Thermal acclimation of leaf respiration of tropical trees and lianas: response to experimental canopy warming,and consequences for tropical forest carbon balance

Climate warming is expected to increase respiration rates of tropical forest trees and lianas, which may negatively affect the carbon balance of tropical forests. Thermal acclimation could mitigate the expected respiration increase, but the thermal acclimation potential of tropical forests remains largely unknown. In a tropical forest in Panama, we experimentally increased nighttime temperatures of upper canopy leaves of three tree and two liana species by on average 3  ° C for 1 week, and quantified temperature responses of leaf dark respiration. Respiration at 25  ° C (R₂₅) decreased with increasing leaf temperature, but acclimation did not result in perfect homeostasis of respiration across temperatures. In contrast, Q₁₀ of treatment and control leaves exhibited similarly high values (range 2.5–3.0) without evidence of acclimation. The decrease in R₂₅ was not caused by respiratory substrate depletion, as warming did not reduce leaf carbohydrate concentration. To evaluate the wider implications of our experimental results, we simulated the carbon cycle of tropical latitudes (24 ° S–24 ° N) from 2000 to 2100 using a dynamic global vegetation model (LM3VN) modified to account for acclimation. Acclimation reduced the degree to which respiration increases with climate warming in the model relative to a no‐acclimation scenario, leading to 21% greater increase in net primary productivity and 18% greater increase in biomass carbon storage over the 21st century. We conclude that leaf respiration of tropical forest plants can acclimate to nighttime warming, thereby reducing the magnitude of the positive feedback between climate change and the carbon cycle.     

A common thermal niche among geographically diverse populations of the widely distributed tree species Eucalyptus tereticornis: No evidence for adaptation to climate‐of‐origin

Impacts of climate warming depend on the degree to which plants are constrained by adaptation to their climate‐of‐origin or exhibit broad climatic suitability. We grew cool‐origin, central and warm‐origin provenances of Eucalyptus tereticornis in an array of common temperature environments from 18 to 35.5°C to determine if this widely distributed tree species consists of geographically contrasting provenances with differentiated and narrow thermal niches, or if provenances share a common thermal niche. The temperature responses of photosynthesis, respiration, and growth were equivalent across the three provenances, reflecting a common thermal niche despite a 2,200 km geographic distance and 13°C difference in mean annual temperature at seed origin. The temperature dependence of growth was primarily mediated by changes in leaf area per unit plant mass, photosynthesis, and whole‐plant respiration. Thermal acclimation of leaf, stem, and root respiration moderated the increase in respiration with temperature, but acclimation was constrained at high temperatures. We conclude that this species consists of provenances that are not differentiated in their thermal responses, thus rejecting our hypothesis of adaptation to climate‐of‐origin and suggesting a shared thermal niche. In addition, growth declines with warming above the temperature optima were driven by reductions in whole‐plant leaf area and increased respiratory carbon losses. The impacts of climate warming will nonetheless vary across the geographic range of this and other such species, depending primarily on each provenance's climate position on the temperature response curves for photosynthesis, respiration, and growth.     

Coupling of respiration, nitrogen, and sugars underlies convergent temperature acclimation in Pinus banksiana across wide-ranging sites and populations

