Atmospheric CO2 as a Global Change Driver Influencing Plant-Animal Interactions

Plants respond to changes in atmospheric carbon dioxide. Toherbivores, the decreased leaf protein contents and increasedC/N ratios common to all leaves under elevated atmospheric carbondioxide imply a reduction in food quality. In addition to thesefine-scale adjustments, the abundance of C₃ and C₄ plants (particularlygrasses) are affected by atmospheric carbon dioxide. C₄ grassescurrently predominate over C₃ grasses in warmer climates andtheir distributions expand as atmospheric carbon dioxide levelsdecreased during glacial periods. C₄ grasses are a less nutritiousfood resource than C₃ grasses both in terms of reduced proteincontent and increased C/N ratios. There is an indication thatas C₄-dominated ecosystems expanded 6–8 Ma b.p., therewere significant species-level changes in mammalian grazers.Today there is evidence that mammalian herbivores differ intheir preference for C₃ versus C₄ food resources, although thefactors contributing to these patterns are not clear. Elevatedcarbon dioxide levels will likely alter food quality to grazersboth in terms of fine-scale (protein content, C/N ratio) andcoarse-scale (C₃ versus C₄) changes.     

The Influence of Plant Nitrogen Status on NO2 Uptake, NO2 Assimilation and on the Gas Exchange Characteristics of Barley Plants Exposed to Atmospheric NO2

Barley plants were grown in nutrient solution at two contrastingnitrate concentrations to produce plants of low or high nitrogen(N) status. Leaves were then exposed continuously to either0.3 mm³ dm^–3 NO₂ or clean air, with the roots and rootingmedium isolated from the polluted air. Uptake of NO₂ was measuredin two ways; as depletion from an air stream containing thegas and using ¹⁵N-labelled NO₂. Results from the two methodsagreed well and demonstrated that the flux of NO₂ into the leavesof N-deficient barley was lower than that of N-sufficient plants.Nevertheless, the relative contribution of¹⁵N derived from ¹⁵NO₂to the N status of the plant was greater in the plants suppliedwith low nitrate. A major factor in regulating NO₂ uptake bybarley leaves appeared to be stomatal conductance, althoughinternal conductance may also be involved. The effects of NO₂exposure of barley on carbon dioxide exchange rates, transpirationand water vapour conductance were also influenced by the N statusof the plant. Key words: Hordeum vulgare, ¹⁵N-labelled NO₂, carbon dioxide exchange, transpiration     

Change in Nitrate-Reducing Activity in Squash Seedlings with NO2 Fumigation

Nitrogen dioxide (NO₂) fumigation inhibited nitrate reductase(NR, EC 1.6.6.1 [EC] ) activity assayed by an in vivo system in thecotyledons, but not in the first leaves, of squash (Cucurbitamaxima Duch.) seedlings. The inhibition was recovered when theseedlings were transferred to NO₂-free conditions, indicatingthat the effect of NO₂ was reversible. The NADH content in thecotyledons, photosynthetic O₂ evolution and respiratory O₂ uptakedid not change notably under NO₂ fumigation. Nitrate contentsin the cotyledons and first leaves did not change with NO₂ fumigation,but nitrite, ammonium and rapidly-metabolized amino acids contentsincreased. The inhibitory effect of NO₂ was also observed inthe in vitro assay, though the inhibition rate was smaller thanthat in the in vivo assay. These results indicate that the inhibitoryeffect of NO₂ on NR activity in squash cotyledons was derivedin part from the decrease in the amount of active NR due toammonium and/or amino acids accumulated in the tissue underNO₂ fumigation. (Received February 12, 1985; Accepted May 27, 1985)     

Measurement of the rate of CO2 solubilization in leaf tissue 4

A novel method for the measurement of the rate of CO₂ exchangecaused by solubilization and bicarbonate formation in leavesis presented. The method is based on the measurement of changesin O² concentration, caused by the uptake or evolution of CO₂from the gas flow as a result of solubilization in or desolubilizationfrom leaf tissue, by means of a zirconium-oxide O₂ analyser.The advantages of the new method are a fast response and a lackof interference by the CO₂ contained in the leaf chamber andleaf intercellular spaces. The method can be used for the investigationof the kinetics of CO₂ transport from leaf intercellular spacesto chloroplasts and the role of carbonic anhydrase in this transportand the investigation of mechanistic and ecological aspectsof buffering, proton translocation and other processes controllingpH in chloroplasts of intact leaves. Key words: Leaf, carbon dioxide, pH.     

