Pelagic Bacteria–Particle Interactions and Community-Specific Growth Rates in Four Lakes along a Trophic Gradient

Abstract The relationships between bacterial concentration, bacterial production, and cell-specific activity of both free and attached bacteria and environmental factors such as suspended solids, nutrients, and temperature were examined in four lakes, two in New Zealand and two in Switzerland. Estimates of cell-specific production were obtained by microautoradiographic counts of [³H]thymidine-labeled cells. Bacteria attached to particles accounted for only 1.3 to 11.6% of the total bacterial abundance, but showed overall 20-fold higher specific growth rates and were relatively more active than their free counterparts. On average, 80 to 100% of epibacteria were attached to organic particles. The abundance and production of free and attached bacteria were positively correlated; however, relationships between these fractions and some environmental variables differed. Cell-specific activities of active bacteria were not equivalent to mean cellular activities of the entire bacterial community and differed in their relationship to trophic state. [³H]Thymidine-positive bacteria were more tightly linked to chlorophyll a than were total bacteria. Our findings indicate that production by attached bacteria, fueled by phytoplankton carbon, supplies ``new' free bacteria to the bacterial community. Our results support the idea that particulate organic matter acts as a source of dissolved nutrients to free bacteria. Bottom-up control of bacterial biomass, as shown by regressions of biomass vs production, appeared to be stronger in two ultraoligotrophic lakes than in two more eutrophic ones. Received: 17 April 1998; Accepted: 24 August 1998     

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.     

Attached and free-living bacteria: Production and polymer hydrolysis during a diatom bloom

Abundance, production and extracellular enzymatic activity of free-living and attached bacteria were measured during the development and collapse of a spring bloom in a eutrophic lake. Free-living bacteria accounted for most of the total bacterial production during the first part of the bloom. Their production had a significant positive correlation to chlorophyll (P < .01) and polysaccharide concentration (P < .02) and to potential -glucosidase and aminopeptidase activity (P < .05), suggesting that algal release of dissolved polymeric compounds provided an important carbon source for bacterial production. As the bloom collapsed, we observed a change in the activity and structure of the microbial community. The mean contribution of attached bacteria to total bacterial production increased from 12% during the first part of the bloom to 26% at the end. Also, the extracellular enzymatic activity of attached bacteria increased as the bloom collapsed and constituted up to 75% of the total hydrolytic activity. An estimated disparity between hydrolytic activity and the corresponding carbon demand of attached bacteria suggested a net release of dissolved organic compounds from organic particles via polymer hydrolysis by attached bacteria. Correspondence to: M. Middelboe     

Photosynthetic organic carbon production and respiratory organic carbon consumption in the trophogenic layer of Lake Biwa

Measurement of the photosynthetic production rate in Lake Biwawas camed out from May 1985 to September 1987. In the light-saturatedlayer, the seasonal variation in the photosynthesis rate perchlorophyll a was regulated by water temperature. The depth-integratedphotosynthetic production rate was 0.21-1.48 g C m^–2 day^–1and the maximum value was observed in midsummer when the watertemperature of the mixed surface layer was highesL The criticalnutrient for photosynthesis may be dissolved reactive phosphorus,which was generally <1 µg P 1^–1 throughout theobservation period. In the trophogenic layer, respiratory organiccarbon consumption, estimated from measurement of respiratoiyelectron transport system activity, was 0.35-1.07 (mean 0.66)g C m^–1 day^–1 and corresponded, on average, to 79%of the photosynthetic carbon production rate. This implies thatthe major part of photosynthetic fixed organic matter mightbe recycled in the trophogenic layer. The estimated settlingorganic carbon flux at 20 m depth, from calculation of theseparameters and changes in the particulate organic carbon concentration,was 0.01 (-0.09 to 0.13) g C m^–1 day^–1 The meansettling organic carbon flux measured by sediment trap at 20m was 0.19 (0.09-0.31) g C m^–1 day^–1 higher thanthe estimated value. It seemed that organic matter collectedby sediment trap may contain allochthonous matter and resuspendedepilimnetic sediment matter.     

