Stomatal regulation in a changing climate: a field study using Free Air Temperature Increase (FATI) and Free Air CO2 Enrichment (FACE)

This study investigates effects of climate warming (+ 2.5°C ubove ambient) and elevated CO₂ concentration (600 μmol mol^?1) on the stomatal functioning and the water relations of Lolium perenne, using Free Air Temperature Increase (FATI) and Free Air CO₂ Enrichment (FACE). Compared to growth at ambient temperature, whole-season temperature increase reduced leaf stomatal conductance, but only at the top of the canopy (-14.6 and -8.8% at ambient and elevated CO₂, respectively). However, because higher canopy temperature raised the leaf-to-air vapour pressure difference, leaf transpiration rate increased (+28% at ambient and +48% at elevated CO₂) and instantaneous leaf water use efficiency, derived from short-term measurements of assimilation and transpiration rate, declined (-11% at ambient and -13% at elevated CO₂). Nevertheless, at the stand level, growth at + 2.5°C reduced transpiration due to fewer tillers per plant and a smaller leaf area per tiller. This sparser vegetation was also more closely coupled to the atmosphere and maintained a drier internal microclimate. To assess whether the stomatal behaviour observed in this experiment could be explained by prevailing concepts of stomatal functioning, three models were applied (Cowan 1977; Ball, Woodrow & Berry 1987; Leuning 1995). The latter model accounted for the highest proportion of variability in the data (58%) and was insensitive to CO₂ and temperature regime, which suggests that the principles of stomatal regulation are not affected by changes in CO₂ or climate.     

开放式昼夜不同增温对单季稻影响的试验研究

全球变暖趋势日益明朗,且存在明显的季节性差异和昼夜不对称性。水稻是我国最重要的粮食作物,研究水稻生产力对昼夜不同增温的响应与适应对我国未来粮食安全战略决策至关重要。为此,作者在江苏南京设计我国首个稻田开放式增温（FATI：Free Air Temperature Increased) 系统,在2007—2008年对水稻进行昼夜不同增温（全天增温、白天增温和夜间增温）的试验研究。结果表明,该增温系统可以形成4 m2均匀且稳定的增温范围,全天、白天和夜间增温处理平均分别可以使水稻全生育期冠层日均温升高2.0℃、0.6℃和0.9℃。白天增温使水稻全生育期冠层白天温度平均升高1.1℃,夜间增温使夜间温度平均升高1.8℃,与未来的升温幅度相似。全天、白天和夜间增温处理下,水稻冠层温度日较差变化分别为0.1℃、0.6℃和-0.9℃。同时,在该系统的3种增温情景下,水稻分蘖期、孕穗期和灌浆期的田间冠层温度日变化趋势基本与常规对照区一致,全生育期的日平均温度变化趋势也基本一致。两年的增温试验表明,不同增温情景均对水稻的生育进程、成熟期地上生物量和产量产生了明显的影响。其中全天、白天和夜间增温分别使水稻从移栽到始穗平均的日期缩短3.5 d、2 d和2.5 d,但对始穗至成熟期的影响不明显;全天、白天和夜间增温分别使水稻地上生物量降低7.7%、6.6%和2.8%,但差异均不显著;白天和夜间增温分别使水稻产量下降8.9%和4.5%,而全天增温下水稻产量略有上升,但均未达到显著水平。从产量构成来看,增温下有效穗数和结实率呈现递增趋势,每穗粒数和千粒重呈现下降趋势。增温对水稻株高的影响不明显。上述结果表明,该稻田开放式增温系统能满足水稻系统生产力对未来气候变暖响应与适应的试验研究要求,气候变暖对水稻生产力影响的相关模型分析结果也尚需进一步的田间实际增温试验验证,模型所需的相关参数也需要进一步完善。     

Diverting residual biomass to energy use: Quantifying the global warming potential of biogenic CO2 (GWPbCO2)

