Heskel Mary

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Heskel
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  • Preprint
    Macromolecular rate theory (MMRT) provides a thermodynamics rationale to underpin the convergent temperature response in plant leaf respiration
    ( 2017-10) Liang, Liyin L. ; Arcus, Vickery ; Heskel, Mary ; O'Sullivan, Odhran S. ; Weerasinghe, Lasantha K. ; Creek, Danielle ; Egerton, John J. G. ; Tjoelker, Mark ; Atkin, Owen K. ; Schipper, Louis A.
    Temperature is a crucial factor in determining the rates of ecosystem processes, e.g. leaf respiration (R) − the flux of plant respired CO2 from leaves to the atmosphere. Generally, R increases exponentially with temperature and formulations such as the Arrhenius equation are widely used in earth system models. However, experimental observations have shown a consequential and consistent departure from an exponential increase in R. What are the principles that underlie these observed patterns? Here, we demonstrate that macromolecular rate theory (MMRT), based on transition state theory for enzyme-catalyzed kinetics, provides a thermodynamic explanation for the observed departure and the convergent temperature response of R using a global database. Three meaningful parameters emerge from MMRT analysis: the temperature at which the rate of respiration would theoretically reach a maximum (the optimum temperature, Topt), the temperature at which the respiration rate is most sensitive to changes in temperature (the inflection temperature, Tinf) and the overall curvature of the log(rate) versus temperature plot (the change in heat capacity for the system, ∆Cp‡). On average the highest potential enzyme-catalyzed rates of respiratory enzymes for R is predicted to occur at 67.0±1.2 °C and the maximum temperature sensitivity at 41.4±0.7 °C from MMRT. The average curvature (average negative ∆Cp‡) was -1.2±0.1 kJ.mol-1K-1. Interestingly, Topt, Tinf and ∆Cp‡ appear insignificantly different across biomes and plant functional types (PFTs), suggesting that thermal response of respiratory enzymes in leaves could be conserved. The derived parameters from MMRT can serve as thermal traits for plant leaves that represents the collective temperature response of metabolic respiratory enzymes and could be useful to understand regulations of R under a warmer climate. MMRT extends the classic transition state theory to enzyme-catalyzed reactions and provides an accurate and mechanistic model for the short-term temperature response of R around the globe.
  • Article
    Implications of improved representations of plant respiration in a changing climate
    (Nature Publishing Group, 2017-11-17) Huntingford, Chris ; Atkin, Owen K. ; Martinez-de la Torre, Alberto ; Mercado, Lina M. ; Heskel, Mary ; Harper, Anna B. ; Bloomfield, Keith J. ; O'Sullivan, Odhran S. ; Reich, Peter B. ; Wythers, Kirk R. ; Butler, Ethan E. ; Chen, Ming ; Griffin, Kevin L. ; Meir, Patrick ; Tjoelker, Mark ; Turnbull, Matthew H. ; Sitch, Stephen ; Wiltshire, Andrew J. ; Malhi, Yadvinder
    Land-atmosphere exchanges influence atmospheric CO2. Emphasis has been on describing photosynthetic CO2 uptake, but less on respiration losses. New global datasets describe upper canopy dark respiration (Rd) and temperature dependencies. This allows characterisation of baseline Rd, instantaneous temperature responses and longer-term thermal acclimation effects. Here we show the global implications of these parameterisations with a global gridded land model. This model aggregates Rd to whole-plant respiration Rp, driven with meteorological forcings spanning uncertainty across climate change models. For pre-industrial estimates, new baseline Rd increases Rp and especially in the tropics. Compared to new baseline, revised instantaneous response decreases Rp for mid-latitudes, while acclimation lowers this for the tropics with increases elsewhere. Under global warming, new Rd estimates amplify modelled respiration increases, although partially lowered by acclimation. Future measurements will refine how Rd aggregates to whole-plant respiration. Our analysis suggests Rp could be around 30% higher than existing estimates.