The Labelling of Glucose-6-phosphate in Pea Slices Supplied with 14C Glucose, and the Assessment of the Respiratory Pathways in Air, in 10% CO2 and in 2% O2 in N

A method is proposed for estimating the proportion of CO₂ ofrespiration derived from the PPP and the TCAC by measuring thespecific activity of the respired CO₂ and those of its precursorsin the tissue. In practice the specific activity of C-1 of G-6-Pand of the carboxyl carbon atoms of pyruvate and malate havebeen taken as the precursors. To apply this method there mustbe good evidence that these compounds are not in ‘pools’in the cell. In air in the sliced pea cotyledon 25–40% of the respiredCO₂ arose from the PPP. In both 10% CO₂ in air and in 2% O₂ in N there was a showingin the rate of respiration and a large increase in the proportionof the CO₂ deriving from the PPP. It is argued that in eachcase the cause of this was different. Key words: Carbon dioxide, Respiration, Pisum Sativum, Pentose-phosphate pathway     

CO2 chemosensitivity in Helix aspersa: three potassium currents mediate pH-sensitive neuronal spike timing

Elevated levels of carbon dioxide increase lung ventilation in Helix aspersa. The hypercapnic response originates from a discrete respiratory chemosensory region in the dorsal subesophageal ganglia that contains CO₂-sensitive neurons. We tested the hypothesis that pH-dependent inhibition of potassium channels in neurons in this region mediated the chemosensory response to CO₂. Cells isolated from the dorsal subesophageal ganglia retained CO₂ chemosensitivity and exhibited membrane depolarization and/or an increase in input resistance during an acid challenge. Isolated somata expressed two voltage-dependent potassium channels, an A-type and a delayed-rectifier-type channel (I_KA and I_KDR). Both conductances were inhibited during hypercapnia. The pattern of voltage dependence indicated that I_KA was affected by extracellular or intracellular pH, but the activity of I_KDR was modulated by extracellular pH only. Application of inhibitors of either channel mimicked many of the effects of acidification in isolated cells and neurons in situ. We also detected evidence of a pH-sensitive calcium-activated potassium channel (I_KCa) in neurons in situ. The results of these studies support the hypothesis that I_KA initiates the chemosensory response, and I_KDR and I_KCa prolong the period of activation of CO₂-sensitive neurons. Thus multiple potassium channels are inhibited by acidosis, and the combined effect of pH-dependent inhibition of these channels enhances neuronal excitability and mediates CO₂ chemosensory responses in H. aspersa. We did not find a single "chemosensory channel," and the chemosensitive channels that we did find were not unique in any way that we could detect. The protein "machinery" of CO₂ chemosensitivity is probably widespread among neurons, and the selection process whereby a neuron acts or does not act as a respiratory CO₂ chemosensor probably depends on the resting membrane potential and synaptic connectivity. carbon dioxide     

Effects of Atmospheric NO2 on Azolla-Anabaena Symbiosis

Cultures of the water fern Azolla pinnata R, Br. exposed for1 week to atmospheric NO₂ (50, 100 or 200 nl l^-1) induced additionallevels of nitrate reductase (NaR) protein and nitrite reductase(NiR) activity. At low concentrations of NO₂ (50 nl l^-1), nitratederived from NO₂ provides an alternative N source for Azollabut does not affect rates of acetylene reduction. However, thesymbiotic relationship between Azolla and its endosymbiont,Anabaena azollae is only affected adversely by high concentrations(100 and 200 nl l^-1) of atmospheric NO₂. The resultant decreasesin rate of growth, nitrogen fixation, heterocyst formation,and overall nitrogen cycling are probably due to the additionalaccumulation of N products derived from higher levels of atmosphericNO₂. Parallel increases in levels of polyamines suggest thatAzolla partially alleviates these harmful effects by incorporatingsome of the extra NO₂-induced N into polyamines.Copyright 1994,1999 Academic Press Azolla-Anabaena symbiosis, nitrogen dioxide pollution, nitrogen metabolism, polyamines     