To calculate the global warming potential of biogenic carbon dioxide emissions (GWP_bCO2) associated with diverting residual biomass to bioenergy use, the decay of annual biogenic carbon pulses into the atmosphere over 100 years was compared between biomass use for energy and its business-as-usual decomposition in agricultural, forestry, or landfill sites. Bioenergy use increased atmospheric CO₂ load in all cases, resulting in a ¹⁰⁰GWP_bCO2 (units of g CO₂e/g biomass CO₂ released) of 0.003 for the fast-decomposing agricultural residues to 0.029 for the slow, 0.084–0.625 for forest residues, and 0.368–0.975 for landfill lignocellulosic biomass. In comparison, carbon emissions from fossil fuels have a ¹⁰⁰GWP of 1.0 g (CO₂e/g fossil CO₂). The fast decomposition rate and the corresponding low ¹⁰⁰GWP_bCO2 values of agricultural residues make them a more climate-friendly feedstock for bioenergy production relative to forest residues and landfill lignocellulosic biomass. This study shows that CO₂ released from the combustion of bioenergy or biofuels made from residual biomass has a greenhouse gas footprint that should be considered in assessing climate impacts.     

Potential effects of interaction between CO2 and temperature on forest landscape response to global warming

Projected temperature increases under global warming could benefit southern tree species by providing them the optimal growing temperature and could be detrimental to northern species by exposing them to the supra optimal growing temperatures. This benefit-detriment trade-off could increase the competitive advantage of southern species in the northern species range and cause the increase or even dominance of southern species in the northern domain. However, the optimum temperature for photosynthesis of C3 plants may increase due to CO₂ enrichment. An increase in the optimum temperature could greatly reduce the benefit-detriment effect. In this study, we coupled a forest ecosystem process model (PnET-II) and a forest GAP model (LINKAGES) with a spatially dynamic forest landscape model (LANDIS-II) to study how an optimum temperature increase could affect forest landscape response due to global warming. We simulated 360 years of forest landscape change in the Boundary Water Canoe Area (BWCA) in northern Minnesota, which is transitional between boreal and temperate forest. Our results showed that, under the control scenario of continuing the historic 1984–1993 mean climate (mainly temperature, precipitation and CO₂), the BWCA will become a spruce-fir dominated boreal forest. However, under the scenario of predicted climatic change [the 2000–2099 climates are predicted by Canadian Climate Center (CCC), followed by 200 years of continuing the predicted 2090–2099 mean climate], the BWCA will become a pine-dominated mixed forest. If the optimum temperature increases gradually with [CO₂] (the increase in optimum temperature is assumed to change gradually from 0 °C in year 2000 to 5 °C in year 2099 when [CO₂] reaches 711 ppm and stabilizes at 5 °C after year 2099), the BWCA would remain a fir-dominated boreal forest in areas with relatively high water-holding capacity, but not in areas with relatively low water-holding capacity. Our results suggest that the [CO₂] induced increases in optimum temperature could substantially reduce forest landscape change caused by global warming. However, not all tree species would be able to successfully adapt to future warming as predicted by CCC, regardless of optimum temperature acclimations.     

The effects of elevated CO2 concentrations on the root growth of Lolium perenne and Trifolium repens grown in a FACE* system

Lolium perenne and Trifolium repens were grown in a Free Air CO₂ Enrichment (FACE) system at elevated (600 μimol mol-¹) and ambient (340 μmol mol-¹) carbon dioxide concentrations during a whole growing season. Using a root ingrowth bag technique the extent to which CO₂ enrichment influenced the growth of L, perenne and T. repens roots under two contrasting nutrient regimes was examined. Root ingrowth bags were inserted for a fixed time into the soil in order to trap roots. It was also possible to follow the mortality of roots in bags inserted for different time intervals. Root ingrowth of both L. perenne and T. repens increased under elevated CO₂ conditions. In L. perenne, root ingrowth decreased with increasing nutrient fertilizer level, but for T. repens the root ingrowth was not affected by the nutrient application rate. Besides biomass measurements, root length estimates were made for T, repens. These showed an increase under elevated CO₂ concentrations. Root decomposition appeared to decrease under elevated CO₂ concentrations. A possible explanation for this effect is the observed changes in tissue composition, such as the increase in the carbon: nitrogen ratio in roots of L. perenne at elevated CO₂ concentrations.     

