Disparate effects of global‐change drivers on mountain conifer forests: warming‐induced growth enhancement in young trees vs. CO2 fertilization in old trees from wet sites

Theory predicts that the postindustrial rise in the concentration of CO₂ in the atmosphere (c_a) should enhance tree growth either through a direct fertilization effect or indirectly by improving water use efficiency in dry areas. However, this hypothesis has received little support in cold‐limited and subalpine forests where positive growth responses to either rising c_a or warmer temperatures are still under debate. In this study, we address this issue by analyzing an extensive dendrochronological network of high‐elevation Pinus uncinata forests in Spain (28 sites, 544 trees) encompassing the whole biogeographical extent of the species. We determine if the basal area increment (BAI) trends are linked to climate warming and increased c_a by focusing on region‐ and age‐dependent responses. The largest improvement in BAI over the past six centuries occurred during the last 150 years affecting young trees and being driven by recent warming. Indeed, most studied regions and age classes presented BAI patterns mainly controlled by temperature trends, while growing‐season precipitation was only relevant in the driest sites. Growth enhancement was linked to rising c_a in mature (151–300 year‐old trees) and old‐mature trees (301–450 year‐old trees) from the wettest sites only. This finding implies that any potential fertilization effect of elevated c_a on forest growth is contingent on tree features that vary with ontogeny and it depends on site conditions (for instance water availability). Furthermore, we found widespread growth decline in drought‐prone sites probably indicating that the rise in c_a did not compensate for the reduction in water availability. Thus, warming‐triggered drought stress may become a more important direct driver of growth than rising c_a in similar subalpine forests. We argue that broad approaches in biogeographical and temporal terms are required to adequately evaluate any effect of rising c_a on forest growth.     

Risky future for Mediterranean forests unless they undergo extreme carbon fertilization

Forest performance is challenged by climate change but higher atmospheric [CO₂] (c_a) could help trees mitigate the negative effect of enhanced water stress. Forest projections using data assimilation with mechanistic models are a valuable tool to assess forest performance. Firstly, we used dendrochronological data from 12 Mediterranean tree species (six conifers and six broadleaves) to calibrate a process‐based vegetation model at 77 sites. Secondly, we conducted simulations of gross primary production (GPP) and radial growth using an ensemble of climate projections for the period 2010–2100 for the high‐emission RCP8.5 and low‐emission RCP2.6 scenarios. GPP and growth projections were simulated using climatic data from the two RCPs combined with (i) expected c_a; (ii) constant c_a = 390 ppm, to test a purely climate‐driven performance excluding compensation from carbon fertilization. The model accurately mimicked the growth trends since the 1950s when, despite increasing c_a, enhanced evaporative demands precluded a global net positive effect on growth. Modeled annual growth and GPP showed similar long‐term trends. Under RCP2.6 (i.e., temperatures below +2 °C with respect to preindustrial values), the forests showed resistance to future climate (as expressed by non‐negative trends in growth and GPP) except for some coniferous sites. Using exponentially growing c_a and climate as from RCP8.5, carbon fertilization overrode the negative effect of the highly constraining climatic conditions under that scenario. This effect was particularly evident above 500 ppm (which is already over +2 °C), which seems unrealistic and likely reflects model miss‐performance at high c_a above the calibration range. Thus, forest projections under RCP8.5 preventing carbon fertilization displayed very negative forest performance at the regional scale. This suggests that most of western Mediterranean forests would successfully acclimate to the coldest climate change scenario but be vulnerable to a climate warmer than +2 °C unless the trees developed an exaggerated fertilization response to [CO₂].     