Temperature Effects on Rice at Elevated CO2 Concentration

The continuing increase in atmospheric carbon dioxide concentration([CO₂]) and projections of possible future increases in globalairtemperatures have stimulated interest in the effects of theseclimate variables on agriculturally important food crops. Thisstudywas conducted to determine the effects of [CO₂] and temperatureon rice (Oryza sativa L., cv. IR–30). Rice plants weregrownseason-long in outdoor, naturally sunlit, controlled-environment,plant growth chambers in temperature regimes ranging from 25/18/21°Cto 37/30/34°C (daytime dry bulb air temperature/night-timedry bulb air temperature/paddy water temperature)and [CO₂] of660 µmol CO₂ mol1 air. An ambient chamber was maintainedat a [CO₂] of 330 µmol mol^–1 and temperature regimesof 28/21/25°C. Carbon dioxide enrichment at 28/21/25°Cincreased both biomass accumulation and tillering and increasedgrain yield by 60%. In the 660 µmol mol^–1 [CO₂]treatment, grain yield decreased from 10.4 to 1.0 Mg ha^–1with increasing temperature from 28/21/25°C to the 37/30/34°Ctemperature treatment. Across this temperature range, the numberof panicles plant^–1 nearly doubled while the number ofseeds panicle^–1 declined sharply. These results indicatethat while future increases in atmospheric [CO₂] are likelyto be beneficial to rice growth and yield, potentially largenegative effects on rice yield are possible if air temperaturesalso rise. Key words: Oryza sativa, CO₂, temperature, growth, yield     

CO2 Excretion Across Isolated Perfused Crab Gills: Facilitation by Carbonic Anhydrase

Crab gill carbonic anhydrase is shown to facilitate the excretionof carbondioxide across isolated perfused gills. A techniquefor perfusing crab gills and assessing the metabolic viabilityof perfused gills is also described in detail. The techniqueis used to follow the disappearance of ¹⁴C label as HCO₃^–and CO₂ from internal perfusate passing through the gill. Theexcretion of the label increases with the flow rate of the externalperfusate across the outside of the gills. The addition of carbonican hydrase to the internal perfusate results in a two- to fourfoldincrease in the excretion of label while Diamox (acetazolamide)treatment decreases the excretion of label by half. It is alsosuggested that carbonic anhydrase, present in muscle tissuesof crabs, minimizes the disequilibrium of the hemolymph CO₂system as metabolically produced CO₂ leaves the tissues andenters the hemolymph. Parallels are drawn between the presenceof carbonic anhydrase in the crab gill system and the presenceof this enzyme in the respiratory organs of both aquatic andterrestrial animals.     

Specific inhibition of photosystem II activity in chloroplasts by fumigation of spinach leaves with SO2

The phytotoxic effects of sulfur dioxide (SO₂) were investigatedby fumigating spinach plants with SO₂. Inhibition of 2,6-dichloroindophenol(DCIP) photoreduction was observed in spinach chloroplasts isolatedfrom fumigated leaves. NADP and DCIP photoreductions were inhibitedto a similar extent by fumigation with 2.0 ppm SO₂ but electronflow from reduced DCIP to NADP was not affected. When electronflow from H₂O to NADP was inhibited by 36%, a 39% inhibitionof non-cyclic photophosphorylation was observed. However, phenazinemethosulfate(PMS)-catalyzed cyclic photophosphorylation wasas active as in the control chloroplasts. Moreover, in the presenceof PMS, no significant suppression was observed in the extentof light-induced H⁺ uptake or in the rate of H⁺ efflux in chloroplasts.From these results, it can be concluded that SO₂ inhibits theelectron flow driven by photosystem II when plants have beenfumigated with SO₂. In spinach leaves fumigated with SO₂, the rate of photosyntheticO₂ evolution was reduced under light-limited conditions, whilethe rate of respiratory O₂ uptake changed slightly. (Received February 8, 1979; )     

