Stable carbon isotopes in tree rings indicate improved water use efficiency and drought responses of a tropical dry forest tree species

Understanding the responses of tropical trees to increasing [CO₂] and climate change is important as tropical forests play an important role in carbon and hydrological cycles. We used stable carbon isotopes (δ¹³C) in tree rings to study the physiological responses of a tropical dry forest tree species in southern Mexico, Mimosa acantholoba to changes in atmospheric [CO₂] and variation in climate. Based on annual records of tree ring δ¹³C, we calculated intrinsic water use efficiency (W _i) and intercellular [CO₂] (c _i). Our results showed that trees responded strongly to the increase in atmospheric [CO₂] over the last four decades; W _i increased dramatically by 40%, while c _i remained largely constant. The maintenance of a constant c _i indicates that photosynthetic rates are unlikely to have increased in response to higher [CO₂], and that improvements in W _i are probably due to a reduction in stomatal conductance. This may have large consequences for the hydrological cycle. Inter-annual variation in c _i was strongly correlated with total annual rainfall (r = 0.70), and not influenced by temperature, solar radiation or cloud cover. Our results show that δ¹³C in tree rings of tropical dry forest trees may be a powerful tool to evaluate long-term responses of trees to increasing [CO₂] and to variation in climate.     

Influence of cold soil and snowcover on photosynthesis and leaf conductance in two Rocky Mountain conifers

Summary The influence of cold soil and snowcover on photosynthesis and conductance of Picea engelmannii and Pinus contorta was investigated early in the growing season in the Medicine Bow Mountains, Wyoming, USA. Trees of both species growing in cold soil (<1°C) associated with snowpack had 25–40% lower leaf photosynthesis than trees in warm soils (>10°C). In cold soils leaf conductance of both species was lower, but more so in Pinus, leading to lower intercellular CO₂ concentrations and greater stomatal limitation of photosynthesis. Soil temperature had no effect on predawn and midday shoot water potentials of Pinus and Picea and lower photosynthesis and conductance did not appear to be a result of lower bulk shoot water potential. Predawn, as well as midday, water potentials of Pinus were consistently higher than Picea suggesting that Pinus may have deeper roots, although trenching experiments indicated young Picea trees have more extensive lateral root systems than similar sized Pinus trees. Young Picea trees (<2 m in height) in snowbanks were capable of utilizing warmer soil 4 m from their base. Under similar conditions Pinus in snowbanks had lower photosynthesis and conductance than controls and Pinus did not appear capable of utilizing warmer soils nearby. Under full sunlight, PPFD reflected from the snow surface was 400–1400 mol m^-2 s^-1 higher than from snow-free surfaces. This reflected light resulted in a 10%–20% increase in photosynthesis of Picea. The beneficial effect of reflected light was apparent whether or not photosynthesis was reduced by low soil temperatures.     

Variation in the δ13C of foliage of Pinus sylvestris L. in relation to climate and additions of nitrogen: analysis of a 32-year chronology

