Expressing leaf water and cellulose oxygen isotope ratios as enrichment above source water reveals evidence of a Péclet effect

Abstract There is an increasing ecological interest in understanding the gradients in H₂ ¹⁸O enrichment in leaf water (i.e. a Péclet effect), because an appreciation of the significance of the Péclet effect is important for improving our understanding of the mechanistic processes affecting the ¹⁸O composition of leaf water and plant organic material. In data sets where both source water and leaf water ¹⁸O data are available, we can evaluate the potential contribution of a Péclet effect. As an example, we recalculate data published earlier by Roden and Ehleringer (1999, Oecologia 121:467–477) as enrichments in leaf water (_L) and cellulose (_cell) above source water. Based on these recalculations, we present support for the relevance of a Péclet effect in leaves. Further, we demonstrate that the subtle variations in _L and _cell caused by a Péclet effect may be masked in experimental systems in which variation in the source water oxygen isotope ratio is considerable.     

Spatial variation in photosynthetic CO(2) carbon and oxygen isotope discrimination along leaves of the monocot triticale (Triticum × Secale) relates to mesophyll conductance and the Péclet effect

Carbon and oxygen isotope discrimination of CO(2) during photosynthesis (Δ(13)C(obs) and Δ(18)O(obs)) were measured along a monocot leaf, triticale (Triticum × Secale). Both Δ(13)C(obs) and Δ(18)O(obs) increased towards the leaf tip. While this was expected for Δ(18)O(obs) , because of progressive enrichment of leaf water associated with the Péclet effect, the result was surprising for Δ(13) C(obs). To explore parameters determining this pattern, we measured activities of key photosynthetic enzymes [ribulose bis-phosphate carboxylase-oxygenase (Rubisco), phosphoenolpyruvate carboxylase (PEPC) and carbonic anhydrase) as well as maximum carboxylation and electron transport rates (V(cmax) and J(max)) along the leaf. Patterns in leaf internal anatomy along the leaf were also quantified. Mesophyll conductance (g(m)) is known to have a strong influence on Δ(13)C(obs) , so we used three commonly used estimation methods to quantify variation in g(m) along the leaf. Variation in Δ(13)C(obs) was correlated with g(m) and chloroplast surface area facing the intercellular air space, but unrelated to photosynthetic enzyme activity. The observed variation could cause errors at higher scales if the appropriate portion of a leaf is not chosen for leaf-level measurements and model parameterization. Our study shows that one-third of the way from the base of the leaf represents the most appropriate portion to enclose in the leaf chamber.     

The dependence of leaf hydraulic conductance on irradiance during HPFM measurements: any role for stomatal response?

This paper examines the dependence of whole leaf hydraulic conductance to liquid water (K(L)) on irradiance when measured with a high pressure flowmeter (HPFM). During HPFM measurements, water is perfused into leaves faster than it evaporates hence water infiltrates leaf air spaces and must pass through stomates in the liquid state. Since stomates open and close under high versus low irradiance, respectively, the possibility exists that K(L) might change with irradiance if stomates close tightly enough to restrict water movement. However, the dependence of K(L) on irradiance could be due to a direct effect of irradiance on the hydraulic properties of other tissues in the leaf. In the present study, K(L) increased with irradiance for 6 of the 11 species tested. Whole leaf conductance to water vapour, g(L), was used as a proxy for stomatal aperture and the time-course of changes in K(L) and g(L) was studied during the transition from low to high irradiance and from high to low irradiance. Experiments showed that in some species K(L) changes were not paralleled by g(L) changes. Measurements were also done after perfusion of leaves with ABA which inhibited the g(L) response to irradiance. These leaves showed the same K(L) response to irradiance as control leaves. These experimental results and theoretical calculations suggest that the irradiance dependence of K(L) is more consistent with an effect on extravascular (and/or vascular) tissues rather than stomatal aperture. Irradiance-mediated stimulation of aquaporins or hydrogel effects in leaf tracheids may be involved.     

Can CO2 enrichment modify the effect of water and high light stress on biomass allocation and relative growth rate of cork oak seedlings?