Patterns and mechanisms of short‐term temperature acclimation and long‐term climatic adaptation of respiration among intraspecific populations are poorly understood, but both are potentially important in constraining respiratory carbon flux to climate warming across large geographic scales, as well as influencing the metabolic fitness of populations. Herein we report on leaf dark respiration of 33‐year‐old trees of jack pine (Pinus banksiana Lamb.) grown in three contrasting North American common gardens (0.9, 4.6, and 7.9 °C, mean annual temperature) comprised of identical populations of wide‐ranging geographic origins. We tested whether respiration rates in this evergreen conifer acclimate to prevailing ambient air temperatures and differ among populations. At each of the common gardens, observed population differences in respiration rates measured at a standard temperature (20 °C) were comparatively small and largely unrelated to climate of seed‐source origin. In contrast, respiration in all populations exhibited seasonal acclimation at all sites. Specific respiration rates at 20 °C inversely tracked seasonal variation in ambient air temperature, increasing with cooler temperatures in fall and declining with warmer temperatures in spring and summer. Such responses were similar among populations and sites, thus providing a general predictive equation regarding temperature acclimation of respiration for the species. Temperature acclimation was associated with variation in nitrogen (N) and soluble carbohydrate concentrations, supporting a joint enzyme and substrate‐based model of respiratory acclimation. Regression analyses revealed convergent relationships between respiration and the combination of needle N and soluble carbohydrate concentrations and between N‐based respiration (R_N, μmol mol N^{? 1} s^{? 1}) and soluble carbohydrate concentrations, providing evidence for general predictive relationships across geographically diverse populations, seasons, and sites. Overall, these findings demonstrate that seasonal acclimation of respiration modulates rates of foliar respiratory carbon flux in a widely distributed evergreen species, and does so in a predictable way. Genetic differences in specific respiration rate appear less important than temperature acclimation in downregulating respiratory carbon fluxes with climate warming across wide‐ranging sites.     

The relative impacts of daytime and night-time warming on photosynthetic capacity in Populus deltoides

In order to investigate the relative impacts of increases in day and night temperature on tree carbon relations, we measured night‐time respiration and daytime photosynthesis of leaves in canopies of 4‐m‐tall cottonwood (Populus deltoides Bartr. ex Marsh) trees experiencing three daytime temperatures (25, 28 or 31 °C) and either (i) a constant nocturnal temperature of 20 °C or (ii) increasing nocturnal temperatures (15, 20 or 25 °C). In the first (day warming only) experiment, rates of night‐time leaf dark respiration (R_dark) remained constant and leaves displayed a modest increase (11%) in light‐saturated photosynthetic capacity (A_max) during the day (1000–1300 h) over the 6 °C range. In the second (dual night and day warming) experiment, R_dark increased by 77% when nocturnal temperatures were increased from 15 °C (0·36 µmol m^?2 s^?1) to 25 °C (0·64 µmol m^?2 s^?1). A_max responded positively to the additional nocturnal warming, and increased by 38 and 64% in the 20/28 and 25/31 °C treatments, respectively, compared with the 15/25 °C treatment. These increases in photosynthetic capacity were associated with strong increases in the maximum carboxylation rate of rubisco (V_cmax) and ribulose‐1,5‐bisphosphate (RuBP) regeneration capacity mediated by maximum electron transport rate (J_max). Leaf soluble sugar and starch concentration, measured at sunrise, declined significantly as nocturnal temperature increased. The nocturnal temperature manipulation resulted in a significant inverse relationship between A_max and pre‐dawn leaf carbohydrate status. Independent measurements of the temperature response of photosynthesis indicated that the optimum temperature (T_opt) acclimated fully to the 6 °C range of temperature imposed in the daytime warming. Our findings are consistent with the hypothesis that elevated night‐time temperature increases photosynthetic capacity during the following light period through a respiratory‐driven reduction in leaf carbohydrate concentration. These responses indicate that predicted increases in night‐time minimum temperatures may have a significant influence on net plant carbon uptake.     

Contrasting thermal acclimation of leaf dark respiration and photosynthesis of Antarctic vascular plant species exposed to nocturnal warming