We report an analysis of both the long‐ and short‐term drivers of the carbon (C) isotope composition (δ¹³C) values of current year needles of Pinus sylvestris L. linked to changing atmospheric carbon dioxide (CO₂) concentrations (c_a) and climate using data from a uniquely long‐term nitrogen (N) fertilization experiment in the north of Sweden (consisting of three N dosage levels and a control treatment) from 1970 until 2002. N loading produced trees with less negative δ¹³C of foliage, by around 0.45‰ on average, with the difference in δ¹³C between control and N treatments not dependant upon N dosage. The average δ¹³C values decreased at a rate of around 0.03‰ yr⁻¹, even after accounting for the Suess effect (the decrease in the atmospheric CO₂δ¹³C due to anthropogenic emissions of isotopically light CO₂). This decrease is large enough to cause a significant, progressive change in the δ¹³C down through a soil profile. Modelled values of plant intrinsic water use efficiency (WUE_i) and the ratio of leaf internal to external [CO₂] (c_i/c_a) showed that this was the result of c_i increasing in parallel with c_a (while c_i/c_a increased), thus causing little change in WUE_i over the 32 years of study. The residuals from the relationships between year and δ¹³C were used to examine the impact of climate on the interannual variation of C isotope composition of needles. This included the use of a fire hazard index (FHI) model, which integrates climatic factors known to influence plant stomatal conductance and hence δ¹³C. The FHI produced the best fit with δ¹³C values when climate data were averaged over the whole growth season (for control plots) and for July for all the N treatments, explaining ca. 60% of the total interannual variation in δ¹³C. Further, trees from the N treatments appeared more susceptible to air‐humidity‐based climate parameters, as seen from higher correlation coefficients, than were control trees. Thus, our data suggest the possibility of increased susceptibility to drought conditions in ecosystems with moderate to high N deposition rates. Also, there is the possibility that, because there was no apparent change in WUE_i of P. sylvestris in this ecosystem over the last 32 years, the rate of sequestration of C into boreal ecosystems may not increase with c_a, as has been predicted.     

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.     

Homeostatic gas-exchange parameters inferred from 13C/12C in tree rings of conifers

The CO₂ concentration of the atmosphere has increased by almost 30% in the past two centuries, with most of the increase (>5 Pa) during the past 60 years. Controlled environment studies of crop plants dependent on the C₃ photosynthetic pathway indicate that an increase of this magnitude would enhance net photosynthesis, reduce stomatal conductance, and increase the difference in CO₂ concentration across the stomata, i.e., CO₂ concentration outside the leaf to that within (c _a-c _i). Here we report evidence, based on stable isotope composition of tree rings from three species of field-grown, native conifer trees, that the trees have indeed responded. However, rather than increasing c _a-c _i, intercellular CO₂ concentrations have shifted upward to match the rise in atmospheric concentrations, holding c _a-c _i constant. No differences were detected among Douglas-fir (Pseudotsuga menziesii), ponderosa pine (Pinus ponderosa), or western white pine (Pinus monticola). The values of c _a-c _i were inferred from stable carbon isotope ratio (¹³C) of tree ring holocellulose adjusted for the 0.6–2.6 difference between holocellulose and whole sapwood. The cellulose extraction removed contaminants deposited in the tree ring after it formed and the adjustment corrected for the enrichment of cellulose relative to whole tissue. The whole sapwood values were then adjusted for bublished estimates of past atmospheric ¹³CO₂ and CO₂ concentrations. To avoid confounding tree age with CO₂, cellulose deposited by saplings in the 1980s was compared to cellulose deposited in the inner rings of nature trees when the mature trees were saplings, between 1910–1929 and 1941–1970; thus saplings were compared to saplings. In a separate analysis, the juvenile effect, which describes the tendency for ¹³C to increase in the first decades of a tree's life, was quantified independent of source CO₂ effects. This study provides evidence that conifers have undergone adjustments in the intercellular CO₂ concentration that have maintained c _a-c _i constant. Based on these results and others, we suggest that c _a-c _i, which has also been referred to as the intrinsic water-use efficiency, should be considered a homeostatic gas-exchange set point for these conifer species.     

The carbon fertilization effect over a century of anthropogenic CO2 emissions: higher intracellular CO2 and more drought resistance among invasive and native grass species contrasts with increased water use efficiency for woody plants in the US Southwest