To test whether the impact of an enriched-CO₂ environment on the growth and biomass allocation of first-season Quercus suber L. seedlings can modify the drought response under shade or sun conditions, seedlings were grown in pots at two CO₂ concentrations × two watering regimes × two irradiances. Compared to CO₂, light and water treatment had greater effects on all morphological traits measured (height, stem diameter, number of leaves, leaf area, biomass fractions). Cork oak showed particularly large increases in biomass in response to elevated CO₂ under low-watered (W−) and high-illuminated conditions (L+). Allocation shifted from shoot to root under increasing irradiance (L+), but was not affected by CO₂. Changes in allocation related to water limitation were only modest, and changed over time. Relative growth rate (RGR) and net assimilation rate (NAR) were significantly greatest in the L+/W+ treatment for both CO₂ concentrations. Changes in RGR were mainly due to NAR. Growth responses to increased light, water or CO₂ were strongest with light, medium with water availability and smallest for CO₂, in terms of RGR. The rise in NAR for light and water treatments was counterbalanced by a decrease in SLA (specific leaf area) and LMF (leaf mass fraction). Results suggest that elevated CO₂ caused cork oak seedlings to improve their performance in dry and high light environments to a greater extent than in well-irrigated and low light ones, thus ameliorating the effects of soil water stress and high light loads on growth.     

Nitrogen fertilization and δ18O of CO2 have no effect on 18O‐enrichment of leaf water and cellulose in Cleistogenes squarrosa (C4) – is VPD the sole control?

The oxygen isotope composition of cellulose (δ¹⁸O_Cel) archives hydrological and physiological information. Here, we assess previously unexplored direct and interactive effects of the δ¹⁸O of CO₂ (δ¹⁸O_CO2), nitrogen (N) fertilizer supply and vapour pressure deficit (VPD) on δ¹⁸O_Cel, ¹⁸O‐enrichment of leaf water (Δ¹⁸O_LW) and cellulose (Δ¹⁸O_Cel) relative to source water, and p_exp_x, the proportion of oxygen in cellulose that exchanged with unenriched water at the site of cellulose synthesis, in a C₄ grass (Cleistogenes squarrosa). δ¹⁸O_CO2 and N supply, and their interactions with VPD, had no effect on δ¹⁸O_Cel, Δ¹⁸O_LW, Δ¹⁸O_Cel and p_exp_x. Δ¹⁸O_Cel and Δ¹⁸O_LW increased with VPD, while p_exp_x decreased. That VPD‐effect on p_exp_x was supported by sensitivity tests to variation of Δ¹⁸O_LW and the equilibrium fractionation factor between carbonyl oxygen and water. N supply altered growth and morphological features, but not ¹⁸O relations; conversely, VPD had no effect on growth or morphology, but controlled ¹⁸O relations. The work implies that reconstructions of VPD from Δ¹⁸O_Cel would overestimate amplitudes of VPD variation, at least in this species, if the VPD‐effect on p_exp_x is ignored. Progress in understanding the relationship between Δ¹⁸O_LW and Δ¹⁸O_Cel will require separate investigations of p_ex and p_x and of their responses to environmental conditions.     

The effect of He−O2 exposure on metabolic rate,thermoregulation and thermal conductance during normothermia and daily torpor

Recently it was proposed that the low metabolic rate during torpor may be better explained by the reduction of thermal conductance than the drop of body temperature or metabolic inhibition. We tested this hypothesis by simultaneously measuring body temperature and metabolic rate as a function of ambient temperature in both torpid and normothermic stripe-faced dunnarts, Sminthopsis macroura (Marsupialia; approx. 25 g body mass), exposed to either air or He–O₂ (21% oxygen in helium) atmospheres. He–O₂ exposure increases the thermal conductance of homeothermic mammals by about twofold in comparison to an air atmosphere without apparent side-effects. Normothermic S. macroura exposed to He–O₂ increased resting metabolic rate by about twofold in comparison to that in air because of the twofold increase in apparent thermal conductance. Torpid S. macroura exposed to He–O₂ at ambient temperatures above the set-point for body temperature showed a completely different metabolic response. In contrast to normothermic individuals, torpid individuals significantly decreased or maintained a similar metabolic rate as those in air although the apparent thermal conductance in He–O₂ was slightly raised. Moreover, the metabolic rate during torpor was only a fraction of that of normothermic individuals although the apparent thermal conductance differed only marginally between normothermia and torpor. Our study shows that a low thermal conductance is not the reason for the low metabolic rates during torpor. It suggests that interrelations between metabolic rate and body temperature of torpid endotherms above the set-point for body temperature differ fundamentally from those of normothermic and homeothermic endotherms.Abbreviations T _a ambient temperature - T _b body temperature - BMR basal metabolic rate - C apparent thermal conductance - He–O ₂ 21% oxygen in helium - MR metabolic rate - MSe mean square-error - RMR festing metabolic rate - TMR metabolic rate during torpor - T difference T _b-T _a - TNZ thermoneutral zone - T _set set-point for body temperature - O ₂ rate of oxygen consumption     

