A model analysis of climate and CO2 controls on tree growth and carbon allocation in a semi-arid woodland

Many studies have failed to show an increase in the radial growth of trees in response to increasing atmospheric CO2 concentration [CO2] despite the expected enhancement of photosynthetic rates and water-use efficiency at high [CO2]. A global light use efficiency model of photosynthesis, coupled with a generic carbon allocation and tree-growth model based on mass balance and tree geometry principles, was used to simulate annual ring-width variations for the gymnosperm Callitris columellaris in the semi-arid Great Western Woodlands, Western Australia, over the past 100 years. Parameter values for the tree-growth model were derived from independent observations except for sapwood specific respiration rate, fine-root turnover time, fine-root specific respiration rate and the ratio of fine-root mass to foliage area (ζ), which were calibrated to the ring-width measurements by Bayesian optimization. This procedure imposed a strong constraint on ζ. Modelled and observed ring-widths showed quantitatively similar, positive responses to total annual photosynthetically active radiation and soil moisture, and similar negative responses to vapour pressure deficit. The model also produced enhanced radial growth in response to increasing [CO2] during recent decades, but the data do not show this. Recalibration in moving 30-year time windows produced temporal shifts in the estimated values of ζ, including an increase by ca 12% since the 1960s, and eliminated the [CO2]-induced increase in radial growth. The potential effect of CO2 on ring-width was thus shown to be small compared to effects of climate variability even in this semi-arid climate. It could be counteracted in the model by a modest allocation shift, as has been observed in field experiments with raised [CO2]

Leaf nitrogen from first principles: field evidence for adaptive variation with climate

Nitrogen content per unit leaf area (Narea) is a key variable in plant functional ecology and biogeochemistry. Narea comprises a structural component, which scales with leaf mass per area (LMA), and a metabolic component, which scales with Rubisco capacity. The co-ordination hypothesis, as implemented in LPJ and related global vegetation models, predicts that Rubisco capacity should be directly proportional to irradiance but should decrease with increases in ci : ca and temperature because the amount of Rubisco required to achieve a given assimilation rate declines with increases in both. We tested these predictions using LMA, leaf δ13C, and leaf N measurements on complete species assemblages sampled at sites on a north–south transect from tropical to temperate Australia. Partial effects of mean canopy irradiance, mean annual temperature, and ci : ca (from δ13C) on Narea were all significant and their directions and magnitudes were in line with predictions. Over 80 % of the variance in community-mean (ln) Narea was accounted for by these predictors plus LMA. Moreover, Narea could be decomposed into two components, one proportional to LMA (slightly steeper in N-fixers), and the other to Rubisco capacity as predicted by the co-ordination hypothesis. Trait gradient analysis revealed ci : ca to be perfectly plastic, while species turnover contributed about half the variation in LMA and Narea. Interest has surged in methods to predict continuous leaf-trait variation from environmental factors, in order to improve ecosystem models. Coupled carbon–nitrogen models require a method to predict Narea that is more realistic than the widespread assumptions that Narea is proportional to photosynthetic capacity, and/or that Narea (and photosynthetic capacity) are determined by N supply from the soil. Our results indicate that Narea has a useful degree of predictability, from a combination of LMA and ci : ca – themselves in part environmentally determined – with Rubisco activity, as predicted from local growing conditions. This finding is consistent with a "plant-centred" approach to modelling, emphasizing the adaptive regulation of traits. Models that account for biodiversity will also need to partition community-level trait variation into components due to phenotypic plasticity and/or genotypic differentiation within species vs. progressive species replacement, along environmental gradients. Our analysis suggests that variation in Narea is about evenly split between these two modes.Ning Dong, Iain Colin Prentice, Bradley J. Evans, Stefan Caddy-Retalic, Andrew J. Lowe and Ian J. Wrigh

Bridging Drought Experiment and Modeling: Representing the Differential Sensitivities of Leaf Gas Exchange to Drought

