CO2 Stimulation Research Articles

Rapid root closure after fire limits fine root responses to elevated atmospheric CO2 in a scrub oak ecosystem in central Florida, USA

AbstractElevated atmospheric carbon dioxide (CO2) often stimulates the growth of fine roots, yet there are few reports of responses of intact root systems to long‐term CO2 exposure. We investigated the effects of elevated CO2 on fine root growth using open top chambers in a scrub oak ecosystem at Kennedy Space Center, Florida for more than 7 years. CO2 enrichment began immediately after a controlled burn, which simulated the natural disturbance that occurs in this system every 10–15 years. We hypothesized that (1) root abundance would increase in both treatments as the system recovered from fire; (2) elevated CO2 would stimulate root growth; and (3) elevated CO2 would alter root distribution. Minirhizotron tubes were used to measure fine root length density (mm cm−2) every three months. During the first 2 years after fire recovery, fine root abundance increased in all treatments and elevated CO2 significantly enhanced root abundance, causing a maximum stimulation of 181% after 20 months. The CO2 stimulation was initially more pronounced in the top 10 cm and 38–49 cm below the soil surface. However, these responses completely disappeared during the third year of experimental treatment: elevated CO2 had no effect on root abundance or on the depth distribution of fine roots during years 3–7. The results suggest that, within a few years following fire, fine roots in this scrub oak ecosystem reach closure, defined here as a dynamic equilibrium between production and mortality. These results further suggest that elevated CO2 hastens root closure but does not affect maximum root abundance. Limitation of fine root growth by belowground resources – particularly nutrients in this nutrient‐poor soil – may explain the transient response to elevated CO2.

Genioglossal Muscle Response to CO2 Stimulation During NREM Sleep

The objective was to evaluate the responsiveness of upper airway muscles to hypercapnia with and without intrapharyngeal negative pressure during non-rapid eye movement (NREM) sleep and wakefulness. We assessed the genioglossal muscle response to CO2 off and on continuous positive airway pressure (CPAP) (to attenuate negative pressure) during stable NREM sleep and wakefulness in the supine position. Laboratory of the Sleep Medicine Division, Brigham and Women's Hospital. Eleven normal healthy subjects. During wakefulness and NREM sleep, we measured genioglossal electromyography (EMG) on and off CPAP at the normal eupneic level and at levels 5 and 10 mm Hg above the awake eupneic level. We observed that CO2 could increase upper-airway muscle activity during NREM sleep and wakefulness in the supine position with and without intrapharyngeal negative pressure. The application of nasal CPAP significantly decreased genioglossal EMG at all 3 levels of PETCO2 during NREM sleep (13.0 +/- 4.9% vs. 4.6 +/- 1.6% of maximal EMG, 14.6 +/- 5.6% vs. 7.1 +/- 2.3% of maximal EMG, and 17.3 +/- 6.3% vs. 10.2 +/- 3.1% of maximal EMG, respectively). However, the absence of negative pressure in the upper airway did not significantly affect the slope of the pharyngeal airway dilator muscle response to hypercapnia during NREM sleep (0.72 +/- 0.30% vs. 0.79 +/- 0.27% of maximal EMG per mm Hg PCO2, respectively, off and on CPAP). We conclude that both chemoreceptive and negative pressure reflex inputs to this upper airway dilator muscle are still active during stable NREM sleep.