From 1890 to 2015, anthropogenic carbon dioxide emissions have increased atmospheric CO₂ concentrations from 270 to 400 mol mol^?1. The effect of increased carbon emissions on plant growth and reproduction has been the subject of study of free‐air CO₂ enrichment (FACE) experiments. These experiments have found (i) an increase in internal CO₂ partial pressure (c_i) alongside acclimation of photosynthetic capacity, (ii) variable decreases in stomatal conductance, and (iii) that increases in yield do not increase commensurate with CO₂ concentrations. Our data set, which includes a 115‐year‐long selection of grasses collected in New Mexico since 1892, is consistent with an increased c_i as a response to historical CO₂ increase in the atmosphere, with invasive species showing the largest increase. Comparison with Palmer Drought Sensitivity Index (PDSI) for New Mexico indicates a moderate correlation with Δ¹³C (r² = 0.32, P < 0.01) before 1950, with no correlation (r² = 0.00, P = 0.91) after 1950. These results indicate that increased c_i may have conferred some drought resistance to these grasses through increased availability of CO₂ in the event of reduced stomatal conductance in response to short‐term water shortage. Comparison with C₃ trees from arid environments (Pinus longaeva and Pinus edulis in the US Southwest) as well as from wetter environments (Bromus and Poa grasses in New Mexico) suggests differing responses based on environment; arid environments in New Mexico see increased intrinsic water use efficiency (WUE) in response to historic elevated CO₂ while wetter environments see increased c_i. This study suggests that (i) the observed increases in c_i in FACE experiments are consistent with historical CO₂ increases and (ii) the CO₂ increase influences plant sensitivity to water shortage, through either increased WUE or c_i in arid and wet environments, respectively.     

Beringia as a glacial refugium for boreal trees and shrubs: new perspectives from mapped pollen data

Aim Beringia, far north‐eastern Siberia and north‐western North America, was largely unglaciated during the Pleistocene. Although this region has long been considered an ice‐age refugium for arctic herbs and shrubs, little is known about its role as a refugium for boreal trees and shrubs during the last glacial maximum (LGM, c. 28,000–15,000 calibrated years before present). We examine mapped patterns of pollen percentages to infer whether six boreal tree and shrub taxa (Populus, Larix, Picea, Pinus, Betula, Alnus/Duschekia) survived the harsh glacial conditions within Beringia. Methods Extensive networks of pollen records have the potential to reveal distinctive temporal–spatial patterns that discriminate between local‐ and long‐distance sources of pollen. We assembled pollen records for 149 lake, peat and alluvial sites from the Palaeoenvironmental Arctic Sciences database, plotting pollen percentages at 1000‐year time intervals from 21,000 to 6000 calibrated years before present. Pollen percentages are interpreted with an understanding of modern pollen representation and potential sources of long‐distance pollen during the glacial maximum. Inferences from pollen data are supplemented by published radiocarbon dates of identified macrofossils, where available. Results Pollen maps for individual taxa show unique temporal‐spatial patterns, but the data for each taxon argue more strongly for survival within Beringia than for immigration from outside regions. The first increase of Populus pollen percentages in the western Brooks Ranges is evidence that Populus trees survived the LGM in central Beringia. Both pollen and macrofossil evidence support Larix survival in western Beringia (WB), but data for Larix in eastern Beringia (EB) are unclear. Given the similar distances of WB and EB to glacial‐age boreal forests in temperate latitudes of Asia and North America, the widespread presence of Picea pollen in EB and Pinus pollen in WB indicates that Picea and Pinus survived within these respective regions. Betula pollen is broadly distributed but highly variable in glacial‐maximum samples, suggesting that Betula trees or shrubs survived in restricted populations throughout Beringia. Alnus/Duschekia percentages show complex patterns, but generally support a glacial refugium in WB. Main conclusions Our interpretations have several implications, including: (1) the rapid post‐glacial migration rate reported for Picea in western Canada may be over estimated, (2) the expansion of trees and shrubs within Beringia should have been nearly contemporaneous with climatic change, (3) boreal trees and shrubs are capable of surviving long periods in relatively small populations (at the lower limit of detection in pollen data) and (4) long‐distance migration may not have been the predominant mode of vegetation response to climatic change in Beringia.     