Global climate change is expected to increase drought duration and intensity in certain regions while increasing rainfall in others. The quantitative consequences of increased drought for ecosystems are not easy to predict. Process-based models must be informed by experiments to determine the resilience of plants and ecosystems from different climates. Here, we demonstrate what and how experimentally derived quantitative information can improve the representation of stomatal and non-stomatal photosynthetic responses to drought in large-scale vegetation models. In particular, we review literature on the answers to four key questions: (1) Which photosynthetic processes are affected under short-term drought? (2) How do the stomatal and non-stomatal responses to short-term drought vary among species originating from different hydro-climates? (3) Do plants acclimate to prolonged water stress, and do mesic and xeric species differ in their degree of acclimation? (4) Does inclusion of experimentally based plant functional type specific stomatal and non-stomatal response functions to drought help Land Surface Models to reproduce key features of ecosystem responses to drought? We highlighted the need for evaluating model representations of the fundamental eco-physiological processes under drought. Taking differential drought sensitivity of different vegetation into account is necessary for Land Surface Models to accurately model drought responses, or the drought impacts on vegetation in drier environments may be over-estimated

Precipitation scaling with temperature in warm and cold climates: an analysis of CMIP5 simulations

We investigate the scaling between precipitation and temperature changes in warm and cold climates using six models that have simulated the response to both increased CO2 and Last Glacial Maximum (LGM) boundary conditions. Globally, precipitation increases in warm climates and decreases in cold climates by between 1.5%/°C and 3%/°C. Precipitation sensitivity to temperature changes is lower over the land than over the ocean and lower over the tropical land than over the extratropical land, reflecting the constraint of water availability. The wet tropics get wetter in warm climates and drier in cold climates, but the changes in dry areas differ among models. Seasonal changes of tropical precipitation in a warmer world also reflect this “rich get richer” syndrome. Precipitation seasonality is decreased in the cold-climate state. The simulated changes in precipitation per degree temperature change are comparable to the observed changes in both the historical period and the LGM

Peatlands and Climate Change

This is the author's manuscript version and this version is free to view and download for personal use only. Not for re-distribution, re-sale or use in derivative works.This material is forthcoming in Peatland Restoration and Ecosystem Services Science, Policy and Practice, 9781107619708, © Cambridge University PressThe fundamental reason for the presence of peatlands is a positive balance between plant production and decomposition. Organic matter accumulates in these systems because prolonged waterlogged conditions result in soil anoxia (i.e., exclusion of oxygen), and under these conditions decomposition rates can be lower than those of primary production. Climate therefore plays an important role in peat accumulation, both directly by affecting productivity and decomposition processes, and indirectly through its effects on hydrology/water balance and vegetation (for a summary, refer to Yu, Beilman & Jones 2009). Climate provides broad-scale constraints or controls on peatland extent, types and vegetation, and ultimately, ecosystem functioning, carbon accumulation, greenhouse gas exchange and all of the other ecosystem services that peatlands provide. Peatlands can play a vital role in helping society mitigate and adapt to climate change, because of their carbon and water regulating functions, while at the same time, the climate sensitivity of peatlands makes them potentially vulnerable to future global warming and changes in spatial and temporal patterns of precipitation, especially if they are in a degraded state. Climate change is likely to alter the hydrology and soil temperature of peatlands, with far- reaching consequences for their biodiversity, ecology and biogeochemistry. Their involvement in the global carbon cycle will also be affected, with the possibility of drier conditions allowing peatland erosion and increases in CO2 emissions that would result in a positive feedback to climate change (Turetsky 2010). This highlights all the more the need for restoration to ensure peatlands are resilient to change so that they continue to deliver ecosystem services for human well-being. This chapter describes the interactions between climate and peatlands, in three sections. The first section explains how present climate influences peatlands, by documenting how climate limits peatland geographical extent globally, and how bioclimatic envelope models can predict peatland extent. We indicate how each type of peatland is linked to a specific climate range, and introduce the concept of ecosystem function in relation to climate. The second section looks into the past. It describes how peat preserves a record of past climates and environmental conditions that can be deciphered to reveal the history of peatland vegetation, hydrology and carbon accumulation changes in relation to past changes in climate. We highlight lessons that can be learned from the palaeorecord preserved in peat. The final section discusses the potential effects of present and future climate change on peatlands, their extent, carbon accumulation rates, fire frequency, water table and greenhouse gas exchanges. We also consider how increases in sea level and CO2 concentration, and decreases in the extent of permafrost, are likely to affect peatlands

Changes in biomass allocation buffer low CO2 effects on tree growth during the last glaciation

