics are possible. We also show that competitor-mediated coexistence can occur due to the positive feedback resulting from recycling by the nitrogen-fixing phytoplankton. To show the impact of seasonal change in water depth, we also allow the water depth to vary in an annual cycle and discuss echo blooms in this context.