A Dynamic Model for Photosynthesis

A dynamic mathematical model of the effect of radiant flux densityand CO₂ concentration on the rate of photosynthesis is proposed.An appropriate dynamic experimental method for ecological studiesof this subject is described. The methodology permits the analysisof numerous problems, including the effect of changes in CO₂concentration on photosynthesis and the effectiveness of energyconversion by a leaf of a plant in different environmental conditions. The dynamic model for photosynthesis is composed of two separateinteracting non-linear parts; one describes the dynamics ofthe complex set of light reactions, and the other describesthe dark reactions. The model explains the dynamics of leafphotosynthesis in a closed circuit flow system, and also explainsthe expressions for the equilibrium states of photosyntheticrate widely used in the literature. photosynthesis, mathematical model, carbon dioxide fixation, light reactions     

The Growth and Yield Responses ofFragaria ananassato Elevated CO2and N Supply

Strawberry plants (Fragaria ananassaDuchesne var. Elsanta) weregrown in pots at two concentrations of carbon dioxide (partialpressures of 39 and 56 Pa) and with three rates of nitrogensupply (0.04, 0.4 and 4 mMas nutrient solution) to study theirindividual and interactive effects on plant growth and fruityield. Nitrogen deficiency reduced total dry biomass and relativegrowth rate (RGR), mainly through reductions in leaf area ratio(LAR) and plant N concentration (PNC), although both the netassimilation rate (NAR) and root weight ratio (RWR) increased.Elevated CO₂increased the N productivity (NP) but reduced theLAR. High CO₂increased the fruit yield by 42% at high N supplyand by 17% at low N supply. The CO₂yield enhancement occurredthrough an increase in the flower and fruit number of individualplants. This resulted in an increase in the fruit weight ratio(FWR) of plants at high CO₂. Nitrogen deficiency reduced thefruit yield by about 50% through decreases in fruit size, fruitset and the number of fruits. However, N deficiency increasedthe proportion of total plant dry biomass allocated to fruits.There were no significant interactions between CO₂and N supplyon yield.Copyright 1998 Annals of Botany Company Nitrogen; carbon dioxide; strawberry (Fragaria ananassaDuchesne); fruit yield.     

The Influence of Carbon Dioxide and Daily Photon-flux Density on Optimal Leaf Nitrogen Concentration and Root: Shoot Ratio

Using a cost-benefit model, the leaf nitrogen concentrationand root : shoot ratio that maximize whole-plant relative growthrate are determined as a function of the above-ground environment(integrated daily photon flux density and the concentrationof carbon dioxide at the site of fixation within the leaf).The major advantage of this approach is that it determines theadaptive significance of leaf physiology by considering thefunctional integration of leaves and roots. The predicted responseto increasing daily photon flux densities is an increase inoptimal leaf N concentration (N_opt) and a concomitant increasein root: shoot ratio. Increased carbon dioxide concentrations,on the other hand, reduce N_opt and only slightly change root:shoot ratio. The observed increase in leaf nitrogen concentrationfound in plants growing at high altitudes (low CO₂ partial pressure)is also predicted. Since these responses to light and CO₂ maximizethe whole-plant relative growth rate, the observed adjustmentsthat plants make to light and carbon dioxide concentration appearto be adaptive. We show that the relationship between photosynthesis and leafnitrogen concentration is complex and depends on the light andCO₂ levels at which photosynthesis is measured. The shape ofthis function is important in determining N_opt and the oppositeresponse of leaf nitrogen to light and carbon dioxide is shownto be the result of the different effects of light and CO₂ onthe photosynthesis-leaf nitrogen curve. Plant growth, photosynthesis, leaf nitrogen, biomass allocation, optimization, carbon dioxide light     