  • Preprint
    Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest
    ( 2016-11) Yang, Hualei ; Yang, Xi ; Zhang, Yongguang ; Heskel, Mary ; Lu, Xiaoliang ; Munger, J. William ; Sun, Shucun ; Tang, Jianwu
    Accurate estimation of terrestrial gross primary productivity (GPP) remains a challenge despite its importance in the global carbon cycle. Chlorophyll fluorescence (ChlF) has been recently adopted to understand photosynthesis and its response to the environment, particularly with remote sensing data. However, it remains unclear how ChlF and photosynthesis are linked at different spatial scales across the growing season. We examined seasonal relationships between ChlF and photosynthesis at the leaf, canopy, and ecosystem scales, and explored how leaf-level ChlF was linked with canopy-scale solar induced chlorophyll fluorescence (SIF) in a temperate deciduous forest at Harvard Forest, Massachusetts, USA. Our results show that ChlF captured the seasonal variations of photosynthesis with significant linear relationships between ChlF and photosynthesis across the growing season over different spatial scales (R2=0.73, 0.77 and 0.86 at leaf, canopy and satellite scales, respectively; p<0.0001). We developed a model to estimate GPP from the tower-based measurement of SIF and leaf-level ChlF parameters. The estimation of GPP from this model agreed well with flux tower observations of GPP (R2=0.68; p<0.0001), demonstrating the potential of SIF for modeling GPP. At the leaf scale, we found that leaf Fq’/Fm’, the fraction of absorbed photons that are used for photochemistry for a light adapted measurement from a pulse amplitude modulation fluorometer, was the best leaf fluorescence parameter to correlate with canopy-SIF yield (SIF/APAR, R2=0.79; p<0.0001). We also found that canopy-SIF and SIF-derived GPP (GPPSIF) were strongly correlated to leaf-level biochemistry and canopy structure, including chlorophyll content (R2=0.65 for canopy-GPPSIF and chlorophyll content; p<0.0001), leaf area index (LAI) (R2=0.35 for canopy-GPPSIF and LAI; p<0.0001), and normalized difference vegetation index (NDVI) (R2=0.36 for canopy-GPPSIF and NDVI; p<0.0001). Our results suggest that ChlF can be a powerful tool to track photosynthetic rates at leaf, canopy, and ecosystem scales.
  • Article
    Seasonal variations of leaf and canopy properties tracked by ground-based NDVI imagery in a temperate forest
    (Nature Publishing Group, 2017-04-28) Yang, Hualei ; Yang, Xi ; Heskel, Mary ; Sun, Shucun ; Tang, Jianwu
    Changes in plant phenology affect the carbon flux of terrestrial forest ecosystems due to the link between the growing season length and vegetation productivity. Digital camera imagery, which can be acquired frequently, has been used to monitor seasonal and annual changes in forest canopy phenology and track critical phenological events. However, quantitative assessment of the structural and biochemical controls of the phenological patterns in camera images has rarely been done. In this study, we used an NDVI (Normalized Difference Vegetation Index) camera to monitor daily variations of vegetation reflectance at visible and near-infrared (NIR) bands with high spatial and temporal resolutions, and found that the infrared camera based NDVI (camera-NDVI) agreed well with the leaf expansion process that was measured by independent manual observations at Harvard Forest, Massachusetts, USA. We also measured the seasonality of canopy structural (leaf area index, LAI) and biochemical properties (leaf chlorophyll and nitrogen content). We found significant linear relationships between camera-NDVI and leaf chlorophyll concentration, and between camera-NDVI and leaf nitrogen content, though weaker relationships between camera-NDVI and LAI. Therefore, we recommend ground-based camera-NDVI as a powerful tool for long-term, near surface observations to monitor canopy development and to estimate leaf chlorophyll, nitrogen status, and LAI.
  • Article
    Differential physiological responses to environmental change promote woody shrub expansion
    (John Wiley & Sons, 2013-03-13) Heskel, Mary ; Greaves, Heather ; Kornfeld, Ari ; Gough, Laura ; Atkin, Owen K. ; Turnbull, Matthew H. ; Shaver, Gaius R. ; Griffin, Kevin L.
