Regulation of bacterial secondary production in two eutrophic lakes and in experimental enclosures

Bacterial cell number and secondary production of bacterioplanktonwere measured during two periods of 14 and 19 days, respectively,in experimental enclosures of lake water (each containing about5–7 m³ and in two Danish, eutrophic lakes. Additions ofphosphate and planktivorous fish to some of the enclosures inducedrapid changes in the phytoplankton biomass, primary productionand release of extracellular organic carbon (EOC). In some ofthe enclosures containing fish, release of EOC from phytoplanktoncon stituted from 36 to 55% of the bacterial secondary productiondetermined by means of [³H]thymidine incorporation into DNA.In others, such a relationship was not found, suggesting thatphytoplankton species-specific release rates of compositionof EOC might play a role in the microbial response. In enclosureswithout fish, released dissolved organic carbon from zooplanktongrazing activities could Sustain     

14C-labelling patterns of phytoplankton: specific activity of different product pools

Primary production, release of extracellular products (EOC)and the distribution of ¹⁴C into primary products of photosynthesis(low mol. wt. products, lipids, polysaccarides, and proteins)were investigated during 2 months in spring 1983 in eutrophicLake Hylke, Denmark. After half days in situ incubations, eachfraction was assayed for ¹⁴C activity and the standing stockof carbon. Calculation of specific activity revealed non-equilibriumconditions in all fractions. Since the intracellular fractionsact as precursors of EOC, this means that the release may beseriously underestimated (3–20 times) by assuming isotopicequilibrium between the inorganic carbon pool and the EOC productsin the Extracellular medium. However, measurements of the standingstock of carbon are complicated by inclusion of non-phytoplanktonmaterial, thereby making calculation of specific activity uncertain. Present address: Carbon 14 Centralen, The International Agencyfor ¹⁴C Determination, 11, Agern Allé, DK-2970 Hørsholm,Denmark     

The effect of environmental variables on release of extracellular organic carbon by freshwater macrophytes

The¹⁴C-technique was used to investigate the effect of oxygen, inorganic carbon and light intensity on the release of extracellular organic carbon (EOC) by three submerged macrophytes. At low CO₂ availability and low light intensities the release expressed as a percentage of the total carbon fixation was higher than at more optimal photosynthetic conditions. Oxygen did not influence the release of EOC. As the rates of release generally followed the rates of photosynthesis, it was concluded that release is closely coupled to photosynthesis. Using gel-filtration to separate EOC it was shown that the tested environmental variables did not influence the molecular weight distribution. The release of EOC was low in all species (0.1 to 4% of the total carbon fixation). Due to the low values it is suggested that release of EOC is of no quantitative importance to the carbon budget of the plants.     

Diurnal variations in the contributions of autotrophic picoalgae and heterotrophic bacteria to planktonic production in an upland lake

An investigation of the diurnal variation in contributions toproduction of the autotrophic and heterotrophic components ofthe picoplankton community was carried out during August andSeptember in Llyn Padarn, a mesotrophic upland lake in NorthWales. The picoplankton was separated using 1 µm pore-sizedfilters into the smaller cell sized fraction (<1 µm),the majority of the bacteria and the larger cell sized picoalgae(<3>1 µm), together with some bacteria. The distributionof bacterial heterotrophic activity between these two fractionsof picoplankton was assessed by uptake of [¹⁴C]glucose and differentialfiltration. Thus, the absolute autotrophic production by picoalgaeand the heterotrophic contribution by bacteria to picoplanktoncommunity production via uptake of extracellular organic carbon(EOC) were determined. Rates of picoplankton community productionexhibited diurnal variation with maximum rates of 19.1 mg Cm^–3 h^–1 recorded at 18.00 h at 4 m depth in September.The bacterial contribution to picoplankton community productionincreased markedly between 15.00 and 18.00 h. Rates of absoluteautotrophic production varied less over 24 h than rates of accumulationin bacteria of ¹⁴C-labelled EOC released from the entire phytoplanktoncommunity. Bacteria contributed up to 86–98% of the neworganic carbon within the picoplankton community at the endof the day. The maximum rate of absolute autotrophic productionin the picoplankton was 1.6 mg C m^–3 h^–1 at 18.00h at 1 m in August, and the maximum rate of bacterial accumulationof new organic carbon was 18.5 mg C m^–3 h^–1 at 18.00h in September at 4 m depth. The diurnal pattern of picoplanktoncommunity production involved increasing rates during the daywith a maximum at 18.00 h. Autotrophic processes were dominantin the first 3–6 h of the light cycle and heterotrophicuptake of ¹⁴C-labelled EOC was the major component from 15.00h onwards. Bacterial uptake of newly released EOC by phytoplanktonwas rapid, comprised the majority of picoplankton production,particularly later in the day, and contributed a maximum of60% of the total pariculate primary production in plankton between15.00 and 18.00 h at 4 m in September with a mean contributionof between 6 and 26% over 24 h in these investigations. Theimportance of autotrophic processes in picoplankton communityproduction has been overestimated in previous investigations.Bacteria play a major role in transferring newly produced EOCrapidly from phytoplankton to the picoplankton community. Atthe end of the day, the majority of newly produced organic carbonis in bacterial cells and this production is significant inthe dynamics of carbon production within the entire planktoniccommunity.     