NITROGEN CYCLING DURING SEVEN YEARS OF ATMOSPHERIC CO2ENRICHMENT IN A SCRUB OAK WOODLAND

Experimentally increasing atmospheric CO2 often stimulates plant growth and ecosystem carbon (C) uptake. Biogeochemical theory predicts that these initial responses will immobilize nitrogen (N) in plant biomass and soil organic matter, causing N availability to plants to decline, and reducing the long-term CO2-stimulation of C storage in N limited ecosystems. While many experiments have examined changes in N cycling in response to elevated CO2, empirical tests of this theoretical prediction are scarce. During seven years of postfire recovery in a scrub oak ecosystem, elevated CO2 initially increased plant N accumulation and plant uptake of tracer 15N, peaking after four years of CO2 enrichment. Between years four and seven, these responses to CO2 declined. Elevated CO2 also increased N and tracer 15N accumulation in the O horizon, and reduced 15N recovery in underlying mineral soil. These responses are consistent with progressive N limitation: the initial CO2 stimulation of plant growth immobilized N in plant biomass and in the O horizon, progressively reducing N availability to plants. Litterfall production (one measure of aboveground primary productivity) increased initially in response to elevated CO2, but the CO2 stimulation declined during years five through seven, concurrent with the accumulation of N in the O horizon and the apparent restriction of plant N availability. Yet, at the level of aboveground plant biomass (estimated by allometry), progressive N limitation was less apparent, initially because of increased N acquisition from soil and later because of reduced N concentration in biomass as N availability declined. Over this seven-year period, elevated CO2 caused a redistribution of N within the ecosystem, from mineral soils, to plants, to surface organic matter. In N limited ecosystems, such changes in N cycling are likely to reduce the response of plant production to elevated CO2.

Initial observations of increased requirements for light-energy dissipation in ryegrass (Lolium perenne) when source/sink ratios become high at a naturally grazed free air CO2 enrichment (FACE) site.

Although photosynthetic response to long-term elevated CO2 has been extensively studied, little attention has yet been directed at coordinated adjustments between the use of absorbed light for CO2 fixation, and the dissipation of potentially harmful excess light. In this study, we have performed an initial analysis of photosynthetic light use and excess light dissipation in response to grazing-induced variation in the source / sink ratio in ryegrass (Lolium perenne L.) after 6 years' exposure to Free Air CO2 Enrichment (FACE). Before grazing, when the source / sink ratio was relatively large, significant down-regulation of photosynthetic capacity (Amax) was observed in the FACE leaves compared with control leaves at the same stage of maturity. The decrease in Amax partly offset the direct stimulation of elevated CO2 on light-saturated photosynthesis, and was accompanied by a reduction in photochemical electron flow that was accompanied by a large increase in susceptibility to photoinhibition. This was indicated by large increases in both non-photochemical quenching (NPQ) and the de-epoxidised state of xanthophyll cycle (DEPS), and also by changes in the photochemical reflectance index (PRI). However, no significant increase in the xanthophyll pool size in FACE leaves was observed, despite the apparent large increase in requirements for photodissipation in FACE leaves. After grazing, when the source / sink ratio was relatively small, the CO2 fixation rates in both the FACE and control leaves were, as expected, significantly higher compared with those before grazing, and there was no down-regulation of photosynthetic capacity in the leaves under FACE conditions. In addition, the extent of photodissipation in the FACE and control leaves was not significantly different. Overall, the profile of leaf physiological and biochemical responses supports the hypothesis that the effect of long-term elevated CO2 can be significantly influenced by short-term variation in the source / sink ratio. As the xanthophyll pool size does not change significantly, this poses the question of whether the increased photodissipative demand observed here under even moderately elevated CO2 concentrations may lead to increased plant susceptibility to photoinhibition, and thus to an increased risk of damage to plant function, under conditions of low sink demand. This question clearly deserves further study.

ELEVATED CO2STIMULATES NET ACCUMULATIONS OF CARBON AND NITROGEN IN LAND ECOSYSTEMS: A META-ANALYSIS

The capability of terrestrial ecosystems to sequester carbon (C) plays a critical role in regulating future climatic change yet depends on nitrogen (N) availability. To predict long-term ecosystem C storage, it is essential to examine whether soil N becomes progressively limiting as C and N are sequestered in long-lived plant biomass and soil organic matter. A critical parameter to indicate the long-term progressive N limitation (PNL) is net change in ecosystem N content in association with C accumulation in plant and soil pools under elevated CO2. We compiled data from 104 published papers that study C and N dynamics at ambient and elevated CO2. The compiled database contains C contents, N contents, and C:N ratio in various plant and soil pools, and root:shoot ratio. Averaged C and N pool sizes in plant and soil all significantly increase at elevated CO2 in comparison to those at ambient CO2, ranging from a 5% increase in shoot N content to a 32% increase in root C content. The C and N contents in litter pools are consistently higher in elevated than ambient CO2 among all the surveyed studies whereas C and N contents in the other pools increase in some studies and decrease in other studies. The high variability in CO2-induced changes in C and N pool sizes results from diverse responses of various C and N processes to elevated CO2. Averaged C:N ratios are higher by 3% in litter and soil pools and 11% in root and shoot pools at elevated relative to ambient CO2. Elevated CO2 slightly increases root:shoot ratio. The net N accumulation in plant and soil pools at least helps prevent complete down-regulation of, and likely supports, long-term CO2 stimulation of C sequestration. The concomitant C and N accumulations in response to rising atmospheric CO2 may reflect intrinsic nature of ecosystem development as revealed before by studies of succession over hundreds to millions of years.