Soil warming and CO2 enrichment induce biomass shifts in alpine tree line vegetation

Responses of alpine tree line ecosystems to increasing atmospheric CO₂ concentrations and global warming are poorly understood. We used an experiment at the Swiss tree line to investigate changes in vegetation biomass after 9 years of free air CO₂ enrichment (+200 ppm; 2001–2009) and 6 years of soil warming (+4 °C; 2007–2012). The study contained two key tree line species, Larix decidua and Pinus uncinata, both approximately 40 years old, growing in heath vegetation dominated by dwarf shrubs. In 2012, we harvested and measured biomass of all trees (including root systems), above‐ground understorey vegetation and fine roots. Overall, soil warming had clearer effects on plant biomass than CO₂ enrichment, and there were no interactive effects between treatments. Total plant biomass increased in warmed plots containing Pinus but not in those with Larix. This response was driven by changes in tree mass (+50%), which contributed an average of 84% (5.7 kg m^?2) of total plant mass. Pinus coarse root mass was especially enhanced by warming (+100%), yielding an increased root mass fraction. Elevated CO₂ led to an increased relative growth rate of Larix stem basal area but no change in the final biomass of either tree species. Total understorey above‐ground mass was not altered by soil warming or elevated CO₂. However, Vaccinium myrtillus mass increased with both treatments, graminoid mass declined with warming, and forb and nonvascular plant (moss and lichen) mass decreased with both treatments. Fine roots showed a substantial reduction under soil warming (?40% for all roots <2 mm in diameter at 0–20 cm soil depth) but no change with CO₂ enrichment. Our findings suggest that enhanced overall productivity and shifts in biomass allocation will occur at the tree line, particularly with global warming. However, individual species and functional groups will respond differently to these environmental changes, with consequences for ecosystem structure and functioning.     

Long-term c i /c a response of trees in western North America to atmospheric CO2 concentration derived from carbon isotope chronologies

To evaluate how the land carbon reservoir has been responding to the rising CO₂ concentration of the atmosphere, it is important to study how plants in natural forests adjust physiologically to the changing atmospheric conditions. Many experimental studies have addressed this issue, but it has been difficult to scale short-term experimental observations to long-term ecosystem-level responses. This paper derives carbon-isotope-related variables for the past 100–200 years from measurements on trees from natural forests. Calculations show that the c _i/c _a ratios [c _i/c _a is the ratio of the CO₂ concentration (μmol mol⁻¹) in the intercellular space of leaves to that in the atmosphere] of the trees were constant or increased slightly before the 20th century, but changed more rapidly in the 20th century; some increased, some decreased, and some stayed constant. In contrast, the CO₂ concentration inside plant leaves increased monotonically for all trees. Received: 12 June 1997 / Accepted: 29 June 1998     

Conifer stem growth at the altitudinal treeline in response to four years of CO2 enrichment

Northern latitude and upper altitude climatic treelines have received increasing attention given their potential sensitivity to atmospheric and climate change. While greater radial stem growth at treeline sites in recent decades has been attributed largely to increasing temperature, rising atmospheric CO₂ concentration may also be contributing to this growth stimulation. Tree ring increments of mature Larix decidua and Pinus uncinata were measured over 4 years in a free air CO₂ enrichment experiment at treeline in the Swiss Central Alps (2180 m a.s.l.). In addition, a one‐time defoliation treatment in the second year (2002) of the experiment was used to simulate one of the common natural insect outbreak events. In response to elevated atmospheric CO₂, Larix showed a cumulative 4‐year growth response of+41%, with particularly strong responses in the third and fourth year. This increase in radial stem wood growth was the result of more latewood production, in particular, the formation of larger tracheids, rather than a greater number of cells. In contrast, Pinus showed no change in ring width to elevated [CO₂], neither in each of the treatment years, nor in the cumulative response over 4 years, although an increase in tracheid size was observed in the third year. Defoliation led to a pronounced decrease in annual ring width of both species, marked in particular by less latewood production, in the treatment, as well as subsequent years. There was no significant interaction between defoliation and CO₂ enrichment. Although Pinus showed no growth response to CO₂, the positive growth response observed in Larix after 4 years of CO₂ enrichment implies that the sensitivity of treeline trees to global change may not be purely temperature driven. We conclude that the open sparse canopy in the treeline ecotone favours the indeterminate growth strategy of the early successional Larix when neither weather nor carbon are limiting, whereas the later successional Pinus does not show any indication of more vigorous growth under future higher atmospheric CO₂ concentrations.     