A comparative analysis of the temperature response of leaf elongation in Bromus stamineus and Lolium perenne plants in the field: intrinsic and size-mediated effects

BACKGROUND AND AIMS: Growth of grass species in temperate-humid regions is restricted by low temperatures. This study analyses the origin (intrinsic or size-mediated) and mechanisms (activity of individual meristems vs. number of active meristems) of differences between Bromus stamineus and Lolium perenne in the response of leaf elongation to moderately low temperatures. METHODS: Field experiments were conducted at Balcarce, Argentina over 2 years (2003 and 2004) using four cultivars, two of B. stamineus and two of L. perenne. Leaf elongation rate (LER) per tiller and of each growing leaf, number of growing leaves and total leaf length per tiller were measured on 15-20 tillers per cultivar, for 12 (2003) or 10 weeks (2004) during autumn and winter. KEY RESULTS: LER was faster in B. stamineus than in L. perenne. In part, this was related to size-mediated effects, as total leaf length per tiller correlated with LER and B. stamineus tillers were 71% larger than L. perenne tillers. However, accounting for size effects revealed intrinsic differences between species in their temperature response. These were based on the number of leaf meristems simultaneously active and not on the (maximum) rate at which individual leaves elongated. Species differences were greater at higher temperatures, being barely notable below 5 degrees C (air temperature). CONCLUSIONS: Bromus stamineus can sustain a higher LER per tiller than L. perenne at air temperatures > 6 degrees C. In the field, this effect would be compounded with time as higher elongation rates lead to greater tiller sizes.     

Advances in understanding the impacts of global warming on marine fishes farmed offshore: Sparus aurata as a case study

Monitoring variations in proteins involved in metabolic processes, oxidative stress responses, cell signalling and protein homeostasis is a powerful tool for developing hypotheses of how environmental variations affect marine organisms' physiology and biology. According to the oxygen- and capacity-limited thermal tolerance hypothesis, thermal acclimation mechanisms such as adjusting the activities of enzymes of intermediary metabolism and of antioxidant defence mechanisms, inducing heat shock proteins (Hsps) or activating mitogen-activated protein kinases may all shift tolerance windows. Few studies have, however, investigated the molecular, biochemical and organismal responses by fishes to seasonal temperature variations in the field to link these to laboratory findings. Investigation of the impacts of global warming on fishes farmed offsore, in the open sea, can provide a stepping stone towards understanding effects on wild populations because they experience similar environmental fluctuations. Over the last 30 years, farming of the gilthead sea bream Sparus aurata (Linnaeus 1758) has become widespread along the Mediterranean coastline, rendering this species a useful case study. Based on available information, the prevailing seasonal temperature variations expose the species to the upper and lower limits of its thermal range. Evidence for this includes oxygen restriction, reduced feeding, reduced responsiveness to environmental stimuli, plus a range of molecular and biochemical indicators that change across the thermal range. Additionally, close relationships between biochemical pathways and seasonal patterns of metabolism indicate a connection between energy demand and metabolic processes on the one hand, and cellular stress responses such as oxidative stress, inflammation and autophagy on the other. Understanding physiological responses to temperature fluctuations in fishes farmed offshore can provide crucial background information for the conservation and successful management of aquaculture resources in the face of global change.     

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.     

Adaptive microevolutionary responses to simulated global warming in Simocephalus vetulus: a mesocosm study

Although several studies suggest the occurrence of microevolutionary responses that may allow local persistence of populations under global warming, rigorous experimental proof is lacking. Here, we combined the realism and rigid, replicated experimental design of a large‐scale mesocosm study where populations of the zooplankter Simocephalus vetulus were exposed for 1 year to different global warming scenarios with a life table experiment under laboratory conditions at three temperatures that eliminated confounding, nongenetic factors. Our results provide solid proof for a rapid microevolutionary response to global warming in both survival and the subcomponents of individual performance (age at reproduction and number of offspring), which may allow populations of S. vetulus to persist locally under predicted scenarios of global warming. Such microevolutionary responses may buffer changes in community structure under global warming and help explain the outcome of previous mesocosm studies finding only marginal effects of global warming at the community level.     

