Marco Borghetti

The human footprint in the carbon cycle of temperate and boreal forests

Temperate and boreal forests in the Northern Hemisphere cover an area of about 2 × 107 square kilometres and act as a substantial carbon sink (0.6–0.7 petagrams of carbon per year). Although forest expansion following agricultural abandonment is certainly responsible for an important fraction of this carbon sink activity, the additional effects on the carbon balance of established forests of increased atmospheric carbon dioxide, increasing temperatures, changes in management practices and nitrogen deposition are difficult to disentangle, despite an extensive network of measurement stations. The relevance of this measurement effort has also been questioned, because spot measurements fail to take into account the role of disturbances, either natural (fire, pests, windstorms) or anthropogenic (forest harvesting). Here we show that the temporal dynamics following stand-replacing disturbances do indeed account for a very large fraction of the overall variability in forest carbon sequestration. After the confounding effects of disturbance have been factored out, however, forest net carbon sequestration is found to be overwhelmingly driven by nitrogen deposition, largely the result of anthropogenic activities. The effect is always positive over the range of nitrogen deposition covered by currently available data sets, casting doubts on the risk of widespread ecosystem nitrogen saturation under natural conditions. The results demonstrate that mankind is ultimately controlling the carbon balance of temperate and boreal forests, either directly (through forest management) or indirectly (through nitrogen deposition).

Impact of long-term drought on xylem embolism and growth in Pinus halepensis Mill.

Abstract The present study was carried out to elucidate the response mechanisms of 50-year-old Pinus halepensis Mill. trees to a long-term and severe drought. The amount of water available to trees was artificially restricted for 12 months by covering the soil with a plastic roof. Over the short term a direct and rapid impact of drought was evident on the water relations and gas exchanges of trees: as the soil dried out in the Spring, there was a concurrent decrease of predawn water potential; transpiration was strongly reduced by stomatal closure. Seasonal changes in the water volume fractions of twig and stem xylem were observed and interpreted as the result of cavitation and refilling in the xylem. When droughted trees recovered to a more favourable water status, refilling of embolized xylem was observed; twig predawn water potentials were still negative in the period when the embolism was reversed in the twig xylem. A few months after the removal of the covering, no differences in whole plant hydraulic resistance were observed between droughted and control trees. Needle and shoot elongation and stem radial growth were considerably reduced in droughted trees; no strategy of trees to allocate carbon preferentially to the stem conducting tissues was apparent throughout the experiment. An after-effect of the drought on growth was observed.

Acclimation to changing light conditions of long-term shade-grown beech (Fagus sylvatica L.) seedlings of different geographic origins

Abstract Effects of changing light conditions on the ecophysiological condition behind survival were examined on beech from two different populations. Plants were grown in a greenhouse under simulated understorey and canopy gap light conditions. Upon exposure to high light maximum photosynthesis of shade-acclimated leaves increased followed by a reduction over several days to between high- and low-light control rates. In the reciprocal transfer, the decrease in maximum photosynthesis was rapid during the first 2–3 days and then levelled off to values comparable to low-light controls. Seedlings from Sicily (Madonie) showed generally higher maximum photosynthetic rates than those from Abetone. Leaf conductance varied in the same direction as photosynthesis in high- to low-light seedlings but to a lesser degree. Leaves grown under low light and exposed to high light experienced photoinhibition. The Abetone population was more susceptible to photoinhibitory damage than the seedlings from Sicily. Exposure to high light of shade-acclimated seedlings resulted in intermediate chlorophyll concentrations between levels of the high-light and low-light seedlings. Carotenoid concentration was unaffected by treatments. Seedlings grew more in high light, but had a lower leaf area ratio. Light-limited seedlings showed a shift in carbon allocation to foliage. Leaves formed in the new light regime maintained the same anatomy that had been developed before transfer. Seedlings from Sicily had thicker leaves than those of seedlings from Abetone. Seedlings from Abetone were found to be more susceptible to changing light conditions than seedlings from Sicily. We conclude that small forest gaps may represent a favorable environment for photosynthesis and growth of beech regeneration as a result of the limited ability of seedlings to acclimate to sudden increases in high irradiance and because of the moderate levels of light stress in small gaps.