The carbon balance in natural and disturbed forests of the southern taiga in central Siberia

Abstract. We evaluated the balance of production and decomposition in natural ecosystems of Pinus sylvestris, Larix sibirica and Betula pendula in the southern boreal forests of central Siberia, using the Yenisei transect. We also investigated whether anthropogenic disturbances (logging, fire and recreation pressure) influence the carbon budget. Pinus and Larix stands up to age class VI act as a net sink for atmospheric carbon. Mineralization rates in young Betula forests exceed rates of uptake via photosynthesis assimilation. Old‐growth stands of all three forest types are CO₂ sources to the atmosphere. The prevalence of old‐growth Larix in the southern taiga suggests that Larix stands are a net source of CO₂. The CO₂ flux to the atmosphere exceeds the uptake of atmospheric carbon via photosynthesis by 0.23 t C.ha^‐1.yr^‐1 (47%). Betula and Pinus forests are net sinks, as photosynthesis exceeds respiration by 13% and 16% respectively. The total carbon flux from Pinus, Larix and Betula ecosystems to the atmosphere is 10 387 thousand tons C.yr^‐1. Net Primary Production (0.935 t‐C.ha^‐1) exceeds carbon release from decomposition of labile and mobile soil organic matter (Rh) by 767 thousand tons C (0.064 t‐C.ha^‐1), so that these forests are net C‐sinks. The emissions due to decomposition of slash (101 thousand tons C; 1.0%) and from fires (0.21%) are very small. The carbon balance of human‐disturbed forests is significantly different. A sharp decrease in biomass stored in Pinus and Betula ecosystems leads to decreased production. As a result, the labile organic matter pool decreased by 6–8 times; course plant residues with a low decomposition rate thus dominate this pool. Annual carbon emissions to the atmosphere from these ecosystems are determined primarily by decomposing fresh litterfall. This source comprises 40–79% of the emissions from disturbed forests compared to only 13–28% in undisturbed forests. The ratio of emissions to production (NPP) is 20–30% in disturbed and 52–76% in undisturbed forests.     

CO2 enrichment in a maturing pine forest: are CO2 exchange and water status in the canopy affected?

A _net, leaf net CO₂ assimilation
c_a, CO₂ concentration of air surrounding a leaf
c_i, leaf intercellular CO₂ concentration
Δ, ¹³C isotope discrimination
δ¹³C, relative stable carbon isotope content
?, ratio of A_net at c_a = 560μmol mol^–1 to A_net at c_a = 360 μmol mol^–1
FACE, free-air CO₂ enrichment
g_w, stomatal conductance to water vapour
Π_i, initial leaf osmotic potential
R_t, relative water content at incipient turgor loss
Ψ_l, xylem water potential of leaves
Ψ_m, soil matric potential