Gross primary production responses to warming,elevated CO2, and irrigation: quantifying the drivers of ecosystem physiology in a semiarid grassland

Determining whether the terrestrial biosphere will be a source or sink of carbon (C) under a future climate of elevated CO₂ (eCO₂) and warming requires accurate quantification of gross primary production (GPP), the largest flux of C in the global C cycle. We evaluated 6 years (2007–2012) of flux‐derived GPP data from the Prairie Heating and CO₂ Enrichment (PHACE) experiment, situated in a grassland in Wyoming, USA. The GPP data were used to calibrate a light response model whose basic formulation has been successfully used in a variety of ecosystems. The model was extended by modeling maximum photosynthetic rate (A_max) and light‐use efficiency (Q) as functions of soil water, air temperature, vapor pressure deficit, vegetation greenness, and nitrogen at current and antecedent (past) timescales. The model fits the observed GPP well (R² = 0.79), which was confirmed by other model performance checks that compared different variants of the model (e.g. with and without antecedent effects). Stimulation of cumulative 6‐year GPP by warming (29%, P = 0.02) and eCO₂ (26%, P = 0.07) was primarily driven by enhanced C uptake during spring (129%, P = 0.001) and fall (124%, P = 0.001), respectively, which was consistent across years. Antecedent air temperature (Tair_ant) and vapor pressure deficit (VPD_ant) effects on A_max (over the past 3–4 days and 1–3 days, respectively) were the most significant predictors of temporal variability in GPP among most treatments. The importance of VPD_ant suggests that atmospheric drought is important for predicting GPP under current and future climate; we highlight the need for experimental studies to identify the mechanisms underlying such antecedent effects. Finally, posterior estimates of cumulative GPP under control and eCO₂ treatments were tested as a benchmark against 12 terrestrial biosphere models (TBMs). The narrow uncertainties of these data‐driven GPP estimates suggest that they could be useful semi‐independent data streams for validating TBMs.     

Possible impact of global warming on cabbage root fly (Delia radicum) activity in the UK

A program for simulating the patterns of egg-laying by populations of the cabbage root fly was used to model the effects of global warming on future cabbage root fly attacks. An increase of 3°C in mean daily temperature would cause the cabbage root fly to become active about a month earlier in the year than at present. Under such conditions, the emergence of flies from the overwintering population would be less synchronised, as the completion of diapause and post-diapause development would occur at the same time in different individuals within the population. However, there would continue to be only three generations of fly each year, even in the south of England. With temperature increases of 5°C or 10°C, the fly would complete four generations each year and aestivation would seriously disrupt egg-laying. These rises in temperature would have a major impact on cabbage root fly activity and would require new strategies for controlling this pest.     

Using the output from global circulation models to predict changes in the distribution and abundance of cereal aphids in Canada: a mechanistic modeling approach

Climate change will alter the abundance and distribution of species. Predicting these shifts is a challenge for ecologists and essential information for the formation of public policy. Here, I use a mechanistic mathematical model of the interaction between grass growth physiology and aphid population dynamics, coupled with the climate change projections from the UK's Hadley Centre HadCM3 global circulation model (GCM) and Canada's Center for Climate Modeling and Analysis CGCM2 GCM to predict the changes in the abundance and distribution of summer cereal aphid populations in wheat-growing regions of Canada. When used with the HadCM3 projections, the model predicts a latitudinal shift northward in abundances but there is longitudinal variation as well. However, when used with the CGCM2 projections the model predicts that continental regions will see a decline while coastal regions will see an increase in summer cereal aphid populations. These effects are stronger under the higher emissions scenarios.     

Frost in a future climate: modelling interactive effects of warmer temperatures and rising atmospheric [CO2] on the incidence and severity of frost damage in a temperate evergreen (Eucalyptus pauciflora)