Recanalization of Long-lasting Middle Cerebral Artery Occlusion by a Combination of Surgical and Interventional Approaches: Technical Case Report

In strictly selected cases of middle cerebral artery (MCA) occlusion, revascularization by extracranial-intracranial (EC-IC) bypass can be considered. The interventional recanalization of the occlusion under direct surgical control has not been reported in the literature so far. A 39-year-old Caucasian female patient had experienced an ischemic stroke 15 years before she came to our attention. At that time, occlusion of the right MCA was diagnosed by angiography. Her neurological deficit resolved within 6 months. Fifteen years later, the patient experienced repeated numbness of her left-sided extremities, which was refractory to medical treatment. Angiography revealed an occluded M1 segment of the MCA. Perfusion computed tomography without and after CO2 stimulation disclosed impaired cerebrovascular capacity. The patient was scheduled for EC-IC bypass. The MCA tree was exposed, and the occluded portion was found to be 10 mm long. We then decided to reopen the vessel by balloon dilation under direct visual control. A catheter was advanced to the M1 origin, where a glidewire was passed into the vessel lumen. With only a little help from the surgeon, it was surprisingly easy to direct the glidewire through the occluded segment. At this time, flow through the M1 segment was re-established. Flow through the MCA that had occluded for 15 years was re-established. On the basis of our experience, in nonatherosclerotic occlusions, intravascular intervention may be considered.

Differential responses to CO2 and sympathetic stimulation in the cerebral and femoral circulations in humans

The relative importance of CO2 and sympathetic stimulation in the regulation of cerebral and peripheral vasculatures has not been previously studied in humans. We investigated the effect of sympathetic activation, produced by isometric handgrip (HG) exercise, on cerebral and femoral vasculatures during periods of isocapnia and hypercapnia. In 14 healthy males (28.1 +/- 3.7 (mean +/- S.D.) years), we measured flow velocity (VP; transcranial Doppler ultrasound) in the middle cerebral artery during euoxic isocapnia (ISO, +1 mmHg above rest) and two levels of euoxic hypercapnia (HC5, end-tidal P(CO(2)), P(ET,CO2), = +5 mmHg above ISO; HC10, P(ET,CO2) = +10 above ISO). Each P(ET,CO2) level was maintained for 10 min using the dynamic end-tidal forcing technique, during which increases in sympathetic activity were elicited by a 2-min HG at 30% of maximal voluntary contraction. Femoral blood flow (FBF; Doppler ultrasound), muscle sympathetic nerve activity (MSNA; microneurography) and mean arterial pressure (MAP; Portapres) were also measured. Hypercapnia increased VP and FBF by 5.0 and 0.6% mmHg-1, respectively, and MSNA by 20-220%. Isometric HG increased MSNA by 50% and MAP by 20%, with no differences between ISO, HC5 and HC10. During the ISO HG there was an increase in cerebral vascular resistance (CVR; 20 +/- 11%), while VP remained unchanged. During HC5 and HC10 HG, VP increased (13% and 14%, respectively), but CVR was unchanged. In contrast, HG-induced sympathetic stimulation increased femoral vascular resistance (FVR) during ISO, HC5 and HC10 (17-41%), while there was a general decrease in FBF below ISO. The HG-induced increases in MSNA were associated with increases in FVR in all conditions (r = 0.76-0.87), whereas increases in MSNA were associated with increases in CVR only during ISO (r = 0.91). In summary, in the absence of hypercapnia, HG exercise caused cerebral vasoconstriction, myogenically and/or neurally, which was reflected by increases in CVR and a maintained VP. In contrast, HG increased FVR during conditions of ISO, HC5 and HC10. Therefore, the cerebral circulation is more responsive to alterations in PCO2, and less responsive to sympathetic stimulation than the femoral circulation.