Pelagic carbon metabolism in a eutrophic lake during a clear-water phase

Dissolved and paniculate organic carbon (DOC and POC, respectively),primary production, bacterial production, bacterial carbon demandand community grazing were measured for 9 weeks in eutrophicFrederiksborg Slotssø. The period covered the declineof the spring bloom, a clear-water phase and a summer phasewith increasing phytoplankton biomass. The process rates andchanges in pools of organic carbon were combined in a carbonbudget for the epilimnion. The POC budget showed a close balancefor both the post-spring bloom and the clear-water phase, whilea surplus was found in the summer phase. Production of POC wasdominated by phytoplankton (2/3) compared to bacteria (1/3)during all phases, and there was a significant correlation betweenphytoplankton and bacterial production rates (r² = 0.48, P <0.039). Bacterial demand for DOC was balanced by productionand changes in the pool of DOC during the decline of the springbloom, but the calculated demand exceeded the supply by 81 and167%, respectively, during the other two periods. The discrepancywas most probably due to an underestimation of bacterial growthefficiency and an overestimation of in situ bacterial productionin carbon units. Production of bacterial substrate by zooplanktonactivity was estimated to be higher than the direct excretionof organic carbon from phytoplankton. The biological successionwas regulated by the balance between area primary productionand community grazing. The clear-water phase was initiated bya combination of low primary production due to low surface irradianceand high community grazing (100 mmol C m^–2 day^–1),which caused a decrease in phytoplankton biomass. However, dueto the high initial phytoplankton biomass, community grazingwas not high enough to cause a significant decrease in areaprimary production. The summer phase was initiated by a decreasein community grazing followed by an increase in phytoplanktonbiomass. Based on these observations and calculations of areaprimary production as a function of chlorophyll concentrations,we suggest that the possibility for zooplankton to regulatephytoplankton biomass in temperate lakes decreases with increasingnutrient level.     

Organic carbon release by phytoplankton: its composition and utilization by bacterioplankton

The present study characterized the rate of production of extracellularlyreleased organic carbon (ROC) by phytoplankton, its molecularweight distribution, subsequent utilization and transformationby bacterioplankton in situ. Primary production rate of phytoplanktonwas high during the study due to continuous blooms of smalldinoflagellates and ranged from 59.8 to 298.7µg CI^–1h^–1. The rate of organic carbon release varied from 1.3to 123.7 µg CI^–1 h^–1 and constituted from4.0 to 68.9% of the total carbon fixed in photosynthesis. TheROC was fractionated on molecular weight (MW) basis. A low MWfraction less than 500 daltons (18.5% of ROC), a fraction ofMW 10 000–30 000 daltons (30% of ROC), and high MW fractionof > 300 000 daltons (15.4% of ROC), were the most dominantin ROC. Bacterioplankton utilized a significant portion of ROC,ranging from 18 to 77%. Part of the utilized ROC incorporatedby the bacterioplankton (31 – 56%), and the remainderwas respired (mineralized). ROC not utilized by bacteria wascomposed of high MW compounds. The dynamics of the in situ utilizationof ROC and its role as a link between autotrophic and heterotrophicprocesses in the estuary are described. ¹Present address: Department of Environmental Microbiology,Institute of Microbiology, Warsaw University, Warsaw, Poland     

Increase in the proportion of metabolically active bacteria along gradients of enrichment in freshwater and marine plankton: implications for estimates of bacterial growth and production rates

It is generally recognized that a fraction of all bacterioplanktoncells enumerated using conventional epifluorescence techniquesis neither growing, dividing nor metabolically active, but thevariation in the proportion of active cells among aquatic systemsis not well understood. Here, we hypothesize that the proportionof metabolically active cells increases systematically alonggradients of enrichment, and to test this hypothesis the numberand proportion of metabolically active planktonic bacteria wereinvestigated during the summer in a set of 24 temperate lakes,which span a considerable range in productivity. The tetrazoliumsalt 5-cyano-2,3-ditolyl tetrazolium chloride (CTC) was usedas an indicator of cells with an active electron transport system.The total number of bacteria ranged from 1.88x10⁶ to 7.70x10⁶ml^–1, whereas the number of active cells was more variableand ranged from 0.37x10⁶ to 2.18x10⁶ ml^–1 in the studylakes. The proportion of metabolically active cells ranged from15 to 33%, and tended to increase with nutrient and chlorophyllconcentrations, but not with dissolved organic carbon (DOC).Data on the number of total and active bacteria culled fromthe literature for marine, estuarine and freshwater systemsshow that the trends we measured in lakes are valid for pelagicsystems in general. Over a broad range of aquatic systems, thetotal number of bacteria varied by three orders of magnitude,whereas the number of active bacteria varied by four ordersof magnitude as system productivity increased. The proportionof active cells increased from ultraoligotrophic openocean areas(<5%) to highly productive estuaries (>50%). Our resultssuggest that in most aquatic systems there is a pool of rapidlygrowmg cells, embedded in a usually larger matrix of inactivebacteria, and that the relative size of the active and inactivepools varies systematically among bacterial populations alonggradients of enrichment.     