Leaf respiration and photosynthesis will respond differently to an increase in temperature during night, which can be more relevant in sensitive ecosystems such as Antarctica. We postulate that the plant species able to colonize the Antarctic Peninsula – Colobanthus quitensis (Kunth) Bartl. and Deschampsia antarctica Desv. – are able to acclimate their foliar respiration and to maintain photosynthesis under nocturnal warming to sustain a positive foliar carbon balance. We conducted a laboratory experiment to evaluate the effect of time of day (day and night) and nocturnal warming on dark respiration. Short (E₀ and Q₁₀) and long‐term acclimation of respiration, leaf carbohydrates, photosynthesis (A_sat) and foliar carbon balance (R/A) were evaluated. The results suggest that the two species have differential thermal acclimation respiration, where D. antarctica showed more thermosensitivity to short‐term changes in temperature than C. quitensis. Experimental nocturnal warming affected respiration at daytime differentially between the two species, with a significant increase of R₁₀ and A_sat in D. antarctica, while no changes on respiration were observed in C. quitensis. Long thermal treatments of the plants indicated that nocturnal but not diurnal respiration could acclimate in both species, and to a greater extent in C. quitensis. Non‐structural carbohydrates were related with respiration in C. quitensis but not in D. antarctica, suggesting that respiration in the former species is likely controlled by total soluble sugars and starch during day and night, respectively. Finally, foliar carbon balance was differentially improved under warming conditions in Antarctic plants by different mechanisms, with C. quitensis deploying respiratory acclimation, while D. antarctica increased its A_sat.     

Thermal acclimation of leaf and root respiration: an investigation comparing inherently fast- and slow-growing plant species

We investigated the extent to which leaf and root respiration (R) differ in their response to short‐ and long‐term changes in temperature in several contrasting plant species (herbs, grasses, shrubs and trees) that differ in inherent relative growth rate (RGR, increase in mass per unit starting mass and time). Two experiments were conducted using hydroponically grown plants. In the long‐term (LT) acclimation experiment, 16 species were grown at constant 18, 23 and 28 °C. In the short‐term (ST) acclimation experiment, 9 of those species were grown at 25/20 °C (day/night) and then shifted to a 15/10 °C for 7 days. Short‐term Q₁₀ values (proportional change in R per 10 °C) and the degree of acclimation to longer‐term changes in temperature were compared. The effect of growth temperature on root and leaf soluble sugar and nitrogen concentrations was examined. Light‐saturated photosynthesis (A_sat) was also measured in the LT acclimation experiment. Our results show that Q₁₀ values and the degree of acclimation are highly variable amongst species and that roots exhibit lower Q₁₀ values than leaves over the 15–25 °C measurement temperature range. Differences in RGR or concentrations of soluble sugars/nitrogen could not account for the inter‐specific differences in the Q₁₀ or degree of acclimation. There were no systematic differences in the ability of roots and leaves to acclimate when plants developed under contrasting temperatures (LT acclimation). However, acclimation was greater in both leaves and roots that developed at the growth temperature (LT acclimation) than in pre‐existing leaves and roots shifted from one temperature to another (ST acclimation). The balance between leaf R and A_sat was maintained in plants grown at different temperatures, regardless of their inherent relative growth rate. We conclude that there is tight coupling between the respiratory acclimation and the temperature under which leaves and roots developed and that acclimation plays an important role in determining the relationship between respiration and photosynthesis.     

High thermal acclimation potential of both photosynthesis and respiration in two lowland Plantago species in contrast to an alpine congeneric

Thermal acclimation of photosynthesis and respiration can enable plants to maintain near constant rates of net CO₂ exchange, despite experiencing sustained changes in daily average temperature. In this study, we investigated whether the degree of acclimation of photosynthesis and respiration of mature leaves differs among three congeneric Plantago species from contrasting habitats [two fast‐growing lowland species (Plantago major and P. lanceolata), and one slow‐growing alpine species (P. euryphylla)]. In addition to investigating some mechanisms underpinning variability in photosynthetic acclimation, we also determined whether leaf respiration in the light acclimates to the same extent as leaf respiration in darkness, and whether acclimation reestablishes the balance between leaf respiration and photosynthesis. Three growth temperatures were provided: constant 13, 20, or 27°C. Measurements were made at five temperatures (6–34°C). Little acclimation of photosynthesis and leaf respiration to growth temperature was exhibited by P. euryphylla. Moreover, leaf masses per area (LMA) were similar in 13°C‐grown and 20°C‐grown plants of the alpine species. In contrast, growth at 13°C increased LMA in the two lowland species; this was associated with increased photosynthetic capacity and rates of leaf respiration (both in darkness and in the light). Alleviation of triose phosphate limitation and increased capacity of electron transport capacity relative to carboxylation were also observed. Such changes demonstrate that the lowland species cold‐acclimated. Light reduced the short‐term temperature dependence (i.e. Q₁₀) of leaf respiration in all three species, irrespective of growth temperature. Collectively, our results highlight the tight coupling that exists between thermal acclimation of photosynthetic and leaf respiratory metabolism (both in darkness and in the light) in Plantago. If widespread among contrasting species, such coupling may enable modellers to assume levels of acclimation in one parameter (e.g. leaf respiration) where details are only known for the other (e.g. photosynthesis).     