Isotopic measurements on junipers growing in southern California during the last glacial, when the ambient atmospheric [CO2] (ca) was ~180 ppm, show the leaf-internal [CO2] (ci) was approaching the modern CO2 compensation point for C3 plants. Despite this, stem growth rates were similar to today. Using a coupled light-use efficiency and tree growth model, we show that it is possible to maintain a stable ci/ca ratio because both vapor pressure deficit and temperature were decreased under glacial conditions at La Brea, and these have compensating effects on the ci/ca ratio. Reduced photorespiration at lower temperatures would partly mitigate the effect of low ci on gross primary production, but maintenance of present-day radial growth also requires a ~27% reduction in the ratio of fine root mass to leaf area. Such a shift was possible due to reduced drought stress under glacial conditions at La Brea. The necessity for changes in allocation in response to changes in [CO2] is consistent with increased below-ground allocation, and the apparent homoeostasis of radial growth, as ca increases today

Bioclimatic envelope model of climate change impacts on blanket peatland distribution in Great Britain

Copyright © 2010 Inter-Research.Publisher's version of record available via: doi: 10.3354/cr00911Blanket peatlands are rain-fed mires that cover the landscape almost regardless of topography. The geographical extent of this type of peatland is highly sensitive to climate. We applied a global process-based bioclimatic envelope model, PeatStash, to predict the distribution of British blanket peatlands. The model captures the present areal extent (Kappa = 0.77) and is highly sensitive to both temperature and precipitation changes. When the model is run using the UKCIP02 climate projections for the time periods 2011-2040, 2041-2070 and 2071-2100, the geographical distribution of blanket peatlands gradually retreats towards the north and the west. In the UKCIP02 high emissions scenario for 2071-2100, the blanket peatland bioclimatic space is ∼84% smaller than contemporary conditions (1961-1990); only parts of the west of Scotland remain inside this space. Increasing summer temperature is the main driver of the projected changes in areal extent. Simulations using 7 climate model outputs resulted in generally similar patterns of declining aereal extent of the bioclimatic space, although differing in degree. The results presented in this study should be viewed as a first step towards understanding the trends likely to affect the blanket peatland distribution in Great Britain. The eventual fate of existing blanket peatlands left outside their bioclimatic space remains uncertain. © Inter-Research 2010 .Environment AgencyNatural Environment Research Council (NERC)Royal Societ

Plant respiration : Controlled by photosynthesis or biomass?

Abstract Two simplifying hypotheses have been proposed for whole-plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first-principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carryover of fixed carbon between years, while the second implies far too great an increase in respiration during stand development ? leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration is not linearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.Peer reviewe

Thermal acclimation of leaf photosynthetic traits in an evergreen woodland, consistent with the coordination hypothesis

Ecosystem models commonly assume that key photosynthetic traits, such as carboxylation capacity measured at a standard temperature, are constant in time. The temperature responses of modelled photosynthetic or respiratory rates then depend entirely on enzyme kinetics. Optimality considerations, however, suggest this assumption may be incorrect. The coordination hypothesis (that Rubisco- and electron-transport-limited rates of photosynthesis are co-limiting under typical daytime conditions) predicts, instead, that carboxylation (Vcmax) capacity should acclimate so that it increases somewhat with growth temperature but less steeply than its instantaneous response, implying that Vcmax when normalized to a standard temperature (e.g. 25 °C) should decline with growth temperature. With additional assumptions, similar predictions can be made for electron-transport capacity (Jmax) and mitochondrial respiration in the dark (Rdark). To explore these hypotheses, photosynthetic measurements were carried out on woody species during the warm and the cool seasons in the semi-arid Great Western Woodlands, Australia, under broadly similar light environments. A consistent proportionality between Vcmax and Jmax was found across species. Vcmax, Jmax and Rdark increased with temperature in most species, but their values standardized to 25 °C declined. The ci : ca ratio increased slightly with temperature. The leaf N  :  P ratio was lower in the warm season. The slopes of the relationships between log-transformed Vcmax and Jmax and temperature were close to values predicted by the coordination hypothesis but shallower than those predicted by enzyme kinetics.This research was funded by the Terrestrial Ecosystem Research Network (TERN), Macquarie University and the Australian National University. Henrique Fürstenau Togashi was supported by an international Macquarie University International Research Scholarship (iMQRES). Iain Colin Prentice, Bradley John Evans, and Henrique Fürstenau Togashi were funded by the Ecosystem Modelling and Scaling Infrastructure (eMAST, part of TERN). TERN and eMAST have been supported by the Australian Government through the National Collaborative Research Infrastructure Strategy (NCRIS). Owen Atkin acknowledges the support of the Australian Research Council (DP130101252 and CE140100008)