High Temperature-induced Sink Limitation Alters Growth and Photosynthetic Acclimation to Elevated CO2 in Bean (Phaseolus vulgaris L.)

Average global surface temperatures are predicted to rise due to increasing atmospheric CO2 and other greenhouse gases. Attempts to predict plant response to CO2 must take into account possible temperature effects on phenology and reproductive sink capacity for carbohydrates. In this study, we investigated the effects of atmospheric CO2 partial pressure [35 Pa ambient CO2 (aCO2) vs. 70 Pa elevated CO2 (eCO2)] and temperature (26/15 vs. 35/21 °C day/night) on short- and long-term net CO2 assimilation (An) and growth of red kidney bean (Phaseolus vulgaris). During early vegetative development [14-31 days after planting (DAP)], An, and relative growth rate (RGR) at eCO2 were significantly greater at the supra-optimum (35/21 °C) than at the optimum (26/15 °C) temperature. At 24 DAP, the CO2 stimulation of An by eCO2 was 49% and 89% at optimum and supra-optimum temperature, respectively, and growth enhancement was 48% and 72% relative to plants grown at aCO2. This high temperature-induced growth enhancement was accompanied by an up-regulation of An of eCO2-grown plants. In contrast, during later reproductive stages (31-68 DAP) the eCO2 stimulation of An was significantly less at the supra-optimum than at optimum temperature. This was associated with reduced seed set, greater leaf carbohydrate accumulation, and down-regulation of An at the higher temperature. At final harvest (68 DAP), the eCO2 stimulation of total dry weight was 31% and 14% at optimum and supra-optimum temperature respectively, and eCO2 stimulation of seed dry weight was 39% and -18% at optimum and supra-optimum temperature, respectively. These data indicate substantial shifts in the response to eCO2 during different phenological stages, and suggest that impaired reproductive development at high temperature could reduce the potential for CO2 stimulation of photosynthesis and productivity in bean and possibly other heat-sensitive species.

Interactive Effects of CO2 and O3 on the Growth of Trisetum flavescens and Trifolium pratense Grown in Monoculture or a Bi-Species Mixture

SUMMARY Golden oat grass (Trisetum flavescens L.) and red clover (Trifolium pratense L.) were grown as monocultures or bi-species mixtures under controlled conditions and exposed to ambient (350 ppm) or elevated (580 ppm) CO2, with or without addition of O3 (diel profile with 150 ppb maximum). Shoot biomass measurements after the initial growth and two re-growth periods were used to determine the specific responses of both species, and the difference in the specific response between monocultures and mixtures. T. pratense was much more responsive to CO2, O3, and their combination, compared to T. flavescens. In the case of O3 but not of CO2, the difference in sensitivity between species was larger in mixture than in monoculture. In contrast to elevated CO2, O3 significantly reduced the root:shoot ratio in the mixture, which could explain the increasing negative effect of O3 on clover with progressing harvests. The relative CO2 stimulation of T. pratense and of the cumulative mixture shoot biomass was larger in the presence than in the absence of O3, which was due to an almost complete protection from O3 stress by elevated CO2. In the mixture, the fraction of T. flavescens was small and increased during the experiment; this increase was most pronounced with O3, but any change in mixture biomass was dominated by the response of T. repens. The results confirm that in grass/legume mixtures legumes are most sensitive to elevated CO2 and O3, but the magnitude of specific responses depends on canopy structure and of plant development. Elevated CO2 minimizes the negative impacts of O3 stress on above- and below-ground plant growth.

Chronic fluoxetine microdialysis into the medullary raphe nuclei of the rat, but not systemic administration, increases the ventilatory response to CO2.