Using temperature‐dependent changes in leaf scaling relationships to quantitatively account for thermal acclimation of respiration in a coupled global climate–vegetation model

The response of plant respiration (R) to temperature is an important component of the biosphere's response to climate change. At present, most global models assume that R increases exponentially with temperature and does not thermally acclimate. Although we now know that acclimation does occur, quantitative incorporation of acclimation into models has been lacking. Using a dataset for 19 species grown at four temperatures (7, 14, 21, and 28 °C), we have assessed whether sustained differences in growth temperature systematically alter the slope and/or intercepts of the generalized log–log plots of leaf R vs. leaf mass per unit leaf area (LMA) and vs. leaf nitrogen (N) concentration. The extent to which variations in growth temperature account for the scatter observed in log–log R–LMA–N scaling relationships was also assessed. We show that thermal history accounts for up to 20% of the scatter in scaling relationships used to predict R, with the impact of thermal history on R–LMA–N generalized scaling relationships being highly predictable. This finding enabled us to quantitatively incorporate acclimation of R into a coupled global climate–vegetation model. We show that accounting for acclimation of R has negligible impact on predicted annual rates of global R, net primary productivity (NPP) or future atmospheric CO₂ concentrations. However, our analysis suggests that accounting for acclimation is important when considering carbon fluxes among thermally contrasting biomes (e.g. accounting for acclimation decreases predicted rates of R by up to 20% in high‐temperature biomes). We conclude that acclimation of R needs to be accounted for when predicting potential responses of terrestrial carbon exchange to climatic change at a regional level.     

Predicting ecosystem carbon balance in a warming Arctic: the importance of long‐term thermal acclimation potential and inhibitory effects of light on respiration

The carbon balance of Arctic ecosystems is particularly sensitive to global environmental change. Leaf respiration (R), a temperature‐dependent key process in determining the carbon balance, is not well‐understood in Arctic plants. The potential for plants to acclimate to warmer conditions could strongly impact future global carbon balance. Two key unanswered questions are (1) whether short‐term temperature responses can predict long‐term respiratory responses to growth in elevated temperatures and (2) to what extent the constant daylight conditions of the Arctic growing season inhibit leaf respiration. In two dominant Arctic species E riophorum vaginatum (tussock grass) and B etula nana (woody shrub), we assessed the extent of respiratory inhibition in the light (R _L/R _D), respiratory response to short‐term temperature change, and respiratory acclimation to long‐term warming treatments. We found that R of both species is strongly inhibited by light (averaging 35% across all measurement temperatures). In E . vaginatum both R _L and R _D acclimated to the long‐term warming treatment, reducing the magnitude of respiratory response relative to the short‐term response to temperature increase. In B . nana, both R _L and R _D responded to short‐term temperature increase but showed no acclimation to the long‐term warming. The ability to predict plant respiratory response to global warming with short‐term temperature responses will depend on species‐specific acclimation potential and the differential response of R _L and R _D to temperature. With projected woody shrub encroachment in Arctic tundra and continued warming, changing species dominance between these two functional groups, may impact ecosystem respiratory response and carbon balance.     