Summer nutrient dynamics in the Middle Atlantic Bight: primary production and utilization of phytoplankton carbon

In late summer, production and utilization of carbon in thestratified water of the Middle Atlantic Bight appears to approachsteady-state conditions. In the euphotic zone there is a paniculateorganic carbon (POC) pool of {small tilde}6000 mg Cm^–2.Primary production adds {small tilde}350 mg C m^–2 d^–1,while zooplankton ingestion removes {small tilde}450 mg C m^–2d^–1, of which {small tilde}135 mg C m^–2 d^–1are returned to the POC pool as feces. Sinking of POC averages{small tilde}240 mg C m ^–2 d^–1. Thus, there is anet loss from the euphotic zone of {small tilde}200 mg C m^–2d^–1, which represents a removal rate from the POC poolof {small tilde}3%d^–1. However, sinking losses of phytoplanktoncarbon from the POC pool were small ({small tilde}12 mg C m^–2d^–1), which suggests that most of the primary productionenters the pelagic food chain. This is in sharp contrast tothe conditions in early spring, when the POC pool increasessubstantially, and the main loss is sinking rather than heterotrophicconsumption. ^*This research was supported by the U.S. Department of Energyunder Contract No. DE-AC02-76CH000I6.     

The use of antibiotics to reduce bacterioplankton uptake of phytoplankton extracellular organic carbon (EOC) in the Potomac River estuary

Phytoplankton production and accumulation of extracellular organic carbon (EOC) was tracked during diel intervals in microcosms by inhibiting bacterioplankton assimilation of EOC with streptomycin and kanamycin. Bacterioplankton production (³H-thymidine incorporation) and metabolism (¹⁴C-glucose incorporation) were monitored in samples collected from the Potomac River estuary to determine the effect of the antibiotics. Particulate (i.e., raw water) primary production and EOC (i.e., water passing through 1.0 μm glass fiber filter) production rates were monitored to determine the impact of antibiotics on phytoplankton. In preliminary experiments, neither streptomycin nor kanamycin alone significantly inhibited bacterioplankton activity compared to controls, but when both were present secondary production and metabolism were reduced up to 90%, and remained as such for 45 h. During field evaluations using a streptomycin and kanamycin mixture (50 μM each) particulate primary production and EOC production were not statistically different in control and antibiotic treated samples indicating that the antibiotics did not negatively influence phytoplankton production rates. In the presence of antibiotics dissolved free amino acids (DFAA) and, to a lesser extent, monosaccharides were significantly elevated compared to controls. This study demonstrates that streptomycin and kanamycin are capable of inhibiting bacterioplankton metabolism and uptake of dissolved organic carbon (DOC) in the samples tested so that the contribution of EOC to the DOC pool and to bacterioplankton metabolism could be measured and assessed.     

Release of extracellular organic carbon during a diatom bloom in Lake Mossö: molecular weight fractionation

SUMMARY. Carbon fixation and release of extracellular organic carbon (EOC) during a diatom bloom were measured by the ¹⁴C—technique. Release in light was always a small fraction (1.5-6.0%) of total carbon fixation. Maximum values were found during the stationary phase of growth. A larger (15–50%) fraction of the inorganic carbon assimilated in the dark was released, although the absolute values were lower than in the light.
Gel permeation chromatography showed that low molecular weight products (<700 daltons) dominated EOC released in the light. A shift toward more complex molecules (> 10,000 daltons) was found in the stationary phase. EOC released in the dark had almost equal amounts of low (< 700 daltons), intermediate (˜5,000 daltons) and high (< 10,000 daltons) molecular weight products.
It is shown that with time the molecular weight composition of EOC shifted towards a predominance of larger molecules.     