Environmental Effects on Oxygen Isotope Enrichment of Leaf Water in Cotton Leaves

The oxygen isotope enrichment of bulk leaf water (Δb) was measured in cotton (Gossypium hirsutum) leaves to test the Craig-Gordon and Farquhar-Gan models under different environmental conditions. Δb increased with increasing leaf-to-air vapor pressure difference (VPd) as an overall result of the responses to the ratio of ambient to intercellular vapor pressures (ea/ei) and to stomatal conductance (gs). The oxygen isotope enrichment of lamina water relative to source water \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \(({\bar{{\Delta}}}_{1}),\) \end{document} which increased with increasing VPd, was estimated by mass balance between less enriched water in primary veins and enriched water in the leaf. The Craig-Gordon model overestimated Δb (and \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}),\) \end{document} as expected. Such discrepancies increased with increase in transpiration rate (E), supporting the Farquhar-Gan model, which gave reasonable predictions of Δb and \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}\) \end{document} with an L of 7.9 mm, much less than the total radial effective length Lr of 43 mm. The fitted values of L for \batchmode \documentclass[fleqn,10pt,legalpaper]{article} \usepackage{amssymb} \usepackage{amsfonts} \usepackage{amsmath} \pagestyle{empty} \begin{document} \({\bar{{\Delta}}}_{1}\) \end{document} of individual leaves showed little dependence on VPd and temperature, supporting the assumption that the Farquhar-Gan formulation is relevant and useful in describing leaf water isotopic enrichment.

Ecotypic variation of xylem embolism, phenological traits, growth parameters and allozyme characteristics in Fagus sylvatica.

Variation in late-winter xylem embolism, phenological traits, growth parameters, xylem anatomy and allozyme characteristics were examined in four populations of European beech (Fagus sylvatica) from different geographical origins in Italy. Ultrasonic acoustic emissions from plant stems were measured during winter. Late-winter xylem embolism was quantified before budburst. The timing of budburst was recorded and plant growth parameters were monitored during the spring. The dimension and density of xylem conduits were measured. The genetic variability was investigated using isoenzymes as genetical markers. Significant differences between populations in the rate of acoustic emissions, late-winter embolism, phenological traits and spring growth parameters were observed (...)

Assessing the effects of nitrogen deposition and climate on carbon isotope discrimination and intrinsic water-use efficiency of angiosperm and conifer trees under rising CO2 conditions.

The objective of this study is to globally assess the effects of atmospheric nitrogen deposition and climate, associated with rising levels of atmospheric CO2 , on the variability of carbon isotope discrimination (Δ(13) C), and intrinsic water-use efficiency (iWUE) of angiosperm and conifer tree species. Eighty-nine long-term isotope tree-ring chronologies, representing 23 conifer and 13 angiosperm species for 53 sites worldwide, were extracted from the literature, and used to obtain long-term time series of Δ(13) C and iWUE. Δ(13) C and iWUE were related to the increasing concentration of atmospheric CO2 over the industrial period (1850-2000) and to the variation of simulated atmospheric nitrogen deposition and climatic variables over the period 1950-2000. We applied generalized additive models and linear mixed-effects models to predict the effects of climatic variables and nitrogen deposition on Δ(13) C and iWUE. Results showed a declining Δ(13) C trend in the angiosperm and conifer species over the industrial period and a 16.1% increase of iWUE between 1850 and 2000, with no evidence that the increased rate was reduced at higher ambient CO2 values. The temporal variation in Δ(13) C supported the hypothesis of an active plant mechanism that maintains a constant ratio between intercellular and ambient CO2 concentrations. We defined linear mixed-effects models that were effective to describe the variation of Δ(13) C and iWUE as a function of a set of environmental predictors, alternatively including annual rate (Nrate ) and long-term cumulative (Ncum ) nitrogen deposition. No single climatic or atmospheric variable had a clearly predominant effect, however, Δ(13) C and iWUE showed complex dependent interactions between different covariates. A significant association of Nrate with iWUE and Δ(13) C was observed in conifers and in the angiosperms, and Ncum was the only independent term with a significant positive association with iWUE, although a multi-factorial control was evident in conifers.

Physiological and structural changes in response to altered precipitation regimes in a Mediterranean macchia ecosystem