In conscious rats, focal CO2 stimulation of the medullary raphe increases ventilation, whereas interference with serotonergic function here decreases the ventilatory response to systemic hypercapnia. We sought to determine whether repeated administration of a selective serotonin reuptake inhibitor in this region would increase the ventilatory response to hypercapnia in unanesthetized rats. In rats instrumented with electroencephalogram-electromyogram electrodes, 250 or 500 microM fluoxetine or artificial cerebrospinal fluid (aCSF) was microdialyzed into the medullary raphe for 30 min daily over 15 days. To compare focal and systemic treatment, two additional groups of rats received 10 mg x kg(-1) x day(-1) fluoxetine or vehicle systemically. Ventilation was measured in normocapnia and in 7% CO2 before treatment (day 0), acutely (days 1 or 3), on day 7, and on day 15. There was no change in normocapnic ventilation in any treatment group. Rats that received 250 microM fluoxetine microdialysis showed a significant 13% increase in ventilation in wakefulness during hypercapnia on day 7, due to an increase in tidal volume. In rats microdialyzed with 500 microM fluoxetine, there were 16 and 32% increases in minute ventilation during hypercapnia in wakefulness and sleep on day 7, and 20 and 28% increases on day 15, respectively, again due to increased tidal volume. There was no change in the ventilatory response to CO2 in rats microdialyzed with aCSF or in systemically treated rats. Chronic fluoxetine treatment in the medullary raphe increases the ventilatory response to hypercapnia in an unanesthetized rat model, an effect that may be due to facilitation of chemosensitive serotonergic neurons.

Photosynthesis, light and nitrogen relationships in a young deciduous forest canopy under open‐air CO2 enrichment

AbstractLeaf photosynthesis (Ps), nitrogen (N) and light environment were measured on Populus tremuloides trees in a developing canopy under free‐air CO2 enrichment in Wisconsin, USA. After 2 years of growth, the trees averaged 1·5 and 1·6 m tall under ambient and elevated CO2, respectively, at the beginning of the study period in 1999. They grew to 2·6 and 2·9 m, respectively, by the end of the 1999 growing season. Daily integrated photon flux from cloud‐free days (PPFDday,sat) around the lowermost branches was 16·8 ± 0·8 and 8·7 ± 0·2% of values at the top for the ambient and elevated CO2 canopies, respectively. Elevated CO2 significantly decreased leaf N on a mass, but not on an area, basis. N per unit leaf area was related linearly to PPFDday,sat throughout the canopies, and elevated CO2 did not affect that relationship. Leaf Ps light‐response curves responded differently to elevated CO2, depending upon canopy position. Elevated CO2 increased Pssat only in the upper (unshaded) canopy, whereas characteristics that would favour photosynthesis in shade were unaffected by elevated CO2. Consequently, estimated daily integrated Ps on cloud‐free days (Psday,sat) was stimulated by elevated CO2 only in the upper canopy. Psday,sat of the lowermost branches was actually lower with elevated CO2 because of the darker light environment. The lack of CO2 stimulation at the mid‐ and lower canopy was probably related to significant down‐regulation of photosynthetic capacity; there was no down‐regulation of Ps in the upper canopy. The relationship between Psday,sat and leaf N indicated that N was not optimally allocated within the canopy in a manner that would maximize whole‐canopy Ps or photosynthetic N use efficiency. Elevated CO2 had no effect on the optimization of canopy N allocation.

Understanding Fracture Performance by Integrating Well Testing and Fracture Modeling

Summary This paper presents case histories and field data that detail the application of well test data to calibrate fracture models, understand fracture performance, and optimize treatment designs. The paper includes field results from gas reservoirs in the Rocky Mountains and west Texas. The case histories also illustrate the application of postfracture pressure-buildup data to identify reasons for fracture treatment problems and to confirm treatment success. The paper also includes a novel data set that illustrates fracture performance from a liquid CO2 stimulation (no sand, no water) in west Texas. The case histories illustrate the importance of integrating well test data with fracture treatment optimization, showing how this process can improve our ability to reliably make multimillion-dollar decisions concerning proppant selection and fracture treatment size.

The growth response of plants to elevated CO2 under non-optimal environmental conditions.