The Effect of External pH on the Apparent CO2 Affinity of Chlorella saccharophila

The carbon dioxide compensation point of the unicellular greenalga, Chloretla saccharophila, was determined in aqueous mediumby a gas chromatographic method. Compensation points decreasedmarkedly from 63 cm³ m^–3 at an external pH of 4.0 to 3.2cm³ m^–3 at pH 8.0 and were not affected by the O₂ concentrationof the medium. The calculated CO² concentration required tosupport the half-maximum photosynthetic rate of the algal cellsranged from 6.0 mmol m_–3 at an external pH of 60 to 1.5mmol m^–3 at pH 8.0 and these values were not affectedby O₂ concentration. The K_m(CO₂) of nbulose-l,5-bisphosphatecarboxylase isolated from cells grown either at pH 4.0 or pH8.0 was determined to be 64 mmol m^–3. These results indicatethat loss of CO₂ by photorespiration does not occur in C. saccharophilacells at acid pH and the disparity between the apparent affinityfor CO₂ of the intact cells and that of the carboxylase indicatesthe operation of a ‘CO₂ concentrating mechanism’in this alga at acid pH. Key words: Acidophilic alga, bicarbonate transport, Chlorella saccharophila, compensation point, CO₂ affinity, PH, RuBP carboxylase     

人类新基因netrin-G 2的初步分析

Netrins是与层粘连蛋白相关的、高度保守的小分子分泌蛋白家族成员,在细胞迁移和轴突导向活动中具有重要的作用,其同源物在多种模式动物中均已发现.Netrins分为2个亚家族:netrins和netrin-Gs;其中的netrin-G亚家族各成员之间具有高度的相似性.目前,在人类中得到确证的只有netrin-G₁,推测在人类中应该还有其它的netrin-G亚家族成员以及编码它的基因.从20周龄人胚中提取脑组织的总RNA,用PCR cDNA文库试剂盒制备脑组织cDNA文库,再用特异性引物进行PCR扩增,得到1条人netrin-G的全长cDNA,命名为人netrin-G₂,用Northern杂交研究其表达,用进化树分析其与netrin家族各成员间的关系;证实人netrin-G₂确实为netrin-G亚家族的1个新成员,该基因位于染色体的9q34,大小为2 428 bp,含有1个1 593 bp的假定开放阅读框,起始密码子为86位的甲硫氨酸,终止密码子为1 678位的TGA,可编码1个大小为530个氨基酸的蛋白质;Northern杂交显示,人netrin-G₂在脑组织中特异性表达,而在其它组织中却很少发现,推测人netrin-G₂可能在中枢神经系统的发育过程中具有重要的作用,可能与刺激性神经冲动的传递以及神经调节有关.     

A CO2 Concentrating System in Leaves of Higher C3-Plants Predicted by a Model Based on RuBP Carboxylase/Oxygenase Kinetics and 14CO2/12CO2 Exchange

Mächler, F., Lehnherr, B., Schnyder, H. and Nösberger,J. 1985. A CO₂ concentrating system in leaves of higher C₃-plantspredicted by a model based on RuBP carboxylase/oxygenase kineticsand ¹⁴CO₂/¹²CO₂ exchange.–J. exp. Bot. 36: 1542–1550. A model is presented which compares the ratio of the two activitiesof the enzyme nbulose bisphosphate carboxylase/oxygenase asdetermined in vitro with the ratio of photosynthesis to photorespirationin leaves as determined from differential ¹⁴CO₂/¹²CO₂ uptakeor from CO₂ compensation concentration. Discrepancies betweenmeasurements made in vitro and in vivo are attributed to theeffect of a CO₂ concentrating system in the leaf cells. Interferencefrom dark respiration is discussed. A CO₂ concentrating systemis postulated which is efficient mainly at low temperature andlow CO₂ concentration. Key words: —Photosynthesis, photorespiration, ribulose bisphosphate carboxylase/oxygenase     

On the Use of the Katharometer for the Estimation of Carbon Dioxide Concentrations in Soils