Seasonal acclimation of leaf respiration in Eucalyptus saligna trees: impacts of elevated atmospheric CO2 and summer drought

Understanding the impacts of atmospheric [CO₂] and drought on leaf respiration (R) and its response to changes in temperature is critical to improve predictions of plant carbon‐exchange with the atmosphere, especially at higher temperatures. We quantified the effects of [CO₂]‐enrichment (+240 ppm) on seasonal shifts in the diel temperature response of R during a moderate summer drought in Eucalyptus saligna growing in whole‐tree chambers in SE Australia. Seasonal temperature acclimation of R was marked, as illustrated by: (1) a downward shift in daily temperature response curves of R in summer (relative to spring); (2)≈60% lower R measured at 20^oC (R₂₀) in summer compared with spring; and (3) homeostasis over 12 months of R measured at prevailing nighttime temperatures. R₂₀, measured during the day, was on average 30–40% higher under elevated [CO₂] compared with ambient [CO₂] across both watered and droughted trees. Drought reduced R₂₀ by≈30% in both [CO₂] treatments resulting in additive treatment effects. Although [CO₂] had no effect on seasonal acclimation, summer drought exacerbated the seasonal downward shift in temperature response curves of R. Overall, these results highlight the importance of seasonal acclimation of leaf R in trees grown under ambient‐ and elevated [CO₂] as well as under moderate drought. Hence, respiration rates may be overestimated if seasonal changes in temperature and drought are not considered when predicting future rates of forest net CO₂ exchange.     

Thermal Acclimation of Respiration and Photosynthesis in the Marine Macroalga Gracilaria lemaneiformis (Gracilariales,Rhodophyta)

The responses of respiration and photosynthesis to temperature fluctuations in marine macroalgae have the potential to significantly affect coastal carbon fluxes and sequestration. In this study, the marine red macroalga Gracilaria lemaneiformis was cultured at three different temperatures (12, 19, and 26°C) and at high‐ and low‐nitrogen (N) availability, to investigate the acclimation potential of respiration and photosynthesis to temperature change. Measurements of respiratory and photosynthetic rates were made at five temperatures (7°C–33°C). An instantaneous change in temperature resulted in a change in the rates of respiration and photosynthesis, and the temperature sensitivities (i.e., the Q₁₀ value) for both the metabolic processes were lower in 26°C‐grown algae than 12°C‐ or 19°C‐grown algae. Both respiration and photosynthesis acclimated to long‐term changes in temperature, irrespective of the N availability under which the algae were grown; respiration displayed strong acclimation, whereas photosynthesis only exhibited a partial acclimation response to changing growth temperatures. The ratio of respiration to gross photosynthesis was higher in 12°C‐grown algae, but displayed little difference between the algae grown at 19°C and 26°C. We propose that it is unlikely that respiration in G. lemaneiformis would increase significantly with global warming, although photosynthesis would increase at moderately elevated temperatures.     

Carbon fluxes acclimate more strongly to elevated growth temperatures than to elevated CO2 concentrations in a northern conifer

Increasing temperatures and atmospheric CO₂ concentrations will affect tree carbon fluxes, generating potential feedbacks between forests and the global climate system. We studied how elevated temperatures and CO₂ impacted leaf carbon dynamics in Norway spruce (Picea abies), a dominant northern forest species, to improve predictions of future photosynthetic and respiratory fluxes from high‐latitude conifers. Seedlings were grown under ambient (AC, c. 435 μmol mol^?1) or elevated (EC, 750 μmol mol^?1) CO₂ concentrations at ambient, +4 °C, or +8 °C growing temperatures. Photosynthetic rates (A_sat) were high in +4 °C/EC seedlings and lowest in +8 °C spruce, implying that moderate, but not extreme, climate change may stimulate carbon uptake. A_sat, dark respiration (R_dark), and light respiration (R_light) rates acclimated to temperature, but not CO₂: the thermal optimum of A_sat increased, and R_dark and R_light were suppressed under warming. In all treatments, the Q₁₀ of R_light (the relative increase in respiration for a 10 °C increase in leaf temperature) was 35% higher than the Q₁₀ of R_dark, so the ratio of R_light to R_dark increased with rising leaf temperature. However, across all treatments and a range of 10–40 °C leaf temperatures, a consistent relationship between R_light and R_dark was found, which could be used to model R_light in future climates. Acclimation reduced daily modeled respiratory losses from warm‐grown seedlings by 22–56%. When R_light was modeled as a constant fraction of R_dark, modeled daily respiratory losses were 11–65% greater than when using measured values of R_light. Our findings highlight the impact of acclimation to future climates on predictions of carbon uptake and losses in northern trees, in particular the need to model daytime respiratory losses from direct measurements of R_light or appropriate relationships with R_dark.     

