No carbon storage in growth-limited trees in a semi-arid woodland.

Plant survival depends on a balance between carbon supply and demand. When carbon supply becomes limited, plants buffer demand by using stored carbohydrates (sugar and starch). During drought, NSCs (non-structural carbohydrates) may accumulate if growth stops before photosynthesis. This expectation is pervasive, yet few studies have combined simultaneous measurements of drought, photosynthesis, growth, and carbon storage to test this. Using a field experiment with mature trees in a semi-arid woodland, we show that growth and photosynthesis slow in parallel as [Formula: see text] declines, preventing carbon storage in two species of conifer (J. monosperma and P. edulis). During experimental drought, growth and photosynthesis were frequently co-limited. Our results point to an alternative perspective on how plants use carbon that views growth and photosynthesis as independent processes both regulated by water availability.

Oral famotidine versus placebo in non-hospitalised patients with COVID-19: a randomised, double-blind, data-intense, phase 2 clinical trial.

OBJECTIVE: We assessed whether famotidine improved inflammation and symptomatic recovery in outpatients with mild to moderate COVID-19. DESIGN: Randomised, double-blind, placebo-controlled, fully remote, phase 2 clinical trial (NCT04724720) enrolling symptomatic unvaccinated adult outpatients with confirmed COVID-19 between January 2021 and April 2021 from two US centres. Patients self-administered 80 mg famotidine (n=28) or placebo (n=27) orally three times a day for 14 consecutive days. Endpoints were time to (primary) or rate of (secondary) symptom resolution, and resolution of inflammation (exploratory). RESULTS: Of 55 patients in the intention-to-treat group (median age 35 years (IQR: 20); 35 women (64%); 18 African American (33%); 14 Hispanic (26%)), 52 (95%) completed the trial, submitting 1358 electronic symptom surveys. Time to symptom resolution was not statistically improved (p=0.4). Rate of symptom resolution was improved for patients taking famotidine (p<0.0001). Estimated 50% reduction of overall baseline symptom scores were achieved at 8.2 days (95% CI: 7 to 9.8 days) for famotidine and 11.4 days (95% CI: 10.3 to 12.6 days) for placebo treated patients. Differences were independent of patient sex, race or ethnicity. Five self-limiting adverse events occurred (famotidine, n=2 (40%); placebo, n=3 (60%)). On day 7, fewer patients on famotidine had detectable interferon alpha plasma levels (p=0.04). Plasma immunoglobulin type G levels to SARS-CoV-2 nucleocapsid core protein were similar between both arms. CONCLUSIONS: Famotidine was safe and well tolerated in outpatients with mild to moderate COVID-19. Famotidine led to earlier resolution of symptoms and inflammation without reducing anti-SARS-CoV-2 immunity. Additional randomised trials are required.

Foliar respiration is related to photosynthetic, growth and carbohydrate response to experimental drought and elevated temperature.

Short-term plant respiration (R) increases exponentially with rising temperature, but drought could reduce respiration by reducing growth and metabolism. Acclimation may alter these responses. We examined if species with different drought responses would differ in foliar R response to +4.8°C temperature and -45% precipitation in a field experiment with mature piñon and juniper trees, and if any differences between species were related to differences in photosynthesis rates, shoot growth and nonstructural carbohydrates (NSCs). Short-term foliar R had a Q10 of 1.6 for piñon and 2.6 for juniper. Piñon foliar R did not respond to the +4.8°C temperatures, but R increased 1.4× for juniper. Across treatments, piñon foliage had higher growth, lower NSC content, 29% lower photosynthesis rates, and 44% lower R than juniper. Removing 45% precipitation had little impact on R for either species. Species differences in the response of R under elevated temperature were related to substrate availability and stomatal response to leaf water potential. Despite not acclimating to the higher temperature and having higher R than piñon, greater substrate availability in juniper suggests it could supply respiratory demand for much longer than piñon. Species responses will be critical in ecosystem response to a warmer climate.

Physical structure and biological composition of canopies in tropical secondary and old-growth forests.