Photosynthetic capacity and leaf nitrogen decline along a controlled climate gradient in provenances of two widely distributed Eucalyptus species

Climate is an important factor limiting tree distributions and adaptation to different thermal environments may influence how tree populations respond to climate warming. Given the current rate of warming, it has been hypothesized that tree populations in warmer, more thermally stable climates may have limited capacity to respond physiologically to warming compared to populations from cooler, more seasonal climates. We determined in a controlled environment how several provenances of widely distributed Eucalyptus tereticornis and E. grandis adjusted their photosynthetic capacity to +3.5°C warming along their native distribution range (~16–38°S) and whether climate of seed origin of the provenances influenced their response to different growth temperatures. We also tested how temperature optima (T_opt) of photosynthesis and J_max responded to higher growth temperatures. Our results showed increased photosynthesis rates at a standardized temperature with warming in temperate provenances, while rates in tropical provenances were reduced by about 40% compared to their temperate counterparts. Temperature optima of photosynthesis increased as provenances were exposed to warmer growth temperatures. Both species had ~30% reduced photosynthetic capacity in tropical and subtropical provenances related to reduced leaf nitrogen and leaf Rubisco content compared to temperate provenances. Tropical provenances operated closer to their thermal optimum and came within 3% of the T_opt of J_max during the daily temperature maxima. Hence, further warming may negatively affect C uptake and tree growth in warmer climates, whereas eucalypts in cooler climates may benefit from moderate warming.     

Regional atmospheric CO2 inversion reveals seasonal and geographic differences in Amazon net biome exchange

Understanding tropical rainforest carbon exchange and its response to heat and drought is critical for quantifying the effects of climate change on tropical ecosystems, including global climate–carbon feedbacks. Of particular importance for the global carbon budget is net biome exchange of CO₂ with the atmosphere (NBE), which represents nonfire carbon fluxes into and out of biomass and soils. Subannual and sub‐Basin Amazon NBE estimates have relied heavily on process‐based biosphere models, despite lack of model agreement with plot‐scale observations. We present a new analysis of airborne measurements that reveals monthly, regional‐scale (~1–8 × 10⁶ km²) NBE variations. We develop a regional atmospheric CO₂ inversion that provides the first analysis of geographic and temporal variability in Amazon biosphere–atmosphere carbon exchange and that is minimally influenced by biosphere model‐based first guesses of seasonal and annual mean fluxes. We find little evidence for a clear seasonal cycle in Amazon NBE but do find NBE sensitivity to aberrations from long‐term mean climate. In particular, we observe increased NBE (more carbon emitted to the atmosphere) associated with heat and drought in 2010, and correlations between wet season NBE and precipitation (negative correlation) and temperature (positive correlation). In the eastern Amazon, pulses of increased NBE persisted through 2011, suggesting legacy effects of 2010 heat and drought. We also identify regional differences in postdrought NBE that appear related to long‐term water availability. We examine satellite proxies and find evidence for higher gross primary productivity (GPP) during a pulse of increased carbon uptake in 2011, and lower GPP during a period of increased NBE in the 2010 dry season drought, but links between GPP and NBE changes are not conclusive. These results provide novel evidence of NBE sensitivity to short‐term temperature and moisture extremes in the Amazon, where monthly and sub‐Basin estimates have not been previously available.     

Interactions among shrub cover and the soil microclimate may determine future Arctic carbon budgets

Arctic and Boreal terrestrial ecosystems are important components of the climate system because they contain vast amounts of soil carbon (C). Evidence suggests that deciduous shrubs are increasing in abundance, but the implications for ecosystem C budgets remain uncertain. Using midsummer CO₂ flux data from 21 sites spanning 16° of latitude in the Arctic and Boreal biomes, we show that air temperature explains c. one‐half of the variation in ecosystem respiration (ER) and that ER drives the pattern in net ecosystem CO₂ exchange across ecosystems. Woody sites were slightly stronger C sinks compared with herbaceous communities. However, woody sites with warm soils (> 10 °C) were net sources of CO₂, whereas woody sites with cold soils (< 10 °C) were strong sinks. Our results indicate that transition to a shrub‐dominated Arctic will increase the rate of C cycling, and may lead to net C loss if soil temperatures rise.     