The effect of chemical fertilizer on soil organic carbon renewal and CO2 emission—a pot experiment with maize

Background and Aims

Previous studies have clearly shown substantial increases of soil organic carbon (SOC) in agricultural soils of Yellow River reaches. Those soils did not receive organic fertilizer input, but did receive chemical fertilizer inputs. Thus, to investigate the hypothesis that the observed SOC increases were driven by chemical fertilizer additions, a maize pot experiment was conducted using a Fluvisol that developed under C₃ vegetation in the Yellow River reaches.

Methods

Using the natural ¹³C abundance method we calculated the SOC renewal ratio (C _renewal), and separated total soil organic carbon (TOC) into maize-derived soil organic carbon (SOC_maize) and original soil organic carbon (SOC_original). Carbon dioxide fluxes and microbial biomass carbon (MBC) were determined by closed chamber method and fumigation-extraction method, respectively. The experiment included five treatments: (1) NPK: application of chemical fertilizer NPK; (2) NP, application of chemical fertilizer NP; (3) PK: application of chemical fertilizer PK; (4) NK, application of chemical fertilizer NK; and (5) CK: unfertilized control.

Results

Fertilization increased maize biomass (including grain, straw and root), TOC, C _renewal, SOC_maize, maize-derived carbon (MDC: including SOC_maize, and root and stubble biomass carbon) and MBC, and these values among the treatments ranked NPK>NP>PK>NK>CK. The C _renewal was 5.54–8.50% across the treatments. Fertilization also increased soil CO₂ emission (including root respiration and SOC_original decomposition), while the SOC_original decomposition during the maize growing season only amounted to 74.0–93.4 and 33.5–46.1% of SOC_maize and MDC among the treatments, respectively. Thus input was larger than export, and led to SOC increase. Maize grain and straw biomass were positively and significantly correlated with soil δ¹³C, TOC, C _renewal, SOC_maize, MDC and MBC.

Conclusions

The study suggests that chemical fertilizer application could increase C _renewal by increasing crop-derived C and accelerating original SOC decomposition, and that as long as a certain level of crop yield or aboveground biomass can be achieved, application of chemical fertilizer alone can maintain or increase SOC level in Fluvisol in the Yellow River reaches.     

Observations on the thermal conductance of Adélie (Pygoscelis adeliae) and Humboldt (Spheniscus humboldti) penguins

 Analyses of cooling rates in one Adélie and one Humboldt penguin yielded calculated thermal conductance values of 0.1040 and 0.1672 W(kg ^°C)^-1, respectively. We review the methods used to calculate penguin surface area, an important component in calculating conductance values, and suggest that, in comparative studies of thermal balance, the use of body mass is a better estimator of body size than surface area. Using previously published data on penguin species, we found a significant model to predict thermal conductance from body mass according to: log C= log 0.1083−0.474 log M, where C is minimal specific thermal conductance in W(kg ^°C)^-1 and M is body mass in kilograms. Received: 13 June 1995/Accepted: 11 March 1996     

Gas valves,forests and global change: a commentary on Jarvis (1976) ‘The interpretation of the variations in leaf water potential and stomatal conductance found in canopies in the field’