Acclimation of snow gum (Eucalyptus pauciflora) leaf respiration to seasonal and diurnal variations in temperature: the importance of changes in the capacity and temperature sensitivity of respiration

We investigated the relationship between daily and seasonal temperature variation and dark respiratory CO₂ release by leaves of snow gum (Eucalyptus pauciflora Sieb. ex Spreng) that were grown in their natural habitat or under controlled‐environment conditions. The open grassland field site in SE Australia was characterized by large seasonal and diurnal changes in air temperature. On each measurement day, leaf respiration rates in darkness were measured in situ at 2–3 h intervals over a 24 h period, with measurements being conducted at the ambient leaf temperature. The rate of respiration at a set measuring temperature (i.e. apparent ‘respiratory capacity’) was greater in seedlings grown under low average daily temperatures (i.e. acclimation occurred), both in the field and under controlled‐environment conditions. The sensitivity of leaf respiration to diurnal changes in temperature (i.e. the Q₁₀ of leaf respiration) exhibited little seasonal variation over much of the year. However, Q₁₀ values were significantly greater on cold winter days (i.e. when daily average and minimum air temperatures were below 6° and –1 °C, respectively). These differences in Q₁₀ values were not due to bias arizing from the contrasting daily temperature amplitudes in winter and summer, as the Q₁₀ of leaf respiration was constant over a wide temperature range in short‐term experiments. Due to the higher Q₁₀ values in winter, there was less difference between winter and summer leaf respiration rates measured at 5 °C than at 25 °C. The net result of these changes was that there was relatively little difference in total daily leaf respiratory CO₂ release per unit leaf dry mass in winter and summer. Under controlled‐environment conditions, acclimation of respiration to growth temperature occurred in as little as 1–3 d. Acclimation was associated with a change in the concentration of soluble sugars under controlled conditions, but not in the field. Our data suggest that acclimation in the field may be associated with the onset of cold‐induced photo‐inhibition. We conclude that cold‐acclimation of dark respiration in snow gum leaves is characterized by changes in both the temperature sensitivity and apparent ‘capacity’ of the respiratory apparatus, and that such changes will have an important impact on the carbon economy of snow gum plants.     

Thermal plasticity of photosynthesis: the role of acclimation in forest responses to a warming climate

The increasing air temperatures central to climate change predictions have the potential to alter forest ecosystem function and structure by exceeding temperatures optimal for carbon gain. Such changes are projected to threaten survival of sensitive species, leading to local extinctions, range migrations, and altered forest composition. This study investigated photosynthetic sensitivity to temperature and the potential for acclimation in relation to the climatic provenance of five species of deciduous trees, Liquidambar styraciflua, Quercus rubra, Quercus falcata, Betula alleghaniensis, and Populus grandidentata. Open‐top chambers supplied three levels of warming (+0, +2, and +4 °C above ambient) over 3 years, tracking natural temperature variability. Optimal temperature for CO₂ assimilation was strongly correlated with daytime temperature in all treatments, but assimilation rates at those optima were comparable. Adjustment of thermal optima was confirmed in all species, whether temperatures varied with season or treatment, and regardless of climate in the species' range or provenance of the plant material. Temperature optima from 17° to 34° were observed. Across species, acclimation potentials varied from 0.55 °C to 1.07 °C per degree change in daytime temperature. Responses to the temperature manipulation were not different from the seasonal acclimation observed in mature indigenous trees, suggesting that photosynthetic responses should not be modeled using static temperature functions, but should incorporate an adjustment to account for acclimation. The high degree of homeostasis observed indicates that direct impacts of climatic warming on forest productivity, species survival, and range limits may be less than predicted by existing models.     