Adaptation of soil microbial growth to temperature: Using a tropical elevation gradient to predict future changes

Terrestrial biogeochemical feedbacks to the climate are strongly modulated by the temperature response of soil microorganisms. Tropical forests, in particular, exert a major influence on global climate because they are the most productive terrestrial ecosystem. We used an elevation gradient across tropical forest in the Andes (a gradient of 20°C mean annual temperature, MAT), to test whether soil bacterial and fungal community growth responses are adapted to long‐term temperature differences. We evaluated the temperature dependency of soil bacterial and fungal growth using the leucine‐ and acetate‐incorporation methods, respectively, and determined indices for the temperature response of growth: Q₁₀ (temperature sensitivity over a given 10oC range) and T_min (the minimum temperature for growth). For both bacterial and fungal communities, increased MAT (decreased elevation) resulted in increases in Q₁₀ and T_min of growth. Across a MAT range from 6°C to 26°C, the Q₁₀ and T_min varied for bacterial growth (Q_10–20 = 2.4 to 3.5; T_min = ?8°C to ?1.5°C) and fungal growth (Q_10–20 = 2.6 to 3.6; T_min = ?6°C to ?1°C). Thus, bacteria and fungi did not differ significantly in their growth temperature responses with changes in MAT. Our findings indicate that across natural temperature gradients, each increase in MAT by 1°C results in increases in T_min of microbial growth by approximately 0.3°C and Q_10–20 by 0.05, consistent with long‐term temperature adaptation of soil microbial communities. A 2°C warming would increase microbial activity across a MAT gradient of 6°C to 26°C by 28% to 15%, respectively, and temperature adaptation of microbial communities would further increase activity by 1.2% to 0.3%. The impact of warming on microbial activity, and the related impact on soil carbon cycling, is thus greater in regions with lower MAT. These results can be used to predict future changes in the temperature response of microbial activity over different levels of warming and over large temperature ranges, extending to tropical regions.     

Seed germination and seedling establishment of an invasive tropical tree species under different climate change scenarios

Increasing air temperature and atmospheric CO₂ levels may affect the distribution of invasive species. Whereas there is wide knowledge on the effect of global change on temperate species, responses of tropical invasive species to these two global change drivers are largely unknown. We conducted a greenhouse experiment on Terminalia catappa L. (Combretaceae), an invasive tree species on Brazilian coastal areas, to evaluate the effects of increased air temperature and CO₂ concentration on seed germination and seedling growth on the island of Santa Catarina (Florianópolis, Brazil). Seeds of the invasive tree were subjected to two temperature levels (ambient and +1.6 °C) and two CO₂ levels (ambient and ~650 ppmv) with a factorial design. Increased temperature enhanced germination rate and shortened germination time of T. catappa seeds. It also increased plant height, number of leaves and above‐ground biomass. By contrast, increased atmospheric CO₂ concentration had no significant effects, and the interaction between temperature and CO₂ concentration did not affect any of the measured traits. Terminalia catappa adapts to a relatively broad range of environmental conditions, being able to tolerate cooler temperatures in its invasive range. As T. catappa is native to tropical areas, global warming might favour its establishment along the coast of subtropical South America, while increased CO₂ levels seem not to have significant effects on seed germination or seedling growth.     

Long tree‐ring chronologies reveal 20th century increases in water‐use efficiency but no enhancement of tree growth at five Iberian pine forests

We investigated the tree growth and physiological response of five pine forest stands in relation to changes in atmospheric CO₂ concentration (c_a) and climate in the Iberian Peninsula using annually resolved width and δ¹³C tree‐ring chronologies since ad 1600. ¹³C discrimination (Δ≈c_i/c_a), leaf intercellular CO₂ concentration (c_i) and intrinsic water‐use efficiency (iWUE) were inferred from δ¹³C values. The most pronounced changes were observed during the second half of the 20th century, and differed between stands. Three sites kept a constant c_i/c_a ratio, leading to significant c_i and iWUE increases (active response to c_a); whereas a significant increase in c_i/c_a resulted in the lowest iWUE increase of all stands at a relict Pinus uncinata forest site (passive response to c_a). A significant decrease in c_i/c_a led to the greatest iWUE improvement at the northwestern site. We tested the climatic signal strength registered in the δ¹³C series after removing the low‐frequency trends due to the physiological responses to increasing c_a. We found stronger correlations with temperature during the growing season, demonstrating that the physiological response to c_a changes modulated δ¹³C and masked the climate signal. Since 1970 higher δ¹³C values revealed iWUE improvements at all the sites exceeding values expected by an active response to the c_a increase alone. These patterns were related to upward trends in temperatures, indicating that other factors are reinforcing stomatal closure in these forests. Narrower rings during the second half of the 20th century than in previous centuries were observed at four sites and after 1970 at all sites, providing no evidence for a possible CO₂‘fertilization’ effect on growth. The iWUE improvements found for all the forests, reflecting both a c_a rise and warmer conditions, seem to be insufficient to compensate for the negative effects of the increasing water limitation on growth.     