Microscopic turgor-operated gas valves on leaf surfaces—stomata—facilitate gas exchange between the plant and the atmosphere, and respond to multiple environmental and endogenous cues. Collectively, stomatal activities affect everything from the productivity of forests, grasslands and crops to biophysical feedbacks between land surface vegetation and climate. In 1976, plant physiologist Paul Jarvis reported an empirical model describing stomatal responses to key environmental and plant conditions that predicted the flux of water vapour from leaves into the surrounding atmosphere. Subsequent theoretical advances, building on this earlier approach, established the current paradigm for capturing the physiological behaviour of stomata that became incorporated into sophisticated models of land carbon cycling. However, these models struggle to accurately predict observed trends in the physiological responses of Northern Hemisphere forests to recent atmospheric CO₂ increases, highlighting the need for improved representation of the role of stomata in regulating forest–climate interactions. Bridging this gap between observations and theory as atmospheric CO₂ rises and climate change accelerates creates challenging opportunities for the next generation of physiologists to advance planetary ecology and climate science. This commentary was written to celebrate the 350th anniversary of the journal Philosophical Transactions of the Royal Society.     

The effect of air containing O2 −,O2 +, CO2 − and CO2 + on the growth of seedlings ofHordeum Vulgaris

Equipment was devised which permitted the addition of specific gaseous ions to the atmosphere of plastic chambers in which seedlings of HORDEUM VULGARIS were grown in sand culture supplied with chemically defined nutrient solutions. Moderate densities of O₂ ^– or O₂ ⁺ ions (1.8×10⁴/cm³)in air containing an added 8% of O₂ accelerated the growth rate. A like number of CO₂ ^– or CO₂ ⁺ ions in air containing 8% of CO₂ inhibited growth, impeded the production of chlorophyll and devitalized the young seedlings. Evidence is presented that O₂ ^– and O₂ ⁺ stimulate the production of cytochromes and other Fe-containing enzymes through their action on the plant regulatory system responsible for the control of Fe metabolism. The toxic effect of CO₂ ^– and CO₂ ⁺ cannot be explained as yet.

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Zusammenfassung Eine Apparatur wurde entwickelt, die die Zufuhr von ionisiertem Gas der AtmosphÄre in Kammern gestattet. Darin wurden Keimlinge von HORDEUM VULGARIS in Sand mit chemisch definierten NÄhrlösungen gezüchtet. Konzentrationen von 1,8×10⁴/cm³ O₂ ^– und O₂ ⁺ in Luft mit zusÄtzlich 8% O₂ beschleunigten die Wachstumsrate. Die gleiche Menge CO₂ ^– und CO₂ ⁺ in Luft mit zusÄtzlich 8% CO₂ hemmte die Wachstumsrate, die Bildung von Chlorophyll und entkrÄftigte die Keimlinge. Es wird gezeigt,dass O₂ ^– und O₂ ⁺ die Bildung von Cytochrom und anderen eisenhaltigen Enzymen anregen durcn ihre Wirkung auf das den Fe-Stoffwechsel regulierende System der Pflanze. Die toxische Wirkung von CO₂ ^– und CO₂ ⁺ lÄsst sich noch nicht erklÄren.

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Resume On a construit un appareil permettant d'introduire dans 1'atmosphères des ions de gaz déterminés. On a alors effectué de telles adjonctions à l'air contenu dans des cellules de plastique dans lesquelles on cultivait HORDEUM VULGARIS sur du sable et dans une solution nutritive chimiquement définie. Des densités modérées d'ions O₂ ^– ou O₂ ⁺ (1,8×10⁴/cm³) dans de l'air additionné de 8% d'O₂ accélèrent la croissance. La meme concentration de CO₂ ^– et CO₂ ⁺ additionnée de 8% de CO₂ a ralenti la croissance et la formation de chlorophylle et a diminué la vitalite des plantes nouvellement germées. On démontre que O₂ ^– et O₂ ⁺ active la formation de cytochrome et d'autres enzymes ferreuses par suite de l'action de ces ions sur le système régularisant le métabolisme du fer dans la plante. L'effet toxique du CO₂ ^– et CO₂ ⁺ reste encore inexpliqué.