Elevated CO₂ is expected to reduce forest water use as a result of CO₂-induced stomatal closure, which has implications for ecosystem-scale phenomena controlled by water availability. Leaf-level CO₂ and H₂O exchange responses and plant and soil water relations were examined in a maturing loblolly pine (Pinus taeda L.) stand in a free-air CO₂ enrichment (FACE) experiment in North Carolina, USA to test if these parameters were affected by elevated CO₂. Current-year foliage in the canopy was continuously exposed to elevated CO₂ (ambient CO₂+200μmol mol^–1) in free-air during needle growth and development for up to 400 d. Photosynthesis in upper canopy foliage was stimulated by 50–60% by elevated CO₂ compared with ambient controls. This enhancement was similar in current-year, ambient-grown foliage temporarily measured at elevated CO₂ compared with long-term elevated CO₂ grown foliage. Significant photosynthetic enhancement by CO₂ was maintained over a range of conditions except during peak drought. There was no evidence of water savings in elevated CO₂ plots in FACE compared to ambient plots under drought and non-drought conditions. This was supported by evidence from three independent measures. First, stomatal conductance was not significantly different in elevated CO₂ versus ambient trees of P. taeda. Calculations of time-integrated c_i/c_a ratios from analysis of foliar δ¹³C showed that these ratios were maintained in foliage under elevated CO₂. Second, soil moisture was not significantly different between ambient and elevated CO₂ plots during drought. Third, pre-dawn and mid-day leaf water potentials were also unaffected by the seasonal CO₂ exposure, as were tissue osmotic potentials and turgor loss points. Together the results strongly support the hypothesis that maturing P. taeda trees have low stomatal responsiveness to elevated CO₂. Elevated CO₂ effects on water relations in loblolly pine-dominated forest ecosystems may be absent or small apart from those mediated by leaf area. Large photosynthetic enhancements in the upper canopy of P. taeda by elevated CO₂ indicate that this maturing forest may have a large carbon sequestration capacity with limiting water supply.     

A dynamic leaf gas‐exchange strategy is conserved in woody plants under changing ambient CO2: evidence from carbon isotope discrimination in paleo and CO2 enrichment studies

Rising atmospheric [CO₂], c_a, is expected to affect stomatal regulation of leaf gas‐exchange of woody plants, thus influencing energy fluxes as well as carbon (C), water, and nutrient cycling of forests. Researchers have proposed various strategies for stomatal regulation of leaf gas‐exchange that include maintaining a constant leaf internal [CO₂], c_i, a constant drawdown in CO₂ (c_a ? c_i), and a constant c_i/c_a. These strategies can result in drastically different consequences for leaf gas‐exchange. The accuracy of Earth systems models depends in part on assumptions about generalizable patterns in leaf gas‐exchange responses to varying c_a. The concept of optimal stomatal behavior, exemplified by woody plants shifting along a continuum of these strategies, provides a unifying framework for understanding leaf gas‐exchange responses to c_a. To assess leaf gas‐exchange regulation strategies, we analyzed patterns in c_i inferred from studies reporting C stable isotope ratios (δ¹³C) or photosynthetic discrimination (?) in woody angiosperms and gymnosperms that grew across a range of c_a spanning at least 100 ppm. Our results suggest that much of the c_a‐induced changes in c_i/c_a occurred across c_a spanning 200 to 400 ppm. These patterns imply that c_a ? c_i will eventually approach a constant level at high c_a because assimilation rates will reach a maximum and stomatal conductance of each species should be constrained to some minimum level. These analyses are not consistent with canalization toward any single strategy, particularly maintaining a constant c_i. Rather, the results are consistent with the existence of a broadly conserved pattern of stomatal optimization in woody angiosperms and gymnosperms. This results in trees being profligate water users at low c_a, when additional water loss is small for each unit of C gain, and increasingly water‐conservative at high c_a, when photosystems are saturated and water loss is large for each unit C gain.     

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.     

Recent CO2 rise has modified the sensitivity of tropical tree growth to rainfall and temperature