The area of tropical secondary forests is increasing rapidly, but data on the physical and biological structure of the canopies of these forests are limited. To obtain such data and to measure the ontogeny of canopy structure during tropical rainforest succession, we studied patch-scale (5 m2) canopy structure in three areas of 18-36 year-old secondary forest in Costa Rica, and compared the results to data from old-growth forest at the same site. All stands were sampled with a stratified random design with complete harvest from ground level to the top of the canopy from a modular portable tower. All canopies were organized into distinct high- and low-leaf-density layers (strata), and multiple strata developed quickly with increasing patch height. The relation of total Leaf Area Index (LAI, leaf area per area of ground) to patch canopy height, the existence of distinct high and low leaf- density layers (strata and free air spaces), the depth and LAI of the canopy strata and free air spaces, and the relation of the number of strata to patch canopy height were remarkably constant across the entire successional gradient. Trees were the most important contributor to LAI at all stages, while contribution of palm LAI increased through succession. We hypothesize that canopy physical structure at the patch scale is driven by light competition and discuss how this hypothesis could be tested. That canopy physical structure was relatively independent of the identity of the species present suggests that canopy physical structure may be conserved even as canopy floristics shift due to changing climate.

Plant respiration: Controlled by photosynthesis or biomass?

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 carry-over 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.

Tropical rainforest carbon sink declines during El Niño as a result of reduced photosynthesis and increased respiration rates.

Changes in tropical forest carbon sink strength during El Niño Southern Oscillation (ENSO) events can indicate future behavior under climate change. Previous studies revealed ˜6 Mg C ha-1  yr-1 lower net ecosystem production (NEP) during ENSO year 1998 compared with non-ENSO year 2000 in a Costa Rican tropical rainforest. We explored environmental drivers of this change and examined the contributions of ecosystem respiration (RE) and gross primary production (GPP) to this weakened carbon sink. For 1998-2000, we estimated RE using chamber-based respiration measurements, and we estimated GPP in two ways: using (1) the canopy process model MAESTRA, and (2) combined eddy covariance and chamber respiration data. MAESTRA-estimated GPP did not statistically differ from GPP estimated using approach 2, but was ˜ 28% greater than published GPP estimates for the same site and years using eddy covariance data only. A 7% increase in RE (primarily increased soil respiration) and a 10% reduction in GPP contributed equally to the difference in NEP between ENSO year 1998 and non-ENSO year 2000. A warming and drying climate for tropical forests may yield a weakened carbon sink from both decreased GPP and increased RE. Understanding physiological acclimation will be critical for the large carbon stores in these ecosystems.

Non-structural carbohydrates in woody plants compared among laboratories.

Non-structural carbohydrates (NSC) in plant tissue are frequently quantified to make inferences about plant responses to environmental conditions. Laboratories publishing estimates of NSC of woody plants use many different methods to evaluate NSC. We asked whether NSC estimates in the recent literature could be quantitatively compared among studies. We also asked whether any differences among laboratories were related to the extraction and quantification methods used to determine starch and sugar concentrations. These questions were addressed by sending sub-samples collected from five woody plant tissues, which varied in NSC content and chemical composition, to 29 laboratories. Each laboratory analyzed the samples with their laboratory-specific protocols, based on recent publications, to determine concentrations of soluble sugars, starch and their sum, total NSC. Laboratory estimates differed substantially for all samples. For example, estimates for Eucalyptus globulus leaves (EGL) varied from 23 to 116 (mean = 56) mg g(-1) for soluble sugars, 6-533 (mean = 94) mg g(-1) for starch and 53-649 (mean = 153) mg g(-1) for total NSC. Mixed model analysis of variance showed that much of the variability among laboratories was unrelated to the categories we used for extraction and quantification methods (method category R(2) = 0.05-0.12 for soluble sugars, 0.10-0.33 for starch and 0.01-0.09 for total NSC). For EGL, the difference between the highest and lowest least squares means for categories in the mixed model analysis was 33 mg g(-1) for total NSC, compared with the range of laboratory estimates of 596 mg g(-1). Laboratories were reasonably consistent in their ranks of estimates among tissues for starch (r = 0.41-0.91), but less so for total NSC (r = 0.45-0.84) and soluble sugars (r = 0.11-0.83). Our results show that NSC estimates for woody plant tissues cannot be compared among laboratories. The relative changes in NSC between treatments measured within a laboratory may be comparable within and between laboratories, especially for starch. To obtain comparable NSC estimates, we suggest that users can either adopt the reference method given in this publication, or report estimates for a portion of samples using the reference method, and report estimates for a standard reference material. Researchers interested in NSC estimates should work to identify and adopt standard methods.