西南喀斯特石漠化环境下土地利用变化对土壤有机碳及其组分的影响

土壤有机碳是喀斯特生态系统中碳转移的动力学媒介和碳流通的主要途径,土壤有机碳及其组分是土壤碳循环的重要组成部分,然而目前缺乏关于喀斯特地区土壤有机碳及其组分的研究。本研究以西南典型喀斯特石漠化区——贵州关岭花江6种典型土地利用方式下(花椒林、火龙果林、花椒火龙果混交林、圆柏林、圆柏女贞混交林和坡耕地)的土壤为研究对象,分析土地利用方式变化对土壤有机碳含量(SOC)和储量(SOCS)、土壤水溶性有机碳(WSOC)、易氧化有机碳(EOC)、颗粒有机碳(POC)、轻组有机碳(LFOC)及重组有机碳(HFOC)含量及其分配比例的影响。结果表明: 6种土地利用方式下SOC和SOCS均表现为圆柏林、圆柏女贞混交林和花椒林显著大于火龙果林、花椒火龙果混交林和坡耕地;在0～20 cm土层,土壤SOCS平均值为花椒林＞圆柏林＞圆柏女贞混交林＞坡耕地＞花椒火龙果混交林＞火龙果林。土壤WSOC、EOC、POC、LFOC和HFOC含量均表现为圆柏林、圆柏女贞混交林和花椒林大于其他3种土地类型。土壤SOC与其各组分(WSOC、EOC、POC、LFOC和HFOC)均呈显著正相关,且各组分两两之间也呈显著正相关。花椒林可作为中国西南喀斯特石漠化生态恢复和山地农业发展优先考虑的经济物种。土壤WSOC、EOC、POC、LFOC和HFOC可作为反映土壤有机碳库的有效指标。     

川西沿海拔梯度典型植被类型土壤活性有机碳分布

研究土壤活性有机碳含量及分配比例是揭示土壤碳库周转及调控机理的重要途径,通过利用高锰酸钾氧化法获得易氧化有机碳、湿筛法获得颗粒有机碳和密度分离法获得轻组有机碳3项指标探讨沿海拔梯度不同植被类型间(山地常绿阔叶林、常绿落叶阔叶混交林、落叶阔叶林、针阔混交林、暗针叶林)土壤活性有机碳含量差异及调控因子,结果表明:随土层加深,土壤颗粒和轻组有机碳含量及分配比例均降低,土壤易氧化有机碳含量降低而分配比例保持较稳定水平。高海拔植被类型具有较高的土壤活性有机碳含量和分配比例。不同活性有机碳含量之间均呈显著线性相关(P0.05)表明活性有机碳起源的类似。活性有机碳与土壤粘粒+粉粒含量百分比呈显著负相关(P0.05)表明活性有机碳趋向分布于土壤大团聚体当中。年均温与不同植被类型间表层土壤活性有机碳含量和分配比例成负相关趋势,但可能由于取样点较少的缘故而在统计上不显著。年均温与土壤非保护性有机碳向保护性有机碳的转化速率常数(K)接近于显著负相关(P=0.062)。     

Environmental Factors Affecting the Loss of Carbon from the Roots of Wheat and Barley Seedlings

The amounts of carbon released into soil from roots of wheatand barley seedlings grown under three environmental conditionsfor 3 weeks with shoots in constant specific activity ¹⁴CO₂are reported. This carbon loss was measured as respired ¹⁴CO₂from both the root and the accompanying microbial populationand as root derived ¹⁴C-labelled organic C compounds in thesoil. With a 16 h photoperiod, growth at 15 ?C constant or 18?C day/14 ?C night gave a loss of 33–40% of the totalnet fixed carbon (defined as ¹⁴C retained in the plant plus¹⁴C lost from the root). The proportion of ¹⁴C translocatedto the roots that was released into the soil did not changewith temperature, so carbon distribution within the plant musthave changed. With a 12 h photoperiod and a temperature regimeof 18 ?C/14 ?C carbon loss from the roots was decreased to 17–25%of the total fixed carbon. Key words: Cereals, Roots, Carbon loss     