Variability and predictability of the empirical conversion factor for converting 3H-thymidine uptake into bacterial carbon production for a eutrophic lake

To assess the variability and predictability of the empiricalconversion factor (CF) for converting the uptake of ³H-thymidine(TDR) to bacterial carbon production rates in eutrophic environments,we performed 11 dilution culture experiments and results wererelated to a set of important concomitantly recorded environmentalvariables. TDR incorporation data from a field study were thenconverted to carbon production with the established empiricalCF and a theoretical CF and compared to production estimatesderived from leucine incorporation. Mean empirical CF variedbetween 0.5 and 7.0 x 10⁶ cells (pmol TDR)^–1 over theyear and showed highly significant negative correlation withTDR incorporation rate and a highly significant positive correlationto temperature. Values of carbon production derived with thevariable empirical CF were lower than the values obtained bythe use of the theoretical CF of 1 x 10⁶ cells (pmol TDR)^–1.They showed less seasonal variation than values obtained byleucine incorporation, and periods of uncoupling were observed.However, when the empirical CF was calculated from the multipleregression equation including TDR incorporation and temperature,the resulting carbon production rates showed a high correspondencewith the leucine-derived production rates. Results of the analyseswere interpreted as an indication that under favourable conditions(resulting in high TDR incorporation) bacteria may be able tooptimize DNA duplication over protein synthesis as a possiblestrategy to persist and to maintain their potential to divideunder limiting conditions (e.g. decrease in temperature andsubstrate availability). In unfavourable conditions (resultingin low incorporation rates), bacteria may then use already producedDNA copies for rapid growth, when the environmental conditionsturn favourable again. Thus, an experimental set-up causingnutrient enrichment of the sampled water by autoclaving andfiltering, as generally used for dilution culture experiments,does not always reflect in situ situations, especially duringperiods of low nutrient concentrations.     

Bacteria associated with granular activated carbon particles in drinking water

A sampling protocol was developed to examine particles released from granular activated carbon filter beds. A gauze filter/Swinnex procedure was used to collect carbon fines from 201 granular activated carbon-treated drinking water samples over 12 months. Application of a homogenization procedure (developed previously) indicated that 41.4% of the water samples had heterotrophic plate count bacteria attached to carbon particles. With the enumeration procedures described, heterotrophic plate count bacteria were recovered at an average rate of 8.6 times higher than by conventional analyses. Over 17% of the samples contained carbon particles colonized with coliform bacteria as enumerated with modified most-probable-number and membrane filter techniques. In some instances coliform recoveries were 122 to 1,194 times higher than by standard procedures. Nearly 28% of the coliforms attached to these particles in drinking water exhibited the fecal biotype. Scanning electron micrographs of carbon fines from treated drinking water showed microcolonies of bacteria on particle surfaces. These data indicate that bacteria attached to carbon fines may be an important mechanism by which microorganisms penetrate treatment barriers and enter potable water supplies.     

Bacteria associated with granular activated carbon particles in drinking water.

A sampling protocol was developed to examine particles released from granular activated carbon filter beds. A gauze filter/Swinnex procedure was used to collect carbon fines from 201 granular activated carbon-treated drinking water samples over 12 months. Application of a homogenization procedure (developed previously) indicated that 41.4% of the water samples had heterotrophic plate count bacteria attached to carbon particles. With the enumeration procedures described, heterotrophic plate count bacteria were recovered at an average rate of 8.6 times higher than by conventional analyses. Over 17% of the samples contained carbon particles colonized with coliform bacteria as enumerated with modified most-probable-number and membrane filter techniques. In some instances coliform recoveries were 122 to 1,194 times higher than by standard procedures. Nearly 28% of the coliforms attached to these particles in drinking water exhibited the fecal biotype. Scanning electron micrographs of carbon fines from treated drinking water showed microcolonies of bacteria on particle surfaces. These data indicate that bacteria attached to carbon fines may be an important mechanism by which microorganisms penetrate treatment barriers and enter potable water supplies.     