Carbohydrate regulation of photosynthesis and respiration from branch girdling in four species of wet tropical rain forest trees.

How trees sense source-sink carbon balance remains unclear. One potential mechanism is a feedback from non-structural carbohydrates regulating photosynthesis and removing excess as waste respiration when the balance of photosynthesis against growth and metabolic activity changes. We tested this carbohydrate regulation of photosynthesis and respiration using branch girdling in four tree species in a wet tropical rainforest in Costa Rica. Because girdling severs phloem to stop carbohydrate export while leaving xylem intact to allow photosynthesis, we expected carbohydrates to accumulate in leaves to simulate a carbon imbalance. We varied girdling intensity by removing phloem in increments of one-quarter of the circumference (zero, one--quarter, half, three-quarters, full) and surrounded a target branch with fully girdled ones to create a gradient in leaf carbohydrate content. Light saturated photosynthesis rate was measured in situ, and foliar respiration rate and leaf carbohydrate content were measured after destructive harvest at the end of the treatment. Girdling intensity created no consistent or strong responses in leaf carbohydrates. Glucose and fructose slightly increased in all species by 3.4% per one-quarter girdle, total carbon content and leaf mass per area increased only in one species by 5.4 and 5.5% per one-quarter girdle, and starch did not change. Only full girdling lowered photosynthesis in three of four species by 59-69%, but the decrease in photosynthesis was unrelated to the increase in glucose and fructose content. Girdling did not affect respiration. The results suggest that leaf carbohydrate content remains relatively constant under carbon imbalance, and any changes are unlikely to regulate photosynthesis or respiration. Because girdling also stops the export of hormones and reactive oxygen species, girdling may induce physiological changes unrelated to carbohydrate accumulation and may not be an effective method to study carbohydrate feedback in leaves. In three species, removal of three-quarters of phloem area did not cause leaf carbohydrates to accumulate nor did it change photosynthesis or respiration, suggesting that phloem transport is flexible and transport rate per unit phloem can rapidly increase under an increase in carbohydrate supply relative to phloem area. Leaf carbohydrate content thus may be decoupled from whole plant carbon balance by phloem transport in some species, and carbohydrate regulation of photosynthesis and respiration may not be as common in trees as previous girdling studies suggest. Further studies in carbohydrate regulation should avoid using girdling as girdling can decrease photosynthesis through unintended means without the tested mechanisms of accumulating leaf carbohydrates.

Global variability in leaf respiration in relation to climate, plant functional types and leaf traits.

Leaf dark respiration (Rdark ) is an important yet poorly quantified component of the global carbon cycle. Given this, we analyzed a new global database of Rdark and associated leaf traits. Data for 899 species were compiled from 100 sites (from the Arctic to the tropics). Several woody and nonwoody plant functional types (PFTs) were represented. Mixed-effects models were used to disentangle sources of variation in Rdark . Area-based Rdark at the prevailing average daily growth temperature (T) of each site increased only twofold from the Arctic to the tropics, despite a 20°C increase in growing T (8-28°C). By contrast, Rdark at a standard T (25°C, Rdark (25) ) was threefold higher in the Arctic than in the tropics, and twofold higher at arid than at mesic sites. Species and PFTs at cold sites exhibited higher Rdark (25) at a given photosynthetic capacity (Vcmax (25) ) or leaf nitrogen concentration ([N]) than species at warmer sites. Rdark (25) values at any given Vcmax (25) or [N] were higher in herbs than in woody plants. The results highlight variation in Rdark among species and across global gradients in T and aridity. In addition to their ecological significance, the results provide a framework for improving representation of Rdark in terrestrial biosphere models (TBMs) and associated land-surface components of Earth system models (ESMs).

Variation in foliar respiration and wood CO2 efflux rates among species and canopy layers in a wet tropical forest.