黄土高原4种典型植被对土壤活性有机碳及土壤碳库的影响

对黄土高原丘陵沟壑区4种典型植被(荒草地、文冠果林地、柠条灌丛、沙棘林地)进行了活性有机碳(MBC,Microbial Biomass Carbon;EOC,Easily Oxidated Carbon;POC,Particle Organic Carbon)及有机碳的测定,结果表明:4种植被土壤活性有机碳含量随着土层深度的增加呈逐渐降低的趋势,土壤有机碳为文冠果林地荒草地沙棘林地柠条灌丛,且差异显著(P0.05)。荒草地0—40 cm范围内土壤MBC含量分别比文冠果林地、柠条灌丛、沙棘林地显著降低了8.61%、23.84%、41.42%(P0.05);荒草地土壤POC含量分别比文冠果林地、沙棘林地降低了14.47%、16.67%,比柠条灌丛POC高出了25.00%;但荒草地土壤EOC含量、碳库活度及碳库管理指数均显著高于其他3种植被类型(P0.05)。4中植被类型中沙棘林地土壤微生物熵(SME,Soil Microbial Entropy)最大,而柠条灌丛土壤有机碳储量最小。土壤SOC与土壤EOC、有机碳储量(SOCS,Soil Organic Carbon Storage)显著相关(P0.05),而土壤SOC与POC呈极显著的相关(P0.01);土壤EOC与土壤POC显著相关(P0.05),但土壤MBC与SOC、POC、EOC、SOCS均呈现出不同程度的负相关性。因此土壤活性有机碳能客观反映土壤肥力和土壤质量的变化情况,是描述土壤质量和评价土壤管理的重要指标。     

Pelagic food web processes in an oligotrophic lake

Major pelagic carbon pathways, including primary production, release of extracellular products (EOC), bacterial production and zooplankton grazing were measured in oligotrophic Lake Almind (Denmark) and in enclosures (7 m³) subjected to artificial eutrophication. Simultaneous measurements at three days interval of carbon exchange rates and pools allowed the construction of carbon flow scenarios over a nineteen day experimental period.The flow of organic carbon was dominated by phytoplankton EOC release, which amounted from 44 to 58% of the net fixation of inorganic carbon. Gross bacterial production accounted for 33 to 75% of the primary production. The lower values of EOC release (44%) and bacterial production (33%) were found in the enclosures with added nutrients. The release of recently fixed photosynthetic products was the most important source of organic carbon to the bacterioplankton. Uptake of dissolved free amino acids was responsible for 52 to 62% of the gross bacterial production. Thus, amino acids constituted a significant proportion of the EOC. Zooplankton (< 50 µm) grazing on algae and bacteria accounted only for a minor proportion of the particulate production in May. Circumstantial evidence is presented that suggests the chrysophycean alga Dinobryon was the most important bacterial remover.The results clearly demonstrated EOC release and bacterial metabolism to be key processes in pelagic carbon cycling in this oligotrophic lake.     

Biogeochemistry of platelet ice: its influence on particle flux under fast ice in the Weddell Sea, Antarctica

An array of four sediment traps and one current meter was deployed under a well-developed platelet layer for 15 days in the Drescher Inlet in the Riiser Larsen ice shelf, in February 1998. Traps were deployed at 10 m (just under the platelet layer), 112 m (above the thermocline), 230 m (below thermocline) and 360 m (close to sea floor). There was a substantial flux of particulate organic material out of the platelet layer, although higher amounts were collected in the traps either side of the thermocline. Material collected was predominantly composed of faecal pellets containing diatom species growing within the platelet layer. The size classes of these pellets suggest they derive from protists grazing rather than from larger metazoans. Sediment trap material was analysed for particulate organic carbon/nitrogen/phosphorus (POC/PON/POP) and '¹³C_POC (carbon isotopic composition of POC). These were compared with organic matter in the overlying platelet layer and the water column. In turn, the biogeochemistry of the platelet layer and water column was investigated and the organic matter characteristics related to inorganic nutrients (nitrate, nitrite, ammonium, silicate, phosphate), dissolved organic carbon/nitrogen (DOC/DON), pH, dissolved inorganic carbon (DIC), oxygen and '¹³C_DIC (carbon isotopic composition dissolved inorganic carbon).     