Atmospheric CO₂ (c_a) rise changes the physiology and possibly growth of tropical trees, but these effects are likely modified by climate. Such c_a × climate interactions importantly drive CO₂ fertilization effects of tropical forests predicted by global vegetation models, but have not been tested empirically. Here we use tree‐ring analyses to quantify how c_a rise has shifted the sensitivity of tree stem growth to annual fluctuations in rainfall and temperature. We hypothesized that c_a rise reduces drought sensitivity and increases temperature sensitivity of growth, by reducing transpiration and increasing leaf temperature. These responses were expected for cooler sites. At warmer sites, c_a rise may cause leaf temperatures to frequently exceed the optimum for photosynthesis, and thus induce increased drought sensitivity and stronger negative effects of temperature. We tested these hypotheses using measurements of 5,318 annual rings from 129 trees of the widely distributed (sub‐)tropical tree species, Toona ciliata. We studied growth responses during 1950–2014, a period during which c_a rose by 28%. Tree‐ring data were obtained from two cooler (mean annual temperature: 20.5–20.7°C) and two warmer (23.5–24.8°C) sites. We tested c_a × climate interactions, using mixed‐effect models of ring‐width measurements. Our statistical models revealed several significant and robust c_a × climate interactions. At cooler sites (and seasons), c_a × climate interactions showed good agreement with hypothesized growth responses of reduced drought sensitivity and increased temperature sensitivity. At warmer sites, drought sensitivity increased with increasing c_a, as predicted, and hot years caused stronger growth reduction at high c_a. Overall, c_a rise has significantly modified sensitivity of Toona stem growth to climatic variation, but these changes depended on mean climate. Our study suggests that effects of c_a rise on tropical tree growth may be more complex and less stimulatory than commonly assumed and require a better representation in global vegetation models.     

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 acclimation to long-term exposure to elevated CO2 concentration in Pinus radiata D. Don. is related to age of needles

The effects of CO₂ enrichment on photosynthesis and ribulose-1,5-bisphosphate carboxylase/ oxygenase (Rubisco) in current year and 1-year-old needles on the same branch were studied on Pinus radiata D. Don. trees growing for 4 years in large, open-top chambers at ambient (36 Pa) and elevated (65 Pa) CO₂ partial pressures. At this age trees were 3·5–4 m tall. Measurements made late in the growing cycle (March) showed that photosynthetic rates at the growth CO₂ concentration [(pCO₂)_a] were lower in 1-year-old needles of trees grown at elevated CO₂ concentrations than in those of trees grown at ambient (pCO₂)_a. At elevated CO₂ concentrations V_cmax (maximum carboxylation rate) was reduced by 13% and J_max (RuBP regeneration capacity mediated by maximum electron transport rate) by 17%. This corresponded with photosynthetic rates at the growth (pCO₂)_a of 4·68 ± 0·41 μmol m^–2 s^–1 and 6·15 ± 0·46 μmol m^–2 s^–1 at 36 and 65 Pa, respectively (an enhancement of 31%). In current year needles photosynthetic rates at the growth (pCO₂)_a were 6·2 ± 0·72 μmol m^–2 s^–1 at 36 Pa and 10·15 ± 0·64 μmol m^–2 s^–1 at 65 Pa (an enhancement of 63%). The smaller enhancement of photosynthesis in 1-year-old needles at 65 Pa was accompanied by a reduction in Rubisco activity (39%) and content (40%) compared with that at 36 Pa. Starch and sugar concentrations in 1-year-old needles were not significantly different in the CO₂ treatments. There was no evidence in biochemical parameters for down-regulation at elevated (pCO₂)_a in fully fexpanded needles of the current year cohort. These data show that enhancement of photosynthesis continues to occur in needles after 4 years’ exposure to elevated CO₂ concentrations. Photosynthetic acclimation reduces the degree of this enhancement, but only in needles after 1 year of growth. Thus, responses to elevated CO₂ concentration change during the lifetime of needles, and acclimation may not be apparent in current year needles. This transitory effect is most probably attributable to the effects of developmental stage and proximity to actively growing shoots on sink strength for carbohydrates. The implications of such age-dependent responses are that older trees, in which the contribution of older needles to the photosynthetic biomass is greater than in younger trees, may become progressively more acclimated to elevated CO₂ concentration.     