Quantifying the effects of drought on abrupt growth decreases of major tree species in Switzerland

Drought entails important effects on tree physiology, which may result in short‐ to long‐term radial growth decreases. While the majority of studies have focused on annual drought‐related variability of growth, relatively little is known about sustained growth decreases following drought years. We apply a statistical framework to identify climatic factors that induce abrupt growth decreases and may eventually result in tree mortality. We used tree‐ring data from almost 500 standing dead trees and 200 living trees in eight sites of the Swiss network of strict forest reserves, including four of the most important Central European tree species (Abies alba, Picea abies, Fagus sylvatica and Quercus spp.). First, to assess short‐term growth responses to drought under various climate and site conditions, we calculated correlations and linear mixed‐effects models between ring‐width indices (RWIs) and drought based on the Standardized Precipitation Evapotranspiration Index (SPEI). Second, to quantify drought effects on abrupt growth decreases, we applied distributed lag nonlinear models (DLNMs), which account for both delayed effects and the nonlinear relationship between the SPEI and the occurrence of abrupt growth decreases. Positive correlations between RWIs and the SPEI indicated short‐term growth responses of all species, particularly at arid sites. Results of the DLNMs revealed species‐specific growth responses to drought. For Quercus spp., abrupt growth decreases were more likely to occur several years following severe drought, whereas for P. abies, A. alba, and F. sylvatica abrupt growth decreases started frequently immediately in the drought year. We conclude that the statistical framework allows for quantifying the effects of drought intensity on the probability of abrupt growth decreases, which ultimately contributes to an improved understanding of climate impacts on forest community dynamics.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.     

Growth, carbon-isotope discrimination, and drought-associated mortality across a Pinus ponderosa elevational transect

Drought‐ and insect‐associated tree mortality at low‐elevation ecotones is a widespread phenomenon but the underlying mechanisms are uncertain. Enhanced growth sensitivity to climate is widely observed among trees that die, indicating that a predisposing physiological mechanism(s) underlies tree mortality. We tested three, linked hypotheses regarding mortality using a ponderosa pine (Pinus ponderosa) elevation transect that experienced low‐elevation mortality following prolonged drought. The hypotheses were: (1) mortality was associated with greater growth sensitivity to climate, (2) mortality was associated with greater sensitivity of gas exchange to climate, and (3) growth and gas exchange were correlated. Support for all three hypotheses would indicate that mortality results at least in part from gas exchange constraints. We assessed growth using basal area increment normalized by tree basal area [basal area increment (BAI)/basal area (BA)] to account for differences in tree size. Whole‐crown gas exchange was indexed via estimates of the CO₂ partial pressure difference between leaf and atmosphere (p_a?p_c) derived from tree ring carbon isotope ratios (δ¹³C), corrected for temporal trends in atmospheric CO₂ and δ¹³C and elevation trends in pressure. Trees that survived the drought exhibited strong correlations among and between BAI, BAI/BA, p_a?p_c, and climate. In contrast, trees that died exhibited greater growth sensitivity to climate than trees that survived, no sensitivity of p_a?p_c to climate, and a steep relationship between p_a?p_c and BAI/BA. The p_a?p_c results are consistent with predictions from a theoretical hydraulic model, suggesting trees that died had a limited buffer between mean water availability during their lifespan and water availability during drought – i.e., chronic water stress. It appears that chronic water stress predisposed low‐elevation trees to mortality during drought via constrained gas exchange. Continued intensification of drought in mid‐latitude regions may drive increased mortality and ecotone shifts in temperate forests and woodlands.     