Significant decrease in precipitation up to 15–20% has been observed in the Mediterranean area in the last two decades as a consequence of climate change. To simulate an analogous scenario, the precipitation regime was altered in replicated experimental plots in a Mediterranean macchia dominated by Arbutus unedo L. species. Two different levels of soil water content (SWC) were obtained during the summer: a mean value of 7% was obtained in water-depleted (D) plots by a partial (−20%) rain exclusion treatment using rain gutters; while a mean value of 14% in SWC was obtained in watered (W) plots supplying water by a sprinkler net. The physiological and structural changes were investigated over the course of two consecutive years by measurement of water potential, gas exchange leaf carbon isotopes, leaf pigments and growth. Apart from short-term responses, mainly related to the elastic response of stomatal conductance to soil water, a more long-lasting and significant acclimation to water availability was observed as a result of the increase in hydraulic resistance in the soil–plant continuum, which persisted even after the return to full water availability during the fall and winter. This response involved the permanent down-regulation of stomatal conductance and photosynthesis, accumulation of photo-protective pigments, as well as a reduction in shoot growth, leaf area index and an increase in shoot-bearing flowers in D plots. This acclimation response prevented the onset of any run-away damage thereby reducing the forest vulnerability to drought. Furthermore, the imposed drought induced a slight increase or no change in intrinsic water-use efficiency (WUEint), as a result of the parallel increase in stomatal and non-stomatal limitations; conversely integrated WUE (i.e., estimated from leaf carbon isotopes) was not affected by drought.

Assessing temporal variation of primary and ecosystem production in two Mediterranean forests using a modified 3-PG model

ContextForest ecosystem carbon uptake is heavily affected by increasing drought in the Mediterranean region.AimsThe objectives of this study were to assess the capacity of a modified 3-PG model to capture temporal variation in gross primary productivity (GPP), and ecosystem net carbon uptake (NEE) in two Mediterranean forest types.MethodsThe model was upgraded from a monthly (3-PG) to a daily time step (3-PGday), and a soil water balance routine was included to better represent soil water availability. The model was evaluated against seasonal GPP and NEE dynamics from eddy covariance measurements.ResultsSimulated and measured soil water content values were congruent throughout the study period for both forest types. 3-PGday effectively described the following: GPP and NEE seasonal patterns; the transition of forest ecosystems from carbon sink to carbon source; however, the model overestimated diurnal ecosystem respiration values and failed to predict ecosystem respiration peaks.ConclusionsThe model served as a rather effective tool to represent seasonal variation in gross primary productivity, and ecosystem net carbon uptake under Mediterranean drought-prone conditions. However, its semi-empirical nature and the simplicity inherent in the original model formulation are obstacles preventing the model working well for short-term daily predictions.

Ecologically implausible carbon response? Reply

Replying to: A. De Schrijver et al. 451, 10.1038/nature06578; W. de Vries et al. 451, 10.1038/nature06579 (2008)Nitrogen (N) deposition alters ecosystem function in several ways, with important effects on N leaching and water quality, as well as on interspecific competition and biodiversity. These changes have been attributed to ecosystem N saturation, defined as the alleviation of N limitations on rates of biological function. After an initial fertilization effect, N saturation has also been suggested to reduce plant function and growth, eventually leading to forest dieback. Although our observation of a substantial positive effect of N deposition on forest carbon (C) sequestration does not imply the absence of nitrate losses or other negative effects, as rightly stressed by De Schrijver et al., the sustained response observed demonstrates that the fear of a generalized forest decline in response to N fertilization could be overstated, at least within the rather broad N deposition range explored in our analysis. The nature of the observed response of forest C sequestration to N deposition, however, has been questioned outright by de Vries et al., who suggested that it could be an artefact resulting from the covariation between N deposition and other environmental variables. The arguments proposed against an overwhelming N effect, however, do not seem to stand up to close scrutiny.

Nutritional regulation in mixotrophic plants: new insights from Limodorum abortivum

Partially mycoheterotrophic (mixotrophic) plants gain carbon from both photosynthesis and their mycorrhizal fungi. This is considered an ancestral state in the evolution of full mycoheterotrophy, but little is known about this nutrition, and especially about the physiological balance between photosynthesis and fungal C gain. To investigate possible compensation between photosynthesis and mycoheterotrophy in the Mediterranean mixotrophic orchid Limodorum abortivum, fungal colonization was experimentally reduced in situ by fungicide treatment. We measured photosynthetic pigments of leaves, stems, and ovaries, as well as the stable C isotope compositions (a proxy for photosynthetic C gain) of seeds and the sizes of ovaries and seeds. We demonstrate that (1) in natural conditions, photosynthetic pigments are most concentrated in ovaries; (2) pigments and photosynthetic C increase in ovaries when fungal C supply is impaired, buffering C limitations and allowing the same development of ovaries and seeds as in natural conditions; and (3) responses to light of pigment and 13C contents in ovaries shift from null responses in natural conditions to responses typical of autotrophic plants in treated L. abortivum, demonstrating photoadaptation and enhanced use of light in the latter. L. abortivum thus preferentially feeds on fungi in natural conditions, but employs compensatory photosynthesis to buffer fungal C limitations and allow seed development.