Although plants are more susceptible to frost damage under elevated atmospheric [CO₂], the importance of frost damage under future, warmer climate scenarios is unknown. Accordingly, we used a model to examine the incidence and severity of frost damage to snow gum (Eucalyptus pauciflora) in a sub‐alpine region of Australia for current and future conditions using the A2 IPCC elevated CO₂ and climate change scenario. An existing model for predicting frost effects on E. pauciflora seedlings was adapted to include effects of elevated [CO₂] on acclimation to freezing temperatures, calibrated with field data, and applied to a study region in Victoria using climate scenario data from CSIRO's Global Climate Model C‐CAM for current (1975–2004) and future (2035–2064) 30 years climate sequences. Temperatures below 0 °C were predicted to occur less frequently while the coldest temperatures (i.e. those below ?8 °C) were almost as common in the future as in the current climate. Both elevated [CO₂] and climate warming affected the timing and rates of acclimation and de‐acclimation of snow gum to freezing temperatures, potentially reducing the length of time that plants are fully frost tolerant and increasing the length of the growing season. Despite fewer days when temperatures fall below 0 °C in the future, with consequently fewer damaging frosts with lower average levels of impact, individual weather sequences resulting in widespread plant mortality may still occur. Furthermore, delayed acclimation due to either warming or rising [CO₂] combined with an early severe frost could lead to more frost damage and higher mortality than would occur in current conditions. Effects of elevated [CO₂] on frost damage were greater in autumn, while warming had more effect in spring. Thus, frost damage will continue to be a management issue for plantation and forest management in regions where frosts persist.     

Elevated carbon dioxide and irrigation effects on water stable aggregates in a Sorghum field: a possible role for arbuscular mycorrhizal fungi

While soil biota and processes are becoming increasingly appreciated as important parameters for consideration in global change studies, the fundamental characteristic of soil structure is a neglected area of research. In a sorghum [Sorghum bicolor (L.) Moench] field experiment in which CO₂[supplied using free‐air CO₂ enrichment (FACE) technology] was crossed factorially with an irrigation treatment, soil aggregate (1–2 mm) water stability increased in response to elevated CO₂. Aggregate water stability was increased by 40% and 20% in response to CO₂, at ample and limited water supply treatments, respectively. Soil hyphal lengths of arbuscular mycorrhizal fungi (AMF) increased strongly (with a threefold increase in the dry treatment) in response to CO₂, and the concentrations of one fraction (easily extractable glomalin, EEG) of the AMF‐produced protein glomalin were also increased. Two fractions of glomalin, and AMF hyphal lengths were all positively correlated with soil aggregate water stability. The present results further support the hypothesis that AMF can become important in global change scenarios. Although in this field study a causal relationship between hyphal length, glomalin and aggregate stability cannot be demonstrated, the present data do suggest that AMF could mediate changes in soil structure under elevated CO₂. This could be of great importance in agricultural systems threatened by erosional soil loss.     

Interactive effects of elevated CO2 and temperature on the leaf-miner Dialectica scalariella Zeller (Lepidoptera: Gracillariidae) in Paterson's Curse, Echium plantagineum (Boraginaceae)

Doubling of the current atmospheric CO₂ concentration, and an increase in global mean annual temperatures of 1.5–6 °C, have been predicted to occur by the end of this century. Whilst the separate effects of CO₂ and temperature on plant–insect interactions have been examined in a number of studies, few have investigated their combined impact. We carried out a factorial experiment to explore the effect of a doubling of CO₂ concentration and a 3 °C temperature increase on the development of a complete generation of the leaf‐miner, Dialectica scalariella, in the host plant Paterson's Curse, Echium plantagineum. Elevated CO₂ increased biomass, reduced leaf N and increased C:N ratios in the host plants. Leaf thickness also increased under elevated CO₂, but only in the high‐temperature treatment. Female D. scalariella did not discriminate between plants grown at the different CO₂ levels when ovipositing, despite the reduction in foliage quality under elevated CO₂. Overall, the negative response of D. scalariella to elevated CO₂ was greater than for many species of free‐living insects, presumably because of the limited mobility imposed by the leaf‐mining habit. Development was accelerated at the high temperature and slowed under elevated CO₂. The net result was a reduction in development time of ～14 days in the elevated CO₂/high temperature treatment, compared to the ambient CO₂/low temperature treatment. Larval survivorship and adult moth weight were both affected by a significant interaction between CO₂ and temperature. At the low temperature, CO₂ had little effect on survivorship, but at the high temperature, survivorship was significantly reduced under elevated CO₂. Similarly, elevated CO₂ had a stronger negative effect on adult moth weight when combined with the high‐temperature treatment. A possible explanation for these results is that the high temperature accelerated insect development to such an extent that the larvae did not have sufficient feeding time to compensate for the poorer quality of the foliage. The frequency with which interactions between CO₂ and temperature affected both plant and insect performance in this study highlights the need for caution when predicting the effects of future climate change on plant–insect interactions from single‐factor experiments.     