Long-term drought effects on the thermal sensitivity of Amazon forest trees

The continued functioning of tropical forests under climate change depends on their resilience to drought and heat. However, there is little understanding of how tropical forests will respond to combinations of these stresses, and no field studies to date have explicitly evaluated whether sustained drought alters sensitivity to temperature. We measured the temperature response of net photosynthesis, foliar respiration and the maximum quantum efficiency of photosystem II (F_v/F_m) of eight hyper-dominant Amazonian tree species at the world's longest-running tropical forest drought experiment, to investigate the effect of drought on forest thermal sensitivity. Despite a 0.6°C–2°C increase in canopy air temperatures following long-term drought, no change in overall thermal sensitivity of net photosynthesis or respiration was observed. However, photosystem II tolerance to extreme-heat damage (T₅₀) was reduced from 50.0 ± 0.3°C to 48.5 ± 0.3°C under drought. Our results suggest that long-term reductions in precipitation, as projected across much of Amazonia by climate models, are unlikely to greatly alter the response of tropical forests to rising mean temperatures but may increase the risk of leaf thermal damage during heatwaves.     

Increased water‐use efficiency translates into contrasting growth patterns of Scots pine and sessile oak at their southern distribution limits

In forests, the increase in atmospheric CO₂ concentrations (C_a) has been related to enhanced tree growth and intrinsic water‐use efficiency (iWUE). However, in drought‐prone areas such as the Mediterranean Basin, it is not yet clear to what extent this “fertilizing” effect may compensate for drought‐induced growth reduction. We investigated tree growth and physiological responses at five Scots pine (Pinus sylvestris L.) and five sessile oak (Quercus petraea (Matt.) Liebl.) sites located at their southernmost distribution limits in Europe for the period 1960–2012 using annually resolved tree‐ring width and δ¹³C data to track ecophysiological processes. Results indicated that all 10 natural stands significantly increased their leaf intercellular CO₂ concentration (C_i), and consequently iWUE. Different trends in the theoretical gas‐exchange scenarios as a response to increasing C_a were found: generally, C_i tended to increase proportionally to C_a, except for trees at the driest sites in which C_i remained constant. C_i from the oak sites displaying higher water availability tended to increase at a comparable rate to C_a. Multiple linear models fitted at site level to predict basal area increment (BAI) using iWUE and climatic variables better explained tree growth in pines (31.9%–71.4%) than in oak stands (15.8%–46.8%). iWUE was negatively linked to pine growth, whereas its effect on growth of oak differed across sites. Tree growth in the western and central oak stands was negatively related to iWUE, whereas BAI from the easternmost stand was positively associated with iWUE. Thus, some Q. petraea stands might have partially benefited from the “fertilizing” effect of rising C_a, whereas P. sylvestris stands due to their strict closure of stomata did not profit from increased iWUE and consequently showed in general growth reductions across sites. Additionally, the inter‐annual variability of BAI and iWUE displayed a geographical polarity in the Mediterranean.     

Family‐level variation in early life‐cycle traits of kelp

Temperate kelp forests (Laminarians) are threatened by temperature stress due to ocean warming and photoinhibition due to increased light associated with canopy loss. However, the potential for evolutionary adaptation in kelp to rapid climate change is not well known. This study examined family‐level variation in physiological and photosynthetic traits in the early life‐cycle stages of the ecologically important Australasian kelp Ecklonia radiata and the response of E. radiata families to different temperature and light environments using a family × environment design. There was strong family‐level variation in traits relating to morphology (surface area measures, branch length, branch count) and photosynthetic performance (F_v/F_m) in both haploid (gametophyte) and diploid (sporophyte) stages of the life‐cycle. Additionally, the presence of family × environment interactions showed that offspring from different families respond differently to temperature and light in the branch length of male gametophytes and oogonia surface area of female gametophytes. Negative responses to high temperatures were stronger for females vs. males. Our findings suggest E. radiata may be able to respond adaptively to climate change but studies partitioning the narrow vs. broad sense components of heritable variation are needed to establish the evolutionary potential of E. radiata to adapt under climate change.     