Phytoplankton photosynthetic efficiency and primary production rates estimated from fast repetition rate fluorometry at coastal embayments affected by upwelling (Rias Baixas, NW of Spain)

Previous studies have not always found a significant relationshipbetween fast repetition rate fluorometry (FRRF)-derived and¹⁴C-based primary production rates. This apparent discrepancymight be related to environmental control of the coupling betweenphotosynthetic electron transport through photosystem II (PSII)and carbon uptake in phytoplankton. In this study, we lookedat this relationship under upwelling conditions favourable forphytoplankton growth. The combination of both techniques allowedthe calculation of the quantum efficiency of carbon fixation,which averaged 0.12 mol C (mol quanta)^–1 indicating thatphytoplankton populations were very efficient in convertingabsorbed photons into photosynthetically produced organic carbon.The tight coupling observed between phytoplankton photosyntheticelectron transport through PSII and carbon uptake resulted ina statistically significant linear relationship between FRRF-derivedand ¹⁴C-based carbon incorporation rates, with a slope of 1.43.In conclusion, the results of the this study indicate that FRRFcan be a useful tool to derive high spatial and temporal resolutionprimary production estimates under environmental conditionsfavouring the close coupling between PSII electron transportand carbon uptake, such as those characteristic of this coastalupwelling system.     

Conspicuous Peak of Oligotrichous Ciliates Following Winter Stratification in a Bog Lake

We analysed early spring protozooplankton peaks following winterstratification in two different basins (northeast, NE; southwest,SW) of an artificially-divided bog lake Große Fuchskuhle(Brandenburg,Germany). The maximum ciliate biomass and numbers (660 and 990µg l^–1 of organic carbon, and 290 and 260 cellsml^–1 in NE and SW, respectively) were reached in the surfacelayer during the ice melting and then continuously decreased.The surface layers were numerically dominated by species ofthe genus Urotricha, while in the ciliate biomass during thefirst part of the study, a mixotrophic oligotrich, Pelagostrombidiummirabile prevailed (C_org up to 940 µg l^–1). We observeda conspicuous ciliate peak that could not be related to thefeeding activity of the dominant ciliate species upon picoplankton.Seemingly, the peak was related to the specific conditions resultingfrom the ice melting where these ciliates could be concentrated.Additionally, the mixotrophy of oligo-trichous species allowedthem to penetrate below the oxycline, although feeding on nanoplanktonsuch as flagellates was also very likely. Our direct measurementsof uptake rates of bacteria showed that the abundant strombidiidswere not nutritionally dependent on bacterivory. However, theciliate uptake rate (up to 670 bacteria ciliate^–1 h^–1)contributed significantly to the overall bacteria mortalityrate. The maximum (community average) cell-specific ciliatefeeding rate of over 200 bacteria ciliates^–1 h^–1was reached, along with an increasing contribution of scuticociliates,paralleled by a drop in large oligotrichs.     

Seasonal variations of bacterial abundance and biomass and their relation to phytoplankton in the hypertrophic tropical lagoon Cienaga Grande de Santa Marta, Colombia

The seasonal development of bacteria was studied in the hypertrophiccoastal lagoon Ciénaga Grande de Santa Marta (Caribbeancoast of Colombia). This large but only 1.5 m deep lagoon issubject to strong seasonal variations of salinity from almostfully marine (April/May) to brackish conditions in October/November.Chlorophyll ranged from 6 to 182 µg L^–1, and grossprimary production amounted to 1690 g C m^–2 per year.Total bacterial number (TBN) ranged from 6.5 to 90.5 x 10⁹ cellsL^–1 and bacterial biomass (BBM) from 77 to 1542 µgC L^–1, which are among the highest ever reported for naturalcoastal waters. Neither TBN nor BBM varied significantly withsalinity, phytoplankton or seston concentrations. Only the bacterialmean cell volume showed a significant relation to salinity,being highest (0.066 µm³) during the period of increasingand lowest (0.032 µm³) during decreasing salinity. Bacterialprotein accounted for 24% (19–26%) and phytoplankton proteinfor 57% (53–71%) of total seston protein. The ratio (annualmean) of bacterial carbon to phytoplankton carbon was 0.44 (range0.04–1.43). At low phytoplankton abundance [chlorophylla (Chl a) < 25 µg L^–1], bacterial carbon wasalmost equal to phytoplankton biomass (i.e. the mean ratio was1.04). In contrast, at Chl a > 100 µg L^–1, BBMwas low compared to phytoplankton biomass (the mean ratio was0.16). In general, BBM varied less than phytoplankton biomass.Most probably, the missing correlation between bacterial andphytoplankton variables was due to (i) organic material partlyderived from allochthonous sources serving as food resourcefor bacteria and (ii) a strong resuspension of bacteria fromthe sediment caused by frequent wind-induced mixing of the veryshallow lagoon.     