Under benign environmental conditions, plant growth is generally stimulated by elevated atmospheric CO2 concentrations. When environmental conditions become sub- or supra-optimal for growth, changes in the biomass enhancement ratio (BER; total plant biomass at elevated CO2 divided by plant biomass at the current CO2 level) may occur. We analysed literature sources that studied CO2×environment interactions on the growth of herbaceous species and tree seedlings during the vegetative phase. For each experiment we calculated the difference in BER for plants that were grown under 'optimal' and 'non-optimal' conditions. Assuming that interactions would be most apparent if the environmental stress strongly diminished growth, we scaled the difference in the BER values by the growth reduction due to the stress factor. In our compilation we found a large variability in CO2×environment interactions between experiments. To test the impact of experimental design, we simulated a range of analyses with a plant-to-plant variation in size common in experimental plant populations, in combination with a number of replicates generally used in CO2×environment studies. A similar variation in results was found as in the compilation of real experiments, showing the strong impact of stochasticity. We therefore caution against strong inferences derived from single experiments and suggest rather a reliance on average interactions across a range of experiments. Averaged over the literature data available, low soil nutrient supply or sub-optimal temperatures were found to reduce the proportional growth stimulation of elevated CO2. In contrast, BER increased when plants were grown at low water supply, albeit relatively modestly. Reduced irradiance or high salinity caused BER to increase in some cases and decrease in others, resulting in an average interaction with elevated CO2 that was not significant. Under high ozone concentrations, the relative growth enhancement by elevated CO2 was strongly increased, to the extent that high CO2 even compensated in an absolute way for the harmful effect of ozone on growth. No systematic difference in response was found between herbaceous and woody species for any of the environmental variables considered.

Tree seedling growth in natural deep shade: functional traits related to interspecific variation in response to elevated CO2.

The mechanisms for species-specific growth responses to changes in atmospheric CO2 concentration within narrow ecological groups of species, such as shade-tolerant, late-successional trees, have rarely been addressed and are not well understood. In this study the underlying functional traits for interspecific variation in the biomass response to elevated CO2 were explored for seedlings of five late-successional temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba). The seedlings were grown in the natural forest understorey in very low and low light microsites (an average of 1.3% and 3.4% full sun in this experiment), and were exposed to either current ambient CO2 concentrations, 500, or 660µlCO2l-1 in 36 open-top chambers (OTC) over two growing seasons. Even across the narrow range of successional status and shade tolerance, the study species varied greatly in photosynthesis, light compensation point, leaf dark respiration (R d), leaf nitrogen concentration, specific leaf area (SLA), leaf area ratio (LAR), and biomass allocation among different plant parts, and showed distinct responses to CO2 in these traits. No single species combined all characteristics traditionally considered as adaptive to low light conditions. At very low light, the CO2 stimulation of seedling biomass was related to increased LAR and decreased R d, responses that were observed only in Fagus and Taxus. At slightly higher light levels, interspecific differences in the biomass response to elevated CO2 were reversed and correlated best with leaf photosynthesis. The data provided here contribute to a mechanistic process-based understanding of distinct response patterns in co-occurring tree species to elevated CO2 in natural deep shade. I conclude that the high variation in physiological and morphological traits among late-successional species, and the consequences for their responses to slight changes in resource availability, have previously been underestimated. The commonly used broad definitions of functional groups of species may not be sufficient for the understanding of recruitment success and dynamic changes in species composition of old-growth forests in response to rising concentrations of atmospheric CO2.

Increased understanding of nutrient immobilization in soil organic matter is critical for predicting the carbon sink strength of forest ecosystems over the next 100 years