Climate-growth relations of congeneric tree species vary across a tropical vegetation gradient in Brazil

Seasonally dry tropical forests are an important global climatic regulator, a main driver of the global carbon sink dynamics and are predicted to suffer future reductions in their productivity due to climate change. Yet, little is known about how interannual climate variability affects tree growth and how climate-growth responses vary across rainfall gradients in these forests. Here we evaluate changes in climate sensitivity of tree growth along an environmental gradient of seasonally dry tropical vegetation types (evergreen forest – savannah – dry forest) in Northeastern Brazil, using congeneric species of two common neotropical genera: Aspidosperma and Handroanthus. We built tree-ring width chronologies for each species × forest type combinations and explored how growth variability correlated with local (precipitation, temperature) and global (the El Niño Southern Oscillation - ENSO) climatic factors. We also assessed how growth sensitivity to climate and the presence of growth deviations varied along the gradient. Precipitation stimulates tree growth and was the main growth-influencing factor across vegetation types. Trees in the dry forest site showed highest growth sensitivity to interannual variation in precipitation. Temperature and ENSO phenomena correlated negatively with growth and sensitivity to both climatic factors were similar across sites. Negative growth deviations were present and found mostly in the dry-forest species. Our results reveal a dominant effect of precipitation on tree growth in seasonally dry tropical forests and suggest that along the gradient, dry forests are the most sensitivity to drought. These forests may therefore be the most vulnerable to the deleterious effects of future climatic changes. These results highlight the importance of understanding the climatic sensitivity of different tropical forests. This understanding is key to predict the carbon dynamics in tropical regions, and sensitivity differences should be considered when prioritizing conservation measures of seasonally dry topical forests.     

Wood anatomical traits in black spruce reveal latent water constraints on the boreal forest

The effects of climate change on high‐latitude forest ecosystems are complex, making forecasts of future scenarios uncertain. The predicted lengthening of the growing season under warming conditions is expected to increase tree growth rates. However, there is evidence of an increasing sensitivity of the boreal forest to drought stress. To assess the influence of temperature and precipitation on the growth of black spruce (Picea mariana), we investigated long‐term series of wood anatomical traits on 20 trees from four sites along 600 km, the latitudinal range of the closed boreal forest in Quebec, Canada. We correlated the anatomical traits resolved at intraring level with daily temperature, vapor pressure deficit (VPD), and precipitation during the 1943–2010 period. Tree‐ring width, number of cells per ring and cell wall thickness were positively affected by spring and summer daily mean and maximum temperature at the northern sites. These results agree with the well‐known positive effect of high temperatures on tree ring formation at high latitudes. However, we captured, for the first time in this region, the latent impact of water availability on xylem traits. Indeed, in all the four sites, cell lumen area showed positive correlations with daily precipitation (mostly at low latitude), and/or negative correlations with daily mean and maximum temperature and VPD (mostly at high latitude). We inferred that drought, due to high temperatures, low precipitations, or both, negatively affects cell enlargement across the closed boreal forest, including the northernmost sites. The production of tracheids with narrower lumen, potentially more resistant to cavitation, could increase xylem hydraulic safety under a warmer and drier climate. However, this would result in lower xylem conductivity, with consequent long‐term hydraulic deterioration, growth decline, and possibly lead to tree dieback, as observed in other forest ecosystems at lower latitudes.     

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.     

Tree growth variation in the tropical forest: understanding effects of temperature,rainfall and CO2