Bacterial consumption of total and dissolved organic carbon in the Great Barrier Reef

Heterotrophic bacteria typically take up directly dissolved organic matter due to the small molecular size, although both particulate and dissolved organic matter have labile (easily consumed) compounds. Tropical coastal waters are important ecosystems because of their high productivity. However, few studies have determined bacterial cycling (i.e. carbon uptake by bacteria and allocation for bacterial biomass and respiration) of dissolved organic carbon in coastal tropical waters, and none has determined bacterial cycling of total and dissolved organic carbon simultaneously. In this study we followed bacterial biomass and production, and organic carbon changes over short-term (12 days) dark incubations with (total organic carbon, TOC) and without particulate organic carbon additions (dissolved organic carbon, DOC). The study was performed at three sites along the middle stretch of the Great Barrier Reef (GBR) during the dry and wet seasons. Our results show that the bacterial growth efficiency is low (0.1–11.5%) compared to other coastal tropical systems, and there were no differences in the carbon cycling between organic matter sources, seasons or locations. Nonetheless, more carbon was consumed in the TOC compared to the DOC incubations, although the proportion allocated to biomass and respiration was similar. This suggests that having more bioavailable substrate in the particulate form did not benefit bacteria. Overall, our study indicates that when comparing the obtained respiration rates with previously measured primary production rates, the GBR is a heterotrophic system. More detailed studies are required to fully explore the mechanisms used by bacteria to cycle TOC and DOC in tropical coastal waters.

     

Mineralization in the Ems-Dollard estuary effects on carbon and oxygen budgets

Summary The Ems estuary is a tidal system, showing typical estuaries characteristics, such as gradients of salinity and suspended matter in the water phase. The tidal amplitude is 2.5–3 m. Tidal flats cover 40% of the total area in the outer part and 75% in the innermost part of the estuary, the Dollard. In the sediment aerobic heterotrophic bacteria are concentrated in the upper 2 cm, the numbers rapidly decreasing with depth.Oxygen production and consumption rates in the sediment, and oxygen consumption in the water are measured, together with environmental parameters and numbers of aerobic heterotrophic bacteria. Using a conversion factor of 12/32, oxygen measurements are translated to organic carbon. Input of organic carbon from external sources (particulate carbon from the river Ems and the North Sea, and organic waste discharge) is calculated for the Dollard. An organic carbon budget for the Dollard, using these calculated figures from one year's measurements, showed that most of the organic carbon entering the systems, originated from external sources.Mineralization was quantitatively more important in the sediment than it was in the water phase. The calculated import and production of organic carbon in the system was larger than the calculated consumption. An explanation for this difference is probably the export of soluble organic carbon, which was not measured, to the adjoining Waddensea, which must be considerable. It was concluded that heterotrophic bacteria must play an important role in carbon fluxes in the Dollard and that studies of growth yield under in situ conditions are necessary for a better understanding of their role in the ecosystem.     

Bacterial response to seasonal changes in primary production and phytoplankton biomass in Lake Constance

Parameters characterizing bacterial biomass and metabolic activityare compared with phytoplankton biomass and daily primary productionrates throughout the year. Between late March (before the onsetof the phytoplankton spring bloom) and mid-July (diatom maximum),bacterial degradation of organic matter was more closely relatedto phytoplankton productivity than during the rest of the year.Bacterial production (as estimated by amino acid net uptake)was significantly correlated with concentrations of chlorophyll and pheopigments. However, bacterial production was correlatedless closely with primary production and only weakly with bacterialbiomass. Bacterial biomass was also only weakly correlated withprimary production but significantly with pheopigments. Numbersof active bacteria as estimated by autoradiography covariedclosely with bacterial production and cell numbers. Wheneverbacterial production was low, enhanced proportions of aminoacids were respired. Oxygen consumption measurements showedthat the size fraction <3 µm contributed 25–75%to total respiration. On average, bacterial biomass comprised11 % of paniculate organic matter and roughly equalled phytoplanktonbiomass. During the growing season, bacterial production inthe uppermost 20 m comprised about 20% of phytoplankton productionwith large seasonal fluctuations. A tentative carbon budgetof the euphotic zone including primary production, zooplanktongrazing, bacterial production and sedimentation is presented. ¹Present address: Institute of Marine Resources A-018, ScrippsInstitution of Oceanography, University of California, San Diego,La Jolla, CA 92093, USA     

Relative contribution of bacterial and photosynthetic production to metazooplankton as carbon sources