     

The effect of O2 and CO2 concentration on photosynthesis and glycolate accumulation in bean leaves treated with α-hydroxy-2-pyridinemethanesulfonic acid (α-HPMS), the glycolate oxidase inhibitor

¹⁴CO₂ assimilation, ¹⁴C incorporation into glycolate and glycolate accumulation in -HPMS treated bean leaves at various O₂ and CO₂ concentrations were studied. In 1% CO₂ oxygen concentration had no significant effect on glycolate accumulation and ¹⁴C incorporation into glycolate. In the CO₂ concentration range of 0.03% to 0.01%, increased oxygen concentration decreased not only ¹⁴CO₂ assimilation but also glycolate accumulation and ¹⁴C incorporation into glycolate. In 1% and 0.1% CO₂, no matter what O₂ concentration was supplied, and in 0.03% CO₂ with 2% and 21% O₂, all of the glycolate accumulated was formed from newly assimilated carbon. In 0.01% CO₂ and 2%, 21% and 100% O₂, and in 0.03% CO₂ with 100% O₂, a substantial portion of the glycolic acid that accumulated in leaves originated from endogenous unlabelled substrates. These findings are discussed in terms of possible changes in the ratio of RuBP carboxylation to RuBP oxygenation and of changes of RuBP pool size, induced by changing O₂ and CO₂ concentrations.This work was supported by the Polish Academy of Sciences, Contract No. 10.2.10.     

The reactions of organosulfur transfer reagents with Fe2(CO)9 and the synthesis of the Fe2(CO)6 derivative of α-lipoic acid

Treatment of Fe₂(CO)₉ with sulfur-transfer reagents of the types ImideSSImide and RSSImide where H-imide = phthalimide, succinimide, benzimidazole, morpholine and piperazine, and R  CH₂Ph and CMe₃ leads to cleavage of both the sulfursulfur bond and the sulfurnitrogen bond to give Fe₃(CO)₉S₂ in varying yields, some Fe₂(CO)₆S₂ plus low yields of the appropriate dimers of the type Fe₂(CO)₆(SR)(SR′), where R = R′ = phthal imido, CH₂Ph, CMe₃ and R = CH₂Ph, CMe₃, R′ = phthalimido. The naturally occurring cyclic disulfide D,L-α-lipoic acid, its methyl ester and amide react with Fe₂(CO)₉ to give Fe₂(CO)₆ derivatives wherein the sulfursufur bond has been broken.     

The effect of elevated CO2 on photochemistry and antioxidative defence capacity in wheat depends on environmental growing conditions – A FACE study

The present study examines photosynthesis, photochemistry and low weight molecular antioxidants (ascorbic acid and glutathione) of two Triticum aestivum L. cultivars (H45 and Yitpi) in response to growth under two CO₂ concentrations (elevated CO₂, e[CO₂] vs. ambient CO₂, a[CO₂]), two sowing times (time of sowing 1, TOS1, less stressful growing conditions vs. time of sowing 2, TOS2, more stressful growing conditions) and two water treatments (rain-fed vs. irrigated). The objective was to evaluate (1) if growth under e[CO₂] will alleviate climate stresses such as higher temperature and/or limited water supply thereby reducing the need for photoprotection and concentrations of low weight molecular antioxidants and (2) cultivar-specific responses to combined climate change factors which may be useful to identify intra-specific variation in stress tolerance for future breeding. We compared gas exchange, chlorophyll fluorescence and antioxidative defence compounds (ascorbic acid, glutathione) of flag leaves of Australian Grains Free Air Carbon dioxide Enrichment (AGFACE) grown wheat. When plants were grown under the less stressful growing conditions of TOS1, e[CO₂] increased light saturated net assimilation rates (A_sat) and quantum yield of PSII electron transport (ΦPSII) but decreased thermal energy dissipation (indicated by increased efficiency of open PSII centres, Fv′/Fm′), while antioxidant concentrations did not change. Under the more stressful growing conditions of TOS2, e[CO₂] also increased A_sat (like at TOS1), however, photochemical processes were not affected while antioxidant concentrations (especially ascorbic acid) were decreased. Cultivar specific responses also varied between sowing dates: Only at TOS2 and additional irrigation, antioxidant concentrations were lower in e[CO₂] grown H45 as compared to Yitpi indicating decreased photo-oxidative pressure in H45. These results suggest a photo-protective role of e[CO₂] as well as some intra-specific variability between investigated cultivars in their stress responsiveness, all strongly modified by environmental growing conditions.     

