Partitioning direct and indirect effects reveals the response of water-limited ecosystems to elevated CO2.

Increasing concentrations of atmospheric carbon dioxide are expected to affect carbon assimilation and evapotranspiration (ET), ultimately driving changes in plant growth, hydrology, and the global carbon balance. Direct leaf biochemical effects have been widely investigated, whereas indirect effects, although documented, elude explicit quantification in experiments. Here, we used a mechanistic model to investigate the relative contributions of direct (through carbon assimilation) and indirect (via soil moisture savings due to stomatal closure, and changes in leaf area index) effects of elevated CO2 across a variety of ecosystems. We specifically determined which ecosystems and climatic conditions maximize the indirect effects of elevated CO2 The simulations suggest that the indirect effects of elevated CO2 on net primary productivity are large and variable, ranging from less than 10% to more than 100% of the size of direct effects. For ET, indirect effects were, on average, 65% of the size of direct effects. Indirect effects tended to be considerably larger in water-limited ecosystems. As a consequence, the total CO2 effect had a significant, inverse relationship with the wetness index and was directly related to vapor pressure deficit. These results have major implications for our understanding of the CO2 response of ecosystems and for global projections of CO2 fertilization, because, although direct effects are typically understood and easily reproducible in models, simulations of indirect effects are far more challenging and difficult to constrain. Our findings also provide an explanation for the discrepancies between experiments in the total CO2 effect on net primary productivity.

Disentangling the net: concomitant xylem and over-bark size measurements reveal the phloem-generated turgor signal behind daytime stem swelling in the mangrove Avicennia marina.

Daytime stem shrinking is a well recorded phenomenon: trees dip into their internal water stores to meet imbalances between water supply and demand. Uncertainty surrounds the mechanisms behind the unusual pattern of daytime stem swelling, presented by species like the mangrove Avicennia marina (Forssk.) Vierh., and which is thought to originate in the osmotic adjustment of storage tissues. We performed on-xylem radius change (XRC) and whole-stem radius change (SRC) measurements with point dendrometers to ascertain if the swelling of the stem is due to an increase in the thickness of the phloem and inner-bark (BRC). We measured leaf water potential (ψleaf), sap flow, leaf turgor and microclimate to understand the coupling between BRC and transpiration-driven changes in XRC. Our results present direct evidence of the type of mechanism responsible for daytime stem swelling. Inner-bark thickness increase concurrent with XRC decrease, sap-flow increase and leaf turgor loss, reveals a phloem-generated turgor signal behind daytime stem swelling of tree stems. On-xylem measurements were highly heterogeneous due to the variability in the three dimensional fish-net wood structure of the stem of A. marina. As daytime decreases in XRC were compensated by BRC and correlated with conditions of high water demand, we suggest a potential role of daytime stem swelling in the hydraulic safety of A. marina.

Daytime stem swelling and seasonal reversal in the peristaltic depletion of stored water along the stem of Avicennia marina (Forssk.) Vierh.

Diurnal courses of stem radial water dynamics represent the sum of all internal and external conditions affecting tree water relations. Changes in stem radius due to early morning water depletion and night time refilling of storage tissues is generally well documented. This study seeks to understand the unusual daytime refilling of stem elastic storage tissues present in the mangrove species Avicennia marina (Forssk.) Vierh, which deviates from our traditional understanding of hydraulics in terrestrial trees. We explored the relationship of this pattern to other water-related physiological processes and environmental variables, and investigated the seasonal changes in the timing and time lags of peak swelling at different stem heights, in order to understand the 'peristaltic' depletion of internally stored water. Our findings show that daytime stem swelling occurs year-round, even on days when leaf water potentials dropped to values lower than -4 MPa. The amplitude of stem swelling was strongly positively correlated to daily light sums more often than to measures of water availability in air and soil, especially in winter. There was also a clear seasonal reversal in the timing and direction of the 'peristaltic' depletion of water along the stem, with an earlier onset of shrinking in the upper (median = 10:00 h) than in the lower stem (median = 12:00 h) in winter, but an earlier onset of shrinking in the lower (median = 08:00 h) than in the upper stem (median = 11:00 h) in summer. This time lag was closely correlated to daily temperature, with a clear switch in the direction of peristaltic stem shrinking at the start of the growing season. We propose that sugar loading/unloading and changes in source-sink activity play a role in the endogenous osmotic adjustment responsible for daytime stem swelling and the seasonal switch in the direction of peristaltic water storage depletion in A. marina.