Short-Term Fixation of 14Carbon by the Submerged Aquatic Angiosperm Potamogeton pectinatus

In ¹⁴C fixation experiments, 3-phosphoglyceric acid was thefirst product of carbon assimilation in the light in Potamogetonpectinatus. The pattern of early ¹⁴C-labelled compounds wasthe same over a range of pH values of the medium from 3.5 to8.1. Rates of ¹⁴C incorporation declined with increasing pHof the medium indicating that free CO₂ is the major exogenoussource of carbon for photosynthesis in Potamogeton pectinatus.     

Organic matter release by dominant hermatypic corals of the Northern Red Sea

Particulate organic matter (POM) and dissolved organic carbon (DOC) release by six dominant hermatypic coral genera (Acropora, Fungia, Goniastrea, Millepora, Pocillopora and Stylophora) were measured under undisturbed conditions by laboratory incubations during four seasonal expeditions to the Northern Red Sea. In addition, the influence of environmental factors (water temperature, light availability and ambient inorganic nutrient concentrations) was evaluated. Particulate organic carbon (POC) and particulate nitrogen (PN) release were always detectable and genus-specific, with Stylophora releasing most POM (6.5 mg POC and 0.5 mg PN m⁻² coral surface area h⁻¹) during all seasons. The fire coral Millepora released significantly less POM (0.3 mg POC and 0.04 mg PN m⁻² coral surface area h⁻¹) than all investigated anthozoan genera. The average POC:PN ratio of POM released by all coral genera was 12 ± 1, indicating high carbon/low nitrogen content of coral-derived organic matter. POM release showed little seasonal variation, but average values of POC and PN release rates correlated with water temperature, light availability and ambient nitrate concentrations. DOC net release and elevated DOC:POC ratios were detectable for Acropora, Goniastrea and Millepora, revealing maximum values for Acropora (30.7 mg DOC m⁻² coral surface area h⁻¹), whilst predominant DOC uptake was observed for Pocillopora, Fungia and Stylophora. Depth-mediated light availability influenced DOC fluxes of Acropora and Fungia, while fluctuations in water temperature and ambient inorganic nutrient concentrations showed no correlation. These comprehensive data provide an important basis for the understanding of coral reef organic matter dynamics and relevant environmental factors.     

围栏禁牧对滇西北高寒湿地土壤活性有机碳的影响

过度放牧导致滇西北高寒湿地碳汇功能逐渐丧失,围栏禁牧作为一种有效的湿地恢复方式,其对滇西北高寒湿地土壤碳库的影响尚缺乏研究。为探明围栏禁牧对滇西北高寒湿地土壤活性有机碳的影响,以纳帕海湿地不同禁牧年限(未禁牧、禁牧3年、禁牧8年、禁牧10年)的草甸和沼泽化草甸作为研究对象,对比分析不同禁牧年限草甸和沼泽化草甸土壤总有机碳(TOC)、颗粒有机碳(POC)、溶解性有机碳(DOC)、易氧化有机碳(EOC)和微生物生物量碳(MBC)含量特征。研究结果表明,随着禁牧年限的增加(<10 a),草甸和沼泽化草甸土壤TOC、DOC、EOC、MBC含量显著增加(P<0.05);随着土壤深度的增加,草甸和沼泽化草甸土壤TOC、EOC和MBC含量逐渐下降,而POC和DOC含量逐渐增加。土壤理化性质显著影响土壤活性有机碳各组分含量,其中总氮(TN)是影响草甸和沼泽化草甸土壤活性有机碳各组分含量的主导因素。围栏禁牧有利于滇西北高寒湿地土壤质量和固碳能力的恢复,且随着禁牧年限的增加,恢复效果越明显。相同恢复年限,纳帕海沼泽化草甸土壤恢复效果比草甸土壤显著。研究可以为放牧湿地恢复研究提供理论基础。     

The Flux of 14C-Labelled Photosynthate through Soyabean Root Nodules during N2 Fixation