As tropical forests respond to environmental change, autotrophic respiration may consume a greater proportion of carbon fixed in photosynthesis at the expense of growth, potentially turning the forests into a carbon source. Predicting such a response requires that we measure and place autotrophic respiration in a complete carbon budget, but extrapolating measurements of autotrophic respiration from chambers to ecosystem remains a challenge. High plant species diversity and complex canopy structure may cause respiration rates to vary and measurements that do not account for this complexity may introduce bias in extrapolation more detrimental than uncertainty. Using experimental plantations of four native tree species with two canopy layers, we examined whether species and canopy layers vary in foliar respiration and wood CO2 efflux and whether the variation relates to commonly used scalars of mass, nitrogen (N), photosynthetic capacity and wood size. Foliar respiration rate varied threefold between canopy layers, ∼0.74 µmol m(-2) s(-1) in the overstory and ∼0.25 µmol m(-2) s(-1) in the understory, but little among species. Leaf mass per area, N and photosynthetic capacity explained some of the variation, but height explained more. Chamber measurements of foliar respiration thus can be extrapolated to the canopy with rates and leaf area specific to each canopy layer or height class. If area-based rates are sampled across canopy layers, the area-based rate may be regressed against leaf mass per area to derive the slope (per mass rate) to extrapolate to the canopy using the total leaf mass. Wood CO2 efflux varied 1.0-1.6 µmol m(-2) s(-1) for overstory trees and 0.6-0.9 µmol m(-2) s(-1) for understory species. The variation in wood CO2 efflux rate was mostly related to wood size, and little to species, canopy layer or height. Mean wood CO2 efflux rate per surface area, derived by regressing CO2 efflux per mass against the ratio of surface area to mass, can be extrapolated to the stand using total wood surface area. The temperature response of foliar respiration was similar for three of the four species, and wood CO2 efflux was similar between wet and dry seasons. For these species and this forest, vertical sampling may yield more accurate estimates than would temporal sampling.

Forest ecosystem respiration estimated from eddy covariance and chamber measurements under high turbulence and substantial tree mortality from bark beetles.

Eddy covariance nighttime fluxes are uncertain due to potential measurement biases. Many studies report eddy covariance nighttime flux lower than flux from extrapolated chamber measurements, despite corrections for low turbulence. We compared eddy covariance and chamber estimates of ecosystem respiration at the GLEES Ameriflux site over seven growing seasons under high turbulence [summer night mean friction velocity (u*) = 0.7 m s(-1)], during which bark beetles killed or infested 85% of the aboveground respiring biomass. Chamber-based estimates of ecosystem respiration during the growth season, developed from foliage, wood, and soil CO2 efflux measurements, declined 35% after 85% of the forest basal area had been killed or impaired by bark beetles (from 7.1 ± 0.22 µmol m(-2) s(-1) in 2005 to 4.6 ± 0.16 µmol m(-2) s(-1) in 2011). Soil efflux remained at ~3.3 µmol m(-2) s(-1) throughout the mortality, while the loss of live wood and foliage and their respiration drove the decline of the chamber estimate. Eddy covariance estimates of fluxes at night remained constant over the same period, ~3.0 µmol m(-2) s(-1) for both 2005 (intact forest) and 2011 (85% basal area killed or impaired). Eddy covariance fluxes were lower than chamber estimates of ecosystem respiration (60% lower in 2005, and 32% in 2011), but the mean night estimates from the two techniques were correlated within a year (r(2) from 0.18 to 0.60). The difference between the two techniques was not the result of inadequate turbulence, because the results were robust to a u* filter of >0.7 m s(-1). The decline in the average seasonal difference between the two techniques was strongly correlated with overstory leaf area (r(2) = 0.92). The discrepancy between methods of respiration estimation should be resolved to have confidence in ecosystem carbon flux estimates.

El Niño Southern Oscillation (ENSO) enhances CO2 exchange rates in freshwater Marsh ecosystems in the Florida everglades.

This research examines the relationships between El Niño Southern Oscillation (ENSO), water level, precipitation patterns and carbon dioxide (CO2) exchange rates in the freshwater wetland ecosystems of the Florida Everglades. Data was obtained over a 5-year study period (2009-2013) from two freshwater marsh sites located in Everglades National Park that differ in hydrology. At the short-hydroperiod site (Taylor Slough; TS) and the long-hydroperiod site (Shark River Slough; SRS) fluctuations in precipitation patterns occurred with changes in ENSO phase, suggesting that extreme ENSO phases alter Everglades hydrology which is known to have a substantial influence on ecosystem carbon dynamics. Variations in both ENSO phase and annual net CO2 exchange rates co-occurred with changes in wet and dry season length and intensity. Combined with site-specific seasonality in CO2 exchanges rates, El Niño and La Niña phases magnified season intensity and CO2 exchange rates at both sites. At TS, net CO2 uptake rates were higher in the dry season, whereas SRS had greater rates of carbon sequestration during the wet season. As La Niña phases were concurrent with drought years and extended dry seasons, TS became a greater sink for CO2 on an annual basis (-11 to -110 g CO2 m-2 yr-1) compared to El Niño and neutral years (-5 to -43.5 g CO2 m-2 yr-1). SRS was a small source for CO2 annually (1.81 to 80 g CO2 m-2 yr-1) except in one exceptionally wet year that was associated with an El Niño phase (-16 g CO2 m-2 yr-1). Considering that future climate predictions suggest a higher frequency and intensity in El Niño and La Niña phases, these results indicate that changes in extreme ENSO phases will significantly alter CO2 dynamics in the Florida Everglades.