Photosynthetic down-regulation in Larrea tridentata exposed to elevated atmospheric CO2: interaction with drought under glasshouse and field (FACE) exposure

The photosynthetic response of Larrea tridentata Cav., an evergreen Mojave Desert shrub, to elevated atmospheric CO₂ and drought was examined to assist in the understanding of how plants from water-limited ecosystems will respond to rising CO₂. We hypothesized that photosynthetic down-regulation would disappear during periods of water limitation, and would, therefore, likely be a seasonally transient event. To test this we measured photosynthetic, water relations and fluorescence responses during periods of increased and decreased water availability in two different treatment implementations: (1) from seedlings exposed to 360, 550, and 700 μmol mol^–1 CO₂ in a glasshouse; and (2) from intact adults exposed to 360 and 550 μmol mol^–1 CO₂ at the Nevada Desert FACE (Free Air CO₂ Enrichment) Facility. FACE and glasshouse well-watered Larrea significantly down-regulated photosynthesis at elevated CO₂, reducing maximum photosynthetic rate (A_max), carboxylation efficiency (CE), and Rubisco catalytic sites, whereas droughted Larrea showed a differing response depending on treatment technique. A_max and CE were lower in droughted Larrea compared with well-watered plants, and CO₂ had no effect on these reduced photosynthetic parameters. However, Rubisco catalytic sites decreased in droughted Larrea at elevated CO₂. Operating C_i increased at elevated CO₂ in droughted plants, resulting in greater photosynthetic rates at elevated CO₂ as compared with ambient CO₂. In well-watered plants, the changes in operating C_i, CE and A_max resulted in similar photosynthetic rates across CO₂ treatments. Our results suggest that drought can diminish photosynthetic down-regulation to elevated CO₂ in Larrea, resulting in seasonally transient patterns of enhanced carbon gain. These results suggest that water status may ultimately control the photosynthetic response of desert systems to rising CO₂.     

Smartphone-integrated field microscopy (SPFM): a low-cost and portable tool to study live biological specimens in the wild

Abstract

Images and videos of organisms recorded in the wild have relevance for biodiversity studies. With the advent of smartphones and their potential integration with microscopy, scientific documentation and recording of organisms has surged to an unprecedented scale. Here we report a novel method, developed by integration of a portable smartphone with a handheld field microscope that we term Smartphone-integrated Field Microscopy (SPFM), to capture images and videos which can be highly useful in field-based biodiversity studies. We firstly describe the design of the method and equipment used, followed by successful field demonstration of the method using a case study of the gymnosperm Ephedra intermedia Schrenk & C. A. Meyer in the Kashmir Himalaya. We then discuss the novelty of our method and its potential applications in biodiversity studies.     

Temperature responses are a window to the physiology of dark respiration: differences between CO2 release and O2 reduction shed light on energy conservation

We showed that temperature responses of dark respiration for foliage of Pinus radiata could be approximated by Arrhenius kinetics, whereby E ₀ determines shape of the exponential response and denotes overall activation energy of respiratory metabolism. Reproducible and predictable deviation from strict Arrhenius kinetics depended on foliage age, and differed between R _CO2 and R _O2. Inhibition of oxygen reduction ( R _O2) by cyanide (inhibiting COX) or SHAM (inhibiting AOX) resulted in reproducible changes of the temperature sensitivity for R _O2, but did not affect R _CO2. Enthalpic growth – preservation of electrons in anabolic products – could be approximated with knowledge of four variables: activation energies ( E ₀) for both R _CO2 and R _O2, and basal rates of respiration at a low reference temperature ( R _REF). Rates of enthalpic growth by P. radiata needles were large in spring due to differences between R _REF of oxidative decarboxylation and that of oxygen reduction, while overall activation energies for the two processes were similar. Later during needle development, enthalpic growth was dependent on differences between E ₀ for R _CO2 as compared with R _O2, and increased E ₀( R _O2) indicated greater contributions of cytochrome oxidase to accompany the switch from carbohydrate sink to source. Temperature-dependent increments in stored energy can be calculated as the difference between R _CO2▵ H _CO2 and R _O2▵ H _O2.