Autumn photosynthetic decline and growth cessation in seedlings of white spruce are decoupled under warming and photoperiod manipulations

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high‐latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down‐regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change.     

Pathways regulating decreased soil respiration with warming in a biocrust‐dominated dryland

A positive soil carbon (C)‐climate feedback is embedded into the climatic models of the IPCC. However, recent global syntheses indicate that the temperature sensitivity of soil respiration (R_S) in drylands, the largest biome on Earth, is actually lower in warmed than in control plots. Consequently, soil C losses with future warming are expected to be low compared with other biomes. Nevertheless, the empirical basis for these global extrapolations is still poor in drylands, due to the low number of field experiments testing the pathways behind the long‐term responses of soil respiration (R_S) to warming. Importantly, global drylands are covered with biocrusts (communities formed by bryophytes, lichens, cyanobacteria, fungi, and bacteria), and thus, R_S responses to warming may be driven by both autotrophic and heterotrophic pathways. Here, we evaluated the effects of 8‐year experimental warming on R_S, and the different pathways involved, in a biocrust‐dominated dryland in southern Spain. We also assessed the overall impacts on soil organic C (SOC) accumulation over time. Across the years and biocrust cover levels, warming reduced R_S by 0.30 μmol CO₂ m^?2 s^?1 (95% CI = ?0.24 to 0.84), although the negative warming effects were only significant after 3 years of elevated temperatures in areas with low initial biocrust cover. We found support for different pathways regulating the warming‐induced reduction in R_S at areas with low (microbial thermal acclimation via reduced soil mass‐specific respiration and β‐glucosidase enzymatic activity) vs. high (microbial thermal acclimation jointly with a reduction in autotrophic respiration from decreased lichen cover) initial biocrust cover. Our 8‐year experimental study shows a reduction in soil respiration with warming and highlights that biocrusts should be explicitly included in modeling efforts aimed to quantify the soil C–climate feedback in drylands.     

After more than a decade of soil moisture deficit,tropical rainforest trees maintain photosynthetic capacity,despite increased leaf respiration

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through‐fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought‐stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought‐induced mortality. We hypothesize that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short‐lived periods of high moisture availability, when stomatal conductance (g_s) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (R_d) was elevated in the TFE‐treated forest trees relative to the control by 28.2 ± 2.8% (mean ± one standard error). This mean R_d value was dominated by a 48.5 ± 3.6% increase in the R_d of drought‐sensitive taxa, and likely reflects the need for additional metabolic support required for stress‐related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity.     

Short‐term acclimation to warmer temperatures accelerates leaf carbon exchange processes across plant types

While temperature responses of photosynthesis and plant respiration are known to acclimate over time in many species, few studies have been designed to directly compare process‐level differences in acclimation capacity among plant types. We assessed short‐term (7 day) temperature acclimation of the maximum rate of Rubisco carboxylation (V_cmax), the maximum rate of electron transport (J_max), the maximum rate of phosphoenolpyruvate carboxylase carboxylation (V_pmax), and foliar dark respiration (R_d) in 22 plant species that varied in lifespan (annual and perennial), photosynthetic pathway (C₃ and C₄), and climate of origin (tropical and nontropical) grown under fertilized, well‐watered conditions. In general, acclimation to warmer temperatures increased the rate of each process. The relative increase in different photosynthetic processes varied by plant type, with C₃ species tending to preferentially accelerate CO₂‐limited photosynthetic processes and respiration and C₄ species tending to preferentially accelerate light‐limited photosynthetic processes under warmer conditions. R_d acclimation to warmer temperatures caused a reduction in temperature sensitivity that resulted in slower rates at high leaf temperatures. R_d acclimation was similar across plant types. These results suggest that temperature acclimation of the biochemical processes that underlie plant carbon exchange is common across different plant types, but that acclimation to warmer temperatures tends to have a relatively greater positive effect on the processes most limiting to carbon assimilation, which differ by plant type. The acclimation responses observed here suggest that warmer conditions should lead to increased rates of carbon assimilation when water and nutrients are not limiting.     