Experiments are described which examine the flux of photosyntheticassimilates from leaves to nodules of soyabean during N₂ fixation.The first part, where the respiratory efflux of ¹⁴CO₂ by noduleswas used as a means of assessing the import of labelled photosynthatefrom leaves, shows that most ¹⁴CO₂ loss from nodulated rootsis due to the metabolic activity of nodules. Much less photosynthatewas imported by nodules if the metabolic activity associatedwith N₂ fixation was inhibited by low O₂ concentration. The second part describes the chemical fate of current photosynthateas it is utilized by nodules. Labelled material was detectedin nodules within c.15 min of supplying ¹⁴CO₂ to the leaf. Thisrose to a maximum at c.70 min before declining by 85% withinthe following 4 h. Most (80%) ¹⁴carbon imported by nodules waseither lost as respiratory ¹⁴CO₂ or re-exported as productsof N₂ fixation. Ten per cent of imported carbon was found asstructural material and 10% as starch. Of the ¹⁴C soluble in ethanol, most was found in the neutralfraction (80% declining to 50% as sucrose) with smaller amountsas amino acids, organic acids (each category rising from 10%to 20%) and phosphate esters (<5%). Comparison of the distribution of ¹⁴C among amino acids, amidesand ureides in the nodules with that of xylem exudates indicatedthat selected compounds were exported from nodules. The ¹⁴Cdata indicate that c.80% of the nitrogen exported from noduleswas in the form of ureides (mainly allantoic acid) and only10–12% as amides. Key words: Nodules, ¹⁴C-photosynthate, Respiration, Carbon flux     

模拟氮沉降对湿地植物生物量与土壤活性碳库的影响

在两种水分条件下(W₁:非淹水,W₂:淹水)分4个氮处理(分别相当于氮沉降率0、1、3、5 g N·m^-2·a^-1)模拟了三江平原典型湿地植物湿草甸小叶章(Deyeuxia angustifolia)植株及土壤活性碳库对氮沉降的响应.结果表明：模拟氮沉降下小叶章的生物量(总生物量、地上生物量、根生物量)均高于对照,其中根生物量的增长程度最大;根中碳含量及分配比例均显著提高,而地上部位的碳含量则显著降低(P<0.05).氮沉降对土壤活性碳库具有显著影响,各活性碳库含量均以5 g N·m^-2·a^-1处理最高,氮沉降对各活性碳库的影响程度依次为CHC(碳水化合物碳)＞LBC(易氧化有机碳)＞DOC(水溶性有机碳)＞MBC(微生物量碳),氮沉降与淹水条件的耦合作用有利于活性碳的释放;回归分析表明,土壤活性碳库与小叶章相关参数间存在显著相关性.氮沉降显著提高了小叶章植株生物量及土壤的活性碳含量.     

Comparison of two methods to measure algel release of dissolved organic carbon and the subsequent uptake by bacteria

Two methods were used to compare phytoplankton release and bacterialuptake of extracellular organic carbon (EOC) products. A sizefractionation and an antibiotic method were used simultaneouslyduring several diel in Situ studies in Danish lakes and onecoastal area. Phytoplankton populations were very sensitiveto even small concentrations of antibiotics (1 µg ml^–1).Generally the bacterial activity was inhibited by 50%. In caseswhere the antibiotics did not affect phytoplankton photosynthesis,the two methods agreed reasonably. Bacterial respiration dataobtained with the antibiotic method were in the range of valuesfrom the literature. This supports the conclusion that the partialuncoupling of EOC release and uptake with antibiotics can beused to follow the flux of EOC in situ. The effect on the phytoplanktonmust however, be evaluated in each experiment. The methodologicalproblems of both approaches are discussed.     

Guard Cell Metabolism in Epidermis of Commelina communis L. During Stomatal Opening and Closing

The rates of CO² incorporation into the epidermis of C. communiswere linear and were similar during the completion of opening(2 h) and closing (1 h) movements of stomata. The kinetics of¹⁴C turnover between metabolites and the rates of ‘leakage’of metabolites were determined for opening and closing movements.When stomata were opening there was a slow turnover of ¹⁴C frommalate chiefly into sugars. Upon stomatal closure ¹⁴C was initiallymainly in sugars, malate, and sugar phosphates. Thereafter,there was a slight loss of label from sugar phosphates witha corresponding increase in malate. Starch became labelled duringopening and closing movements. Rates of incorporation of CO₂found in the ‘leakage’ fraction were greatest whenstomata were opening. Of the labelled compounds Most‘from the tissue, malate was the most highly labelled whetherstomata were opening or closing. Although interpretation of the turnover patterns is difficultwithout knowledge of pool sizes for the metabolites it is suggestedthat a pool of sugars exists within the guard cells, which havefairly direct and reversible access to carbon from starch andmalate. The implications of loss of malate from guard cellsduring stomatal opening and closing are discussed.