The NOXO1β PX domain preferentially targets PtdIns(4,5)P2 and PtdIns(3,4,5)P3

NOXO1β [NOXO1 (Nox organizer 1) β] is a cytosolic protein that, in conjunction with NOXA1 (Nox activator 1), regulates generation of reactive oxygen species by the NADPH oxidase 1 (Nox1) enzyme complex. NOXO1β is targeted to membranes through an N-terminal PX (phox homology) domain. We have used NMR spectroscopy to solve the structure of the NOXO1β PX domain and surface plasmon resonance (SPR) to assess phospholipid specificity. The solution structure of the NOXO1β PX domain shows greatest similarity to that of the phosphatidylinositol 3-kinase-C2α PX domain with regard to the positions and types of residues that are predicted to interact with phosphatidylinositol phosphate (PtdInsP) head groups. SPR experiments identify PtdIns(4,5)P(2) and PtdIns(3,4,5)P(3) as preferred targets of NOXO1β PX. These findings contrast with previous lipid overlay experiments showing strongest binding to monophosphorylated PtdInsP and phosphatidylserine. Our data suggest that localized membrane accumulation of PtdIns(4,5)P(2) or PtdIns(3,4,5)P(2) may serve to recruit NOXO1β and activate Nox1.     

The effect of potato plant transformation with the gene encoding Δ12-acyl-lipid desaturase on the CO2 exchange and activities of antioxidant enzymes under hypothermia

The effects of potato (Solanum tuberosum L., cv. Desnitsa) plant transformation with the desA gene encoding Δ12-acyl-lipid desaturase from Synechocystis sp. PCC 6803 on the regulation of free-radical processes in relation to plant tolerance to hypothermia are considered. It was shown that the content of polyunsaturated fatty acids (PUFA) in transformed plants was higher than in wild-type ones. In particular, the content of linoleic acid in transformants was higher by 35% and the content of linolenic acid was by 41% higher than in untransformed plants. In addition, transformation induced an increase in the absolute content of C₁₆-PUFA and on the whole resulted in a marked accumulation of membrane lipids. As judged from the values of the damage index and the ratio of photosynthesis to respiration in wild-type and transformed plants under cold treatment, these changes in lipid metabolism favored the protection of coupling membranes, thus preventing plants against free-radical oxidation under low-temperature stress. As a result, the intensity of oxidative stress in transformed plants was much lower than in wild-type ones, whereas antioxidant enzymes (superoxide dismutase, catalase, peroxidase) were not substantially activated under hypothermia.     

The effect of some ions on the activity of β-galactosidase and the inhibitory effect of tris (hydroxymethyl) aminomethane

The stimulating effect of phosphate and the inhibitory effect of tris-HCl on the activity of β-galactosidase inEscherichia coli was studied. The phosphate anion antagonizes the inhibitory effect of chloride. Since a similar effect is displayed by sulphate and arsenate no specific “stimulating” effect of phosphate can take place. The tris cation has also an inhibitory effect which is antagonized by univalent cations (K⁺). The resulting β-galactosidase activity reflects the antagonisms between cations and anions present in the reaction medium.     

Development of ectomycorrhizas in model beech–spruce ecosystems on siliceous and calcareous soil: a 4-year experiment with atmospheric CO2 enrichment and nitrogen fertilization

To study responses of forests to global change, model ecosystems consisting of beech and spruce trees were established in open top chambers. The ecosystems were exposed to four conditions for 4 years, each replicated four times: ambient and elevated CO₂, and low and high nitrogen input. At the end of the trial, the trees were 6–8 years old. Each chamber contained two separate compartments with siliceous and calcareous soil. Here, we focus on the development of ectomycorrhizas in the topsoil layer. Ectomycorrhizal fungal biomass associated with the fine roots, estimated by using ergosterol content as a marker, was much higher in the siliceous compared to the richer calcareous soil. Also, in root-free soil samples, the level of ergosterol, indicative of the extraradical mycelium of ectomycorrhizal fungi as well as the mycelium of other fungi, was about six times higher in the siliceous than in the calcareous soil. Conditions of elevated atmospheric CO₂ primarily affected ectomycorrhizas in the calcareous soil. Fungal biomass, calculated per soil volume of the top soil layer, increased significantly, as did the metabolic activity of the ectomycorrhizal fungi, measured as uptake of glucose and synthesis of trehalose. Conditions of nitrogen fertilization affected ectomycorrhizas in the siliceous, nutrient poor soil.