The responses of microbial temperature relationships to seasonal change and winter warming in a temperate grassland

Microorganisms dominate the decomposition of organic matter and their activities are strongly influenced by temperature. As the carbon (C) flux from soil to the atmosphere due to microbial activity is substantial, understanding temperature relationships of microbial processes is critical. It has been shown that microbial temperature relationships in soil correlate with the climate, and microorganisms in field experiments become more warm‐tolerant in response to chronic warming. It is also known that microbial temperature relationships reflect the seasons in aquatic ecosystems, but to date this has not been investigated in soil. Although climate change predictions suggest that temperatures will be mostly affected during winter in temperate ecosystems, no assessments exist of the responses of microbial temperature relationships to winter warming. We investigated the responses of the temperature relationships of bacterial growth, fungal growth, and respiration in a temperate grassland to seasonal change, and to 2 years’ winter warming. The warming treatments increased winter soil temperatures by 5–6°C, corresponding to 3°C warming of the mean annual temperature. Microbial temperature relationships and temperature sensitivities (Q₁₀) could be accurately established, but did not respond to winter warming or to seasonal temperature change, despite significant shifts in the microbial community structure. The lack of response to winter warming that we demonstrate, and the strong response to chronic warming treatments previously shown, together suggest that it is the peak annual soil temperature that influences the microbial temperature relationships, and that temperatures during colder seasons will have little impact. Thus, mean annual temperatures are poor predictors for microbial temperature relationships. Instead, the intensity of summer heat‐spells in temperate systems is likely to shape the microbial temperature relationships that govern the soil‐atmosphere C exchange.     

Thermal acclimation of shoot respiration in an Arctic woody plant species subjected to 22 years of warming and altered nutrient supply

Despite concern about the status of carbon (C) in the Arctic tundra, there is currently little information on how plant respiration varies in response to environmental change in this region. We quantified the impact of long‐term nitrogen (N) and phosphorus (P) treatments and greenhouse warming on the short‐term temperature (T) response and sensitivity of leaf respiration (R), the high‐T threshold of R, and associated traits in shoots of the Arctic shrub Betula nana in experimental plots at Toolik Lake, Alaska. Respiration only acclimated to greenhouse warming in plots provided with both N and P (resulting in a ~30% reduction in carbon efflux in shoots measured at 10 and 20 °C), suggesting a nutrient dependence of metabolic adjustment. Neither greenhouse nor N+P treatments impacted on the respiratory sensitivity to T (Q₁₀); overall, Q₁₀ values decreased with increasing measuring T, from ~3.0 at 5 °C to ~1.5 at 35 °C. New high‐resolution measurements of R across a range of measuring Ts (25–70 °C) yielded insights into the T at which maximal rates of R occurred (T_max). Although growth temperature did not affect T_max, N+P fertilization increased T_max values ~5 °C, from 53 to 58 °C. N+P fertilized shoots exhibited greater rates of R than nonfertilized shoots, with this effect diminishing under greenhouse warming. Collectively, our results highlight the nutrient dependence of thermal acclimation of leaf R in B. nana, suggesting that the metabolic efficiency allowed via thermal acclimation may be impaired at current levels of soil nutrient availability. This finding has important implications for predicting carbon fluxes in Arctic ecosystems, particularly if soil N and P become more abundant in the future as the tundra warms.