Short and long-term carbon balance of bioenergy electricity production fueled by forest treatments.

BACKGROUND: Forests store large amounts of carbon in forest biomass, and this carbon can be released to the atmosphere following forest disturbance or management. In the western US, forest fuel reduction treatments designed to reduce the risk of high severity wildfire can change forest carbon balance by removing carbon in the form of biomass, and by altering future potential wildfire behavior in the treated stand. Forest treatment carbon balance is further affected by the fate of this biomass removed from the forest, and the occurrence and intensity of a future wildfire in this stand. In this study we investigate the carbon balance of a forest treatment with varying fates of harvested biomass, including use for bioenergy electricity production, and under varying scenarios of future disturbance and regeneration. RESULTS: Bioenergy is a carbon intensive energy source; in our study we find that carbon emissions from bioenergy electricity production are nearly twice that of coal for the same amount of electricity. However, some emissions from bioenergy electricity production are offset by avoided fossil fuel electricity emissions. The carbon benefit achieved by using harvested biomass for bioenergy electricity production may be increased through avoided pyrogenic emissions if the forest treatment can effectively reduce severity. CONCLUSION: Forest treatments with the use of harvested biomass for electricity generation can reduce carbon emissions to the atmosphere by offsetting fossil fuel electricity generation emissions, and potentially by avoided pyrogenic emissions due to reduced intensity and severity of a future wildfire in the treated stand. However, changes in future wildfire and regeneration regimes may affect forest carbon balance and these climate-induced changes may influence forest carbon balance as much, or more, than bioenergy production.

Three decades of research at Flakaliden advancing whole-tree physiology, forest ecosystem and global change research.

Nutrient supply often limits growth in forest ecosystems and may limit the response of growth to an increase in other resources, or to more favorable environmental factors such as temperature and soil water. To explore the consequences and mechanisms of optimum nutrient supply for forest growth, the Flakaliden research site was established in 1986 on a young Norway spruce site with nutrient-poor soil. This special section on research at Flakaliden presents five papers that explore different facets of nutrition, atmospheric CO2 concentration, [CO2], and increased temperature treatments, using the original experiment as a base. Research at Flakaliden shows the dominant role of nutrition in controlling the response of growth to the increased photosynthesis promoted by elevated [CO2] and temperature. Experiments with whole-tree chambers showed that all treatments (air temperature warming, elevated [CO2] and optimum nutrition) increased shoot photosynthesis by 30-50%, but growth only increased with [CO2] when combined with the optimum nutrition treatment. Elevated [CO2] and temperature increased shoot photosynthesis by increasing the slope between light-saturated photosynthesis and foliar nitrogen by 122%, the initial slope of the light response curve by 52% and apparent quantum yield by 10%. Optimum nutrition also decreased photosynthetic capacity by 17%, but increased it by 62% in elevated [CO2], as estimated from wood δ(13)C. Elevated air temperature advanced spring recovery of photosynthesis by 37%, but spring frost events remained the controlling factor for photosynthetic recovery, and elevated [CO2] did not affect this. Increased nutrient availability increased wood growth primarily through a 50% increase in tracheid formation, mostly during the peak growth season. Other notable contributions of research at Flakaliden include exploring the role of optimal nutrition in large-scale field trials with foliar analysis, using an ecosystem approach for multifactor experiments, development of whole-tree chambers allowing inexpensive environmental manipulations, long-term deployment of shoot chambers for continuous measurements of gas exchange and exploring the ecosystem response to soil and aboveground tree warming. The enduring legacy of Flakaliden will be the rich data set of long-term, multifactor experiments that has been and will continue to be used in many modeling and cross-site comparison studies.