The calibration of thermocouples for leaf temperature measurements in gas exchange cuvettes

Summary A thermocouple shows a high accuracy in reading temperatures, but it does not always give a true value for the temperature of the leaf against which it is pressed on. Temperature differences between the leaf and the air moving around it cause deviations of the shown temperature from the actual leaf temperature. A method is described to calibrate thermocouples inside gas exchange cuvettes without obstructing the movement of the air around the leaf, so that the heat exchange between thermocouple and air is taken into account. The reading of the leaf temperature in a steady-state porometer was checked by this method at various temperatures of ambient air (T _a ) and of the leaf (T ₁ ) and was found to give an average value of T _a and T ₁ . The effect of incorrect estimation of the leaf temperature on computed diffusive resistances on H₂O (r ^w ) and CO₂ (r ^c ) and intercellular CO₂ partial pressures (p _i ^c ) is discussed.Abbreviations A net CO₂ uptake - E evaporative transpiration - R correlation coefficient - T Temperature - a heat-transfer coefficient - b ratio of heat-transfer coefficients - q heat transfer - p ^c CO₂ partial pressure - p ^w H₂O partial pressure - r ^c diffusive resistance on CO₂ - r ^w diff. resistance on H₂O - a ambient - i intercellular - l leaf - p porometer - s leaf surface (=boundary layer+stomata) - t thermocouple; 1, 2: number of cuvette     

Non-structural carbon compounds in temperate forest trees

The current carbon supply status of temperate forest trees was assessed by analysing the seasonal variation of non‐structural carbohydrate (NSC) concentrations in leaves, branch wood and stem sapwood of 10 tree species (six deciduous broad‐leafed, one deciduous conifer and three evergreen conifer trees) in a temperate forest that is approximately 100 years old. In addition, all woody tissue was analysed for lipids (acylglycerols). The major NSC fractions were starch, sucrose, glucose and fructose, with other carbohydrates (e.g. raffinose and stachyose) and sugar alcohols (cyclitols and sorbitol) playing only a minor quantitative role. The radial distribution of NSC within entire stem cores, assessed here for the first time in a direct interspecific comparison, revealed large differences in the size of the active sapwood fraction among the species, reflecting the specific wood anatomy (ring‐porous versus diffuse‐porous xylem). The mean minimum NSC concentrations in branch wood during the growing season was 55% of maximum, and even high NSC concentrations were maintained during times of extensive fruit production in masting Fagus sylvestris. The NSC in stem sapwood varied very little throughout the season (cross species mean never below 67% of maximum), and the small reductions observed were not significant for any of the investigated species. Although some species contained substantial quantities of lipids in woody tissues (‘fat trees’; Tilia, Pinus, Picea, Larix), the lipid pools did not vary significantly across the growing season in any species. On average, the carbon stores of deciduous trees would permit to replace the whole leave canopy four times. These data imply that there is not a lot of leeway for a further stimulation of growth by ongoing atmospheric CO₂ enrichment. The classical view that deciduous trees rely more on C‐reserves than evergreen trees, seems unwarranted or has lost its justification due to the greater than 30% increase in atmospheric CO₂ concentrations over the last 150 years.     

Share the wealth: Trees with greater ectomycorrhizal species overlap share more carbon

The mutualistic symbiosis between forest trees and ectomycorrhizal fungi (EMF) is among the most ubiquitous and successful interactions in terrestrial ecosystems. Specific species of EMF are known to colonize specific tree species, benefitting from their carbon source, and in turn, improving their access to soil water and nutrients. EMF also form extensive mycelial networks that can link multiple root‐tips of different trees. Yet the number of tree species connected by such mycelial networks, and the traffic of material across them, are just now under study. Recently we reported substantial belowground carbon transfer between Picea, Pinus, Larix and Fagus trees in a mature forest. Here, we analyze the EMF community of these same individual trees and identify the most likely taxa responsible for the observed carbon transfer. Among the nearly 1,200 EMF root‐tips examined, 50%–70% belong to operational taxonomic units (OTUs) that were associated with three or four tree host species, and 90% of all OTUs were associated with at least two tree species. Sporocarp ¹³C signals indicated that carbon originating from labelled Picea trees was transferred among trees through EMF networks. Interestingly, phylogenetically more closely related tree species exhibited more similar EMF communities and exchanged more carbon. Our results show that belowground carbon transfer is well orchestrated by the evolution of EMFs and tree symbiosis.