    Direct and indirect effects of warming are increasingly modifying the carbon-rich vegetation and soils of the Arctic tundra, with important implications for the terrestrial carbon cycle. Understanding the biological and environmental influences on the processes that regulate foliar carbon cycling in tundra species is essential for predicting the future terrestrial carbon balance in this region. To determine the effect of climate change impacts on gas exchange in tundra, we quantified foliar photosynthesis (Anet), respiration in the dark and light (RD and RL, determined using the Kok method), photorespiration (PR), carbon gain efficiency (CGE, the ratio of photosynthetic CO2 uptake to total CO2 exchange of photosynthesis, PR, and respiration), and leaf traits of three dominant species – Betula nana, a woody shrub; Eriophorum vaginatum, a graminoid; and Rubus chamaemorus, a forb – grown under long-term warming and fertilization treatments since 1989 at Toolik Lake, Alaska. Under warming, B. nana exhibited the highest rates of Anet and strongest light inhibition of respiration, increasing CGE nearly 50% compared with leaves grown in ambient conditions, which corresponded to a 52% increase in relative abundance. Gas exchange did not shift under fertilization in B. nana despite increases in leaf N and P and near-complete dominance at the community scale, suggesting a morphological rather than physiological response. Rubus chamaemorus, exhibited minimal shifts in foliar gas exchange, and responded similarly to B. nana under treatment conditions. By contrast, E. vaginatum, did not significantly alter its gas exchange physiology under treatments and exhibited dramatic decreases in relative cover (warming: −19.7%; fertilization: −79.7%; warming with fertilization: −91.1%). Our findings suggest a foliar physiological advantage in the woody shrub B. nana that is further mediated by warming and increased soil nutrient availability, which may facilitate shrub expansion and in turn alter the terrestrial carbon cycle in future tundra environments.
  • Article
    A gradient of nutrient enrichment reveals nonlinear impacts of fertilization on Arctic plant diversity and ecosystem function
    (John Wiley & Sons, 2017-03-22) Prager, Case M. ; Naeem, Shahid ; Boelman, Natalie ; Eitel, Jan U. H. ; Greaves, Heather ; Heskel, Mary ; Magney, Troy ; Menge, Duncan N. L. ; Vierling, Lee ; Griffin, Kevin L.
    Rapid environmental change at high latitudes is predicted to greatly alter the diversity, structure, and function of plant communities, resulting in changes in the pools and fluxes of nutrients. In Arctic tundra, increased nitrogen (N) and phosphorus (P) availability accompanying warming is known to impact plant diversity and ecosystem function; however, to date, most studies examining Arctic nutrient enrichment focus on the impact of relatively large (>25x estimated naturally occurring N enrichment) doses of nutrients on plant community composition and net primary productivity. To understand the impacts of Arctic nutrient enrichment, we examined plant community composition and the capacity for ecosystem function (net ecosystem exchange, ecosystem respiration, and gross primary production) across a gradient of experimental N and P addition expected to more closely approximate warming-induced fertilization. In addition, we compared our measured ecosystem CO2 flux data to a widely used Arctic ecosystem exchange model to investigate the ability to predict the capacity for CO2 exchange with nutrient addition. We observed declines in abundance-weighted plant diversity at low levels of nutrient enrichment, but species richness and the capacity for ecosystem carbon uptake did not change until the highest level of fertilization. When we compared our measured data to the model, we found that the model explained roughly 30%–50% of the variance in the observed data, depending on the flux variable, and the relationship weakened at high levels of enrichment. Our results suggest that while a relatively small amount of nutrient enrichment impacts plant diversity, only relatively large levels of fertilization—over an order of magnitude or more than warming-induced rates—significantly alter the capacity for tundra CO2 exchange. Overall, our findings highlight the value of measuring and modeling the impacts of a nutrient enrichment gradient, as warming-related nutrient availability may impact ecosystems differently than single-level fertilization experiments.