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Elevated carbon dioxide (eCO2) in the atmosphere increases forest biomass productivity but only where soil nutrients, particularly nitrogen (N) and phosphorus (P), are not limiting growth. eCO2, in turn, can impact rhizosphere nutrient availability. Our current understanding of nutrient cycling under eCO2 is mainly derived from surface soil, leaving mechanisms of the impact of eCO2 on rhizosphere nutrient availability at deeper depths unexplored. To investigate the influence of eCO2 on nutrient availability in soil at depth, we studied various C, N, and P pools (extractable, microbial biomass, total soil C and N, and mineral-associated P) and nutrient cycling processes (enzyme activity and gross N mineralisation) associated with C, N, and P cycling in both bulk and rhizosphere soil at different depths at the Free Air CO2 enrichment facility in a native Australian mature Eucalyptus woodland (EucFACE) on a nutrient-poor soil. We found decreasing nutrient availability and gross N mineralisation with depth; however, this depth-associated decrease was reduced under eCO2, which we suggest is due to enhanced root influence. Increases in available PO43-, adsorbed P, and the C : N and C : P ratio of enzyme activity with depth were observed. We conclude that the influences of roots and of eCO2 can affect available nutrient pools and processes well beyond the surface soil of a mature forest ecosystem. Our findings indicate a faster recycling of nutrients in the rhizosphere, rather than additional nutrients becoming available through soil organic matter (SOM) decomposition. If the plant growth response to eCO2 is reduced by the constraints of nutrient limitations, then the current results would call to question the potential for mature tree ecosystems to fix more C as biomass in response to eCO2. Future studies should address how accessible the available nutrients at depth are to deeply rooted plants and if fast recycling of nutrients is a meaningful contribution to biomass production and the accumulation of soil C in response to eCO2.

Land surface models underpin coupled climate model projections of droughts and heatwaves. However, the lack of simultaneous observations of individual components of evapotranspiration, concurrent with root-zone soil moisture, has limited previous model evaluations. Here, we use a comprehensive set of observations from a water-limited site in southeastern Australia including both evapotranspiration and soil moisture to a depth of 4.5 m to evaluate the Community Atmosphere-Biosphere Land Exchange (CABLE) land surface model. We demonstrate that alternative process representations within CABLE had the capacity to improve simulated evapotranspiration, but not necessarily soil moisture dynamics–highlighting problems of model evaluations against water fluxes alone. Our best simulation was achieved by resolving a soil evaporation bias, using a more realistic initialisation of the groundwater aquifer state and higher vertical soil resolution informed by observed soil properties, and further calibrating soil hydraulic conductivity. Despite these improvements, the role of the empirical soil moisture stress function in influencing the simulated water fluxes remained important: using a site-calibrated function reduced the soil water stress on plants by 36 % during drought and 23 % at other times. These changes in CABLE not only improve the seasonal cycle of evapotranspiration but also affect the latent and sensible heat fluxes during droughts and heatwaves. The range of parameterisations tested led to differences of ∼150 W m−2 in the simulated latent heat flux during a heatwave, implying a strong impact of parameterisations on the capacity for evaporative cooling and feedbacks to the boundary layer (when coupled). Overall, our results highlight the opportunity to advance the capability of land surface models to capture water cycle processes, particularly during meteorological extremes, when sufficient observations of both evapotranspiration fluxes and soil moisture profiles are available.

The response of mature forest ecosystems to a rising atmospheric carbon dioxide concentration (Ca) is a major uncertainty in projecting the future trajectory of the Earth's climate. Although leaf-level net photosynthesis is typically stimulated by exposure to elevated Ca (eCa), it is unclear how this stimulation translates into carbon cycle responses at the ecosystem scale. Here we estimate a key component of the carbon cycle, the gross primary productivity (GPP), of a mature native eucalypt forest exposed to free-air CO2 enrichment (the EucFACE experiment). In this experiment, light-saturated leaf photosynthesis increased by 19 % in response to a 38 % increase in Ca. We used the process-based forest canopy model, MAESPA, to upscale these leaf-level measurements of photosynthesis with canopy structure to estimate the GPP and its response to eCa. We assessed the direct impact of eCa, as well as the indirect effect of photosynthetic acclimation to eCa and variability among treatment plots using different model scenarios. At the canopy scale, MAESPA estimated a GPP of 1574 g C m−2 yr−1 under ambient conditions across 4 years and a direct increase in the GPP of +11 % in response to eCa. The smaller canopy-scale response simulated by the model, as compared with the leaf-level response, could be attributed to the prevalence of RuBP regeneration limitation of leaf photosynthesis within the canopy. Photosynthetic acclimation reduced this estimated response to 10 %. After taking the baseline variability in the leaf area index across plots in account, we estimated a field GPP response to eCa of 6 % with a 95 % confidence interval (−2 %, 14 %). These findings highlight that the GPP response of mature forests to eCa is likely to be considerably lower than the response of light-saturated leaf photosynthesis. Our results provide an important context for interpreting the eCa responses of other components of the ecosystem carbon cycle.

Elevated carbon dioxide (CO2) can increase plant growth, but the magnitude of this CO2 fertilization effect is modified by soil nutrient availability. Predicting how nutrient availability affects plant responses to elevated CO2 is a key consideration for ecosystem models, and many modeling groups have moved to, or are moving towards, incorporating nutrient limitation in their models. The choice of assumptions to represent nutrient cycling processes has a major impact on model predictions, but it can be difficult to attribute outcomes to specific assumptions in complex ecosystem simulation models. Here we revisit the quasi-equilibrium analytical framework introduced by Comins and McMurtrie (1993) and explore the consequences of specific model assumptions for ecosystem net primary productivity (NPP). We review the literature applying this framework to plant–soil models and then analyze the effect of several new assumptions on predicted plant responses to elevated CO2. Examination of alternative assumptions for plant nitrogen uptake showed that a linear function of the mineral nitrogen pool or a linear function of the mineral nitrogen pool with an additional saturating function of root biomass yield similar CO2 responses at longer timescales (>5 years), suggesting that the added complexity may not be needed when these are the timescales of interest. In contrast, a saturating function of the mineral nitrogen pool with linear dependency on root biomass yields no soil nutrient feedback on the very-long-term (>500 years), near-equilibrium timescale, meaning that one should expect the model to predict a full CO2 fertilization effect on production. Secondly, we show that incorporating a priming effect on slow soil organic matter decomposition attenuates the nutrient feedback effect on production, leading to a strong medium-term (5–50 years) CO2 response. Models incorporating this priming effect should thus predict a strong and persistent CO2 fertilization effect over time. Thirdly, we demonstrate that using a “potential NPP” approach to represent nutrient limitation of growth yields a relatively small CO2 fertilization effect across all timescales. Overall, our results highlight the fact that the quasi-equilibrium analytical framework is effective for evaluating both the consequences and mechanisms through which different model assumptions affect predictions. To help constrain predictions of the future terrestrial carbon sink, we recommend the use of this framework to analyze likely outcomes of new model assumptions before introducing them to complex model structures.