The terrestrial biosphere is currently thought to be a significant sink for atmospheric carbon (C). However, the future course of this sink under rising [CO2] and temperature is uncertain. Some contrasting possibilities that have been suggested are: that the sink is currently increasing through CO2 fertilization of plant growth but will decline over the next few decades because of CO2 saturation and soil nutrient constraints; that the sink will continue to increase over the next century because rising temperature will stimulate the release of plant-available soil nitrogen (N) through increased soil decomposition; that, alternatively, the sink will not be sustained because the additional soil N released will be immobilized in the soil rather than taken up by plants; or that the sink will soon become negative because loss of soil C through temperature stimulation of soil respiration will override any CO2 or temperature stimulation of plant growth. Soil N immobilization is thus a key process; however, it remains poorly understood. In this paper we use a forest ecosystem model of plant-soil C and N dynamics to gauge the importance of this uncertainty for predictions of the future C sink of forests under rising [CO2] and temperature. We characterize soil N immobilization by the degree of variability of soil N:C ratios assumed in the model. We show that the modeled C sink of a stand of Norway spruce (Picea abies (L.) Karst.) in northern Sweden is highly sensitive to this assumption. Under increasing temperature, the model predicts a strong C sink when soil N:C is inflexible, but a greatly reduced C sink when soil N:C is allowed to vary. In complete contrast, increasing atmospheric [CO2] leads to a much stronger C sink when soil N:C is variable. When both temperature and [CO2] increase, the C sink strength is relatively insensitive to variability in soil N:C; significantly, however, with inflexible soil N:C the C sink is primarily a temperature response whereas with variable soil N:C, it is a combined temperature-CO2 response. Simulations with gradual increases of temperature and [CO2] indicate a sustained C sink over the next 100 years, in contrast to recent claims that the C sink will decline over the next few decades. Nevertheless, in using a relatively simple model, our primary aim is not to make precise predictions of the C sink over the next 100 years, but rather to highlight key areas of model uncertainty requiring further experimental clarification. Here we show that improved understanding of the processes underlying soil N immobilization is essential if we are to predict the future course of the forest carbon sink.

Clearance of filtered fluid from the lung during exercise: role of hyperpnea.

During strenuous exercise in sheep, lung lymph flow increases within seconds and rises to levels 7- to 10-fold over baseline. Concomitant with the flow increase, the lymph protein content rapidly decreases to levels consistent with severe capillary hypertension. This pattern of clearance of filtered fluid is quite different than is seen with the passive capillary hypertension that results from mechanical obstruction of the mitral valve. In passive capillary hypertension, the increase in lymph flow and reduction in lymph protein content develop over several hours. The purpose of this study was to discover if these observed differences in edema clearance are related to the hyperpnea that accompanies exercise. Sheep were instrumented for continuous measurement of pulmonary arterial, left atrial, and systemic pressures, cardiac output by ultrasound, lung lymph flow, and ventilation. First, hemodynamics, ventilatory, and lymph clearance variables were measured during moderate exercise at 2.8 mph on a treadmill. Second, on a separate occasion, sheep were induced to hyperventilate to the same minute ventilation as during exercise, using modest CO2 stimulation. Lymph flow and hemodynamics were unaffected by this hyperpnea. The third arm of the experiment was to raise pulmonary microvascular pressure at rest to the level seen with exercise by means of a balloon catheter placed in the mitral valve. Lymph flow rose and protein content decreased slowly and to a lower degree than seen with exercise despite a comparable microvascular pressure. Finally, left atrial hypertension and induced hyperpnea were combined in sheep at rest, and the resulting lymph flow and protein content were the same as seen with exercise at similar pressures and ventilation. We conclude that hyperpnea is a major mechanism of interstitial liquid clearance during exercise, and may be largely responsible for preventing pulmonary edema that might occur at the high microvascular pressures of strenuous exercise.

Conditional repression of a calcium-activated potassium channel reveals its role in the hypercapnic ventilatory response

An increase in tidal volume is an important feature of the ventilatory response to hypercapnia. The neural substrate for tidal volume augmentation is the frequency of action potentials per inspiratory burst. Small-conductance, calcium-activated potassium channels, which are gated by the increases in intracellular calcium during activation, contribute to the interval between action potentials by generating an afterhyperpolarization current. The generation of mice in which one member of this class of potassium channels (SK3) can be regulated by dietary doxycycline (dox) [1] has enabled its role in the ventilatory response to carbon dioxide to be examined. Experiments were carried out in awake mice at 2 weeks of age. A control period in 100% oxygen was followed by 5 min in 5% CO2, 95% O2. Animals targeted for repression of SK3 (SK3 T/T) on dox showed a greater increase in tidal volume than either SK3 T/T not exposed to dox or wild type. There was no difference in the increase in respiratory rate between the three groups of animals. These results show that SK3 is an important regulator of action potential frequency during CO2 stimulation. They also suggest that medullary neurons which contribute to respiratory pattern may be characterized by SK3 channels while those responsible for rhythm may not.