The method depends upon the difference in thermal conductivitybetween carbon dioxide and other components of the soil atmosphere. The circuit calibration and mode of operation of the instrumentare fully described together with an investigation of the effectof differences in ambient temperature. Values of carbon dioxide concentration obtained in trials onculture and field soils are given. The values lie within rangesreported by earlier workers using gas analysis of samples ofsoil air. katharometer, CO₂ concentration, soil atmosphere     

Acclimation of nitrogen uptake capacity of rice to elevated atmospheric CO2 concentration

Background and Aims: Nitrogen (N) is a major factor affecting yield gain of cropsunder elevated atmospheric carbon dioxide concentrations [CO₂].It is well established that elevated [CO₂] increases root mass,but there are inconsistent reports on the effects on N uptakecapacity per root mass. In the present study, it was hypothesizedthat the responses of N uptake capacity would change with theduration of exposure to elevated [CO₂]. Methods: The hypothesis was tested by measuring N uptake capacity inrice plants exposed to long-term and short-term [CO₂] treatmentsat different growth stages in plants grown under non-limitingN conditions in hydroponic culture. Seasonal changes in photosynthesisrate and transpiration rate were also measured. Key Results: In the long-term [CO₂] study, leaf photosynthetic responsesto intercellular CO₂ concentration (Ci) were not affected byelevated [CO₂] before the heading stage, but the initial slopein this response was decreased by elevated [CO₂] at the grain-fillingstage. Nitrate and ammonium uptake capacities per root dry weightwere not affected by elevated [CO₂] at panicle initiation, butthereafter they were reduced by elevated [CO₂] by 31–41% at the full heading and mid-ripening growth stages. In theshort-term study (24 h exposures), elevated [CO₂] enhanced nitrateand ammonium uptake capacities at the early vegetative growthstage, but elevated [CO₂] decreased the uptake capacities atthe mid-reproductive stage. Conclusions: This study showed that N uptake capacity was downregulated underlong-term exposure to elevated [CO₂] and its response to elevated[CO₂] varied greatly with growth stage.     

PCO2 method for measuring photosynthesis and respiration in freshwater lakes

A method for measuring net community photosynthesis and respirationbased on changes in PCO₂ is described. Samples were incubatedin serum stoppered bottles and successive pCO₂ measurementstaken from the bottle headspace. At the end of the incubation,samples were acidified with concentrated phosphoric acid andtotal dissolved inorganic carbon (DIC) was determined. Alkalinitywas calculated from PCO₂ and DIC. Alkalinity was assumed toremain constant for the duration of the experiment so that DICcould be calculated from PCO₂ and alkalinity for each measurement.Uptake rates were calculated from the slope of     

Light-Dependent CO2 Retrieval in Immature Barley Caryopses

Immature detached caryopses from barley (Hordeum vulgare L.var. distichum cv. Midas) were shown to be capable of light-dependentretrieval of internally-produced CO₂. In the first set of experiments,caryopses were radioactively labelled by supplying (U-14C)-sucroseto detached ears in liquid culture. Caryopses were then removedfrom the ear and given a 12 h chase of non-radioactive sucrosein either the light or dark. More ¹⁴C was recovered in the caryopsesafter the chase in the light than in the dark but the differenceswere not significant. In the second set of experiments, ¹⁴C-labelledcaryopses obtained by a 15 min light incubation in ¹⁴CO₂ weremaintained in either the light or dark for 3 h and any redistributionof label between the tissues recorded. The results show thatunder these conditions, photosynthesis in the Chl-containinggreen layer of the pericarp can prevent losses of internally-producedCO₂, since 3 times as much radiocarbon remained in the caryopsesincubated in the light as in the dark. These differences weresignificant at P=0.001. Experiments with the mutant barley Albinolemma, which has no Chi in the pericarp, showed that there waslittle difference between light and dark treatments. This confirmsthe suggestion that photosynthesis in the pericarp of the normalcultivar Midas may be concerned in the refixation of CO₂. Key words: Barley, pericarp, photosynthesis, carbon dioxide