Percentages of label transfer (PLT) from bacteria and autotrophsto metazooplankton during 4 h were determined in parallel usingdissolved organic and inorganic¹³C tracers, respectively, inin situ batch incubations, to estimate the relative contributionof each production to the metazooplankton food requirement.The batch incubation for each pathway was done eight times,respectively, during 20 days in a mesocosm where continuousspecies succession of organisms was observed. The PLT in thetwo pathways, dependent on metazooplankton abundance and speciescomposition rather than changes in the activities of producers(bacteria or autotrophs), showed a similar tendency of changeand were of a similar magnitude: mean 1.0% (0.09–2.7%)in the photosynthetic pathway and 0.5% (0.22–1.5%) inthe bacterial pathway. This finding suggests that metazooplanktonare supported by entire microbial food webs including both thebacterial and photosynthetic pathways.     

长期施肥条件下水稻土腐殖质组成及稳定性碳同位素特性

利用太湖地区26年水稻土长期定位施肥试验,分析了长期不同施肥处理对土壤剖面有机碳分布和稳定性碳同位素自然丰度(δ¹³C)的影响,以及土壤中不同结合态腐殖质组成的变化.结果表明：长期施肥使水稻土表层土壤有机碳含量显著升高,不同处理土壤剖面有机碳含量与土层深度呈极显著指数负相关(P<0.01);施化肥处理10～30 cm土层和施有机肥处理20～40 cm土层有机碳含量变化相对稳定;随土层深度增加,土壤δ¹³C值逐渐升高,其变化范围在-24‰～-28‰,不同处理土壤剖面有机碳含量与δ¹³C值呈显著线性负相关(P<0.05);0～20 cm土层,仅施有机肥处理(M0)、有机肥+氮+磷处理(MNP)、有机肥+氮+磷+钾处理(MNPK)、有机肥+秸秆+氮处理(MRN)以及秸秆+氮处理(CRN)的δ¹³C值明显降低;30～50 cm土层,除CRN外,有机肥和化肥处理土壤的δ¹³C值均明显升高;不同处理土壤中结合态腐殖质均以紧结合态腐殖质(胡敏素)为主,其含量在50%以上,其余部分为松结合态和稳结合态腐殖质;长期施肥使土壤松结合态腐殖质含量及富啡酸(FA)与胡敏酸(HA)比值(HA/FA)升高.     

Production and Turnover of Planktonic Bacteria in Two Southeastern Blackwater Rivers

Production by attached and free-living planktonic bacteria in two blackwater rivers in the Southeastern United States was measured over a period of 14 months by using the rate of incorporation of [methyl-³H]thymidine into DNA. Production rates and biomass dynamics were compared to determine the potential for in situ production to supply planktonic biomass. Bacterial production in these rivers was moderate and varied seasonally. Rates varied from 0.058 to 2.120 mg of C m⁻³ h⁻¹ in the Ogeechee River and from 0.002 to 2.418 mg of C m⁻³ h⁻¹ in Black Creek. Regressions of growth rate on various environmental variables showed that temperature and total dissolved organic carbon concentration were the best predictors of growth. Although attached bacteria were <21% of the total biomass, they accounted for up to 53% of the total production. Turnover times for attached bacteria ranged from <1 day to >3 years depending on season. Turnover times of free-living bacteria varied from 4.4 days to 11.8 years. Comparisons of biomass with production indicated that during most seasons, the majority of bacterial biomass in these rivers was of allochthonous origin. During summer, when water temperatures were high, bacterial growth in the river may have supplied a greater percentage of the standing stock of bacteria than allochthonous inputs.     

Fatty acid composition in photosynthetic products of natural phytoplankton population in Lake Biwa, Japan

Fatty acid composition of phytoplankton photosynthetic productswas determined by a ¹³C tracer and gas chromatography-mass spectrometry(¹³C-GC-MS) method from August 1985 to June 1986 in Lake Biwa,Japan. The total fatty acid production rate varied from 2.8to 10.9 µg C l^–1 day^–1 at the water surfaceand accounted for 9.1–30% of photosynthetic productionof particulate organic carbon. A high contribution of fattyacid to the particulate organic carbon production rate was noticedduring winter time, and an increase in the fatty acid contributionresulted in an increase in the C/N value in the photosyntheticproducts. The fatty acid composition varied throughout the year,mainly depending on the seasonal change in the dominant phytoplanktonspecies. The contribution of polyunsaturated fatty acids tototal fatty acids was low during the summer period, probablydue to nitrogen limitation of phytoplankton growth.