Constraints to growth of annual nettle (Urtica urens) in an elevated CO2atmosphere: Decreased leaf area ratio and tissue N cannot be explained by ontogenetic drift or mineral N supply

The current literature indicates that the stimulation of relative growth rate (RGR) by an elevated atmospheric CO2concentration is transient.Urtica urensL. was exposed to an elevated atmospheric CO2concentration for 26 days to better understand the factors involved in this constraint to growth. Plants were grown hydroponically without nutrient limitation in controlled‐environment cabinets. Consistent with studies of other C3 species, the initial CO2stimulation of RGR ofU. urenswas not sustained and declined in the early stages of exposure. Whilst the decline in RGR was most strongly linked to a reduction in the CO2stimulation of net assimilation rate (NAR), its initial increase was constrained by an early and persistent reduction in leaf area ratio (LAR) due to a decreased specific leaf area (SLA). The decline in NAR could not be linked to any down‐regulation of photosynthetic capacity of individual leaves, despite an accumulation of soluble sugars in them. The reductions in LAR and SLA reflected an accumulation of structural weight in addition to an accumulation of total non‐structural carbohydrate (TNC). To account for the impact of ontogenetic drift on the partitioning of weight and leaf area, this study extends the usual allometric approach to include an analysis of effects on the vertical placement of regression lines (i.e. their elevations). Using this approach, we argue that CO2‐induced reductions in LAR and SLA cannot be explained by ontogenetic drift. By monitoring the tissue N concentration, external N supply was shown unambiguously to be non‐limiting for growth at any plant size. Nevertheless, tissue N was consistently lower in elevated CO2, independent of both ontogeny and TNC accumulation, raising the possibility that the reductions in NAR, LAR and SLA are related to some internal constraint on N utilization.

Ventilatory vs. dyspneic responses to augmenting and subsequently declining CO2 stimulation in humans.

The PCO2 - ventilation control system is known to reveal an open-loop condition when the subject rebreathes a high level of CO2 (such as 7%) according to the Read method. Equilibrium among receptor tissues, mixed venous and arterial blood as well as end-tidal PCO2 (PETCO2) is assumed to develop rapidly so that steady-state conditions in the stimulus - response relationship can be achieved promptly. In the present experiment, we first observed the effects of an augmenting PETCO2 with the conventional Read method (incrementing run, IR). Subsequently, PETCO2 was lowered at the same rate as that of IR but in an opposite direction (decrementing run, DR). Both the CO2 - ventilatory as well as dyspneic responses were obtained under iso-oxic condition and the results between IR and DR runs were compared.

Photosynthetic responses to CO2 enrichment of four hardwood species in a forest understory.

We compared the CO2- and light-dependence of photosynthesis of four tree species (Acer rubrum, Carya glabra, Cercis canadensis, Liquidambar styraciflua) growing in the understory of a loblolly pine plantation under ambient or ambient plus 200 µl l-1 CO2. Naturally-established saplings were fumigated with a free-air CO2 enrichment system. Light-saturated photosynthetic rates were 159-190% greater for Ce. canadensis saplings grown and measured under elevated CO2. This species had the greatest CO2 stimulation of photosynthesis. Photosynthetic rates were only 59% greater for A. rubrum saplings under CO2 enrichment and Ca. glabra and L. styraciflua had intermediate responses. Elevated CO2 stimulated light-saturated photosynthesis more than the apparent quantum yield. The maximum rate of carboxylation of ribulose-1,5-bisphosphate carboxylase, estimated from gas-exchange measurements, was not consistently affected by growth in elevated CO2. However, the maximum electron transport rate estimated from gas- exchange measurements and from chlorophyll fluorescence, when averaged across species and dates, was approximately 10% higher for saplings in elevated CO2. The proportionately greater stimulation of light-saturated photosynthesis than the apparent quantum yield and elevated rates of maximum electron transport suggests that saplings growing under elevated CO2 make more efficient use of sunflecks. The stimulation of light-saturated photosynthesis by CO2 did not appear to correlate with shade-tolerance ranking of the individual species. However, the species with the greatest enhancement of photosynthesis, Ce. canadensis and L. styraciflua, also invested the greatest proportion of soluble protein in Rubisco. Environmental and endogenous factors affecting N partitioning may partially explain interspecific variation in the photosynthetic response to elevated CO2.