Tropical forest responses to climatic variability have important consequences for global carbon cycling, but are poorly understood. As empirical, correlative studies cannot disentangle the interactive effects of climatic variables on tree growth, we used a tree growth model (IBTREE) to unravel the climate effects on different physiological pathways and in turn on stem growth variation. We parameterized the model for canopy trees of Toona ciliata (Meliaceae) from a Thai monsoon forest and compared predicted and measured variation from a tree‐ring study over a 30‐year period. We used historical climatic variation of minimum and maximum day temperature, precipitation and carbon dioxide (CO₂) in different combinations to estimate the contribution of each climate factor in explaining the inter‐annual variation in stem growth. Running the model with only variation in maximum temperature and rainfall yielded stem growth patterns that explained almost 70% of the observed inter‐annual variation in stem growth. Our results show that maximum temperature had a strong negative effect on the stem growth by increasing respiration, reducing stomatal conductance and thus mitigating a higher transpiration demand, and – to a lesser extent – by directly reducing photosynthesis. Although stem growth was rather weakly sensitive to rain, stem growth variation responded strongly and positively to rainfall variation owing to the strong inter‐annual fluctuations in rainfall. Minimum temperature and atmospheric CO₂ concentration did not significantly contribute to explaining the inter‐annual variation in stem growth. Our innovative approach – combining a simulation model with historical data on tree‐ring growth and climate – allowed disentangling the effects of strongly correlated climate variables on growth through different physiological pathways. Similar studies on different species and in different forest types are needed to further improve our understanding of the sensitivity of tropical tree growth to climatic variability and change.     

Tree growth declines and mortality were associated with a parasitic plant during warm and dry climatic conditions in a temperate coniferous forest ecosystem

Insects and pathogens are widely recognized as contributing to increased tree vulnerability to the projected future increasing frequency of hot and dry conditions, but the role of parasitic plants is poorly understood even though they are common throughout temperate coniferous forests in the western United States. We investigated the influence of western hemlock dwarf mistletoe (Arceuthobium tsugense) on large (≥45.7 cm diameter) western hemlock (Tsuga heterophylla) growth and mortality in a 500 year old coniferous forest at the Wind River Experimental Forest, Washington State, United States. We used five repeated measurements from a long‐term tree record for 1,395 T. heterophylla individuals. Data were collected across a time gradient (1991–2014) capturing temperature increases and precipitation decreases. The dwarf mistletoe rating (DMR), a measure of infection intensity, varied among individuals. Our results indicated that warmer and drier conditions amplified dwarf mistletoe effects on T. heterophylla tree growth and mortality. We found that heavy infection (i.e., high DMR) resulted in reduced growth during all four measurement intervals, but during warm and dry intervals (a) growth declined across the entire population regardless of DMR level, and (b) both moderate and heavy infections resulted in greater growth declines compared to light infection levels. Mortality rates increased from cooler‐wetter to warmer‐drier measurement intervals, in part reflecting increasing mortality with decreasing tree growth. Mortality rates were positively related to DMR, but only during the warm and dry measurement intervals. These results imply that parasitic plants like dwarf mistletoe can amplify the impact of climatic stressors of trees, contributing to the vulnerability of forest landscapes to climate‐induced productivity losses and mortality events.     

Annual wood production in a tropical rain forest in NE Costa Rica linked to climatic variation but not to increasing CO2

Increased atmospheric [CO₂] could theoretically lead to increased forest productivity (‘CO₂ fertilization’). This mechanism was hypothesized as a possible explanation for biomass increases reported from tropical forests in the last 30+ years. We used unique long‐term records of annually measured stands (eighteen 0.5 ha plots, 10 years) and focal tree species (six species, 24 years) to assess the effects of rainfall, temperature, and atmospheric [CO₂] on annual wood production in a neotropical rain forest. Our study area was a meso‐scale section (600 ha) of old‐growth Tropical Wet Forest in NE Costa Rica. Using the repeated remeasurements we directly assessed the relative effects of interannual climatic variation and increasing atmospheric [CO₂] on wood production. A remarkably simple two‐factor model explained 91% of the interannual variance in stand‐level tree growth; the statistically independent factors were total dry season rainfall (positive effect, r²=0.85) and night‐time temperature (negative effect, r²=0.42). Stand‐level tree mortality increased significantly with night‐time temperature. After accounting for dry season rainfall and night‐time temperature, there was no effect of annual [CO₂] on tree growth in either the stand or focal species data. Tree growth in this Tropical Wet Forest was surprisingly sensitive to the current range of dry season conditions and to variations in mean annual night‐time temperature of 1–2°. Our results suggest that wood production in the lowland rainforests of NE Costa Rica (and by extension in other tropical regions) may be severely reduced in future climates that are only slightly drier and/or warmer.