Iron Constraints on Planktonic Primary Production in Oligotrophic Lakes

Phototrophic primary production is a fundamental ecosystem process, and it is ultimately constrained by access to limiting nutrients. Whereas most research on nutrient limitation of lacustrine phytoplankton has focused on phosphorus (P) and nitrogen (N) limitation, there is growing evidence that iron (Fe) limitation may be more common than previously acknowledged. Here we show that P was the nutrient that stimulated phytoplankton primary production most strongly in seven out of nine bioassay experiments with natural lake water from oligotrophic clearwater lakes. However, Fe put constraints on phytoplankton production in eight lakes. In one of these lakes, Fe was the nutrient that stimulated primary production most, and concurrent P and Fe limitation was observed in seven lakes. The effect of Fe addition increased with decreasing lake water concentrations of total phosphorus and dissolved organic matter. Possible mechanisms are low import rates and low bioavailability of Fe in the absence of organic chelators. The experimental results were used to predict the relative strength of Fe, N, and P limitation in 659 oligotrophic clearwater lakes (with total phosphorus ≤ 0.2 μM P and total organic carbon < 6 mg C l⁻¹) from a national lake survey. Fe was predicted to have a positive effect in 88% of these lakes, and to be the nutrient with the strongest effect in 30% of the lakes. In conclusion, Fe, along with P and N, is an important factor constraining primary production in oligotrophic clearwater lakes, which is a common lake-type throughout the northern biomes. This paper is dedicated to the memory of Prof. Peter Blomqvist (deceased 2004).     

A Comparative Study of Primary Production and Related Factors in Four Saline Lakes in Victoria,Australia

The purpose of the study was to compare the primary plankton productivities of lakes of different salinities and to determine the causative factors involved in their production rates. Four lakes (specific conductivity —mS cm⁻¹ at 18°C) were initially chosen: Coragulac (9), Red Rock (25), Corangamite (38), Pink (250). Sampling and production measurements were made every two to three weeks. Three lakes were dominated by specific phytoplankton blooms: Red Rock (Anabaena spiroides), Corangamite (Nodularia spumigena). Pink (Dunaliella salina). Coragulac Lake had more diverse populations. Red Rock Tarn had some of the highest production values ever recorded. Extremely high soluble phosphate and inorganic carbon concentrations were the most important causative factors. Pink Lake had very low production rates. High salinity and low nutrient concentrations were limiting factors. The other lakes were intermediate in production and nutrient levels. Zooplankton populations were also determined.     

Relationships between picophytoplankton and environmental variables in lakes along a gradient of water colour and nutrient content

SUMMARY 1. Biomass and production of picophytoplankton, phytoplankton and heterotrophic bacterioplankton were measured in seven lakes, exhibiting a broad range in water colour because of humic substances. The aim of the study was to identify environmental variables explaining the absolute and relative importance of picophytoplankton. In addition, two dystrophic lakes were fertilised with inorganic phosphorus and nitrogen, to test eventual nutrient limitation of picophytoplankton in these systems.
2. Picophytoplankton biomass and production were highest in lakes with low concentrations of dissolved organic carbon (DOC), and DOC proved the factor explaining most variation in picophytoplankton biomass and production. The relationship between picophytoplankton and lake trophy was negative, most likely because much P was bound in humic complexes. Picophytoplankton biomass decreased after the additions of P and N.
3. Compared with heterotrophic bacterioplankton, picophytoplankton were most successful at the clearwater end of the lake water colour gradient. Phytoplankton dominated over heterotrophic bacteria in the clearwater systems possibly because heterotrophic bacteria in such lakes are dependent on organic carbon produced by phytoplankton.
4. Compared with other phytoplankton, picophytoplankton did best at intermediate DOC concentrations; flagellates dominated in the humic lakes and large autotrophic phytoplankton in the clearwater lakes.
5. Picophytoplankton were not better competitors than large phytoplankton in situations when heterotrophic bacteria had access to a non-algal carbon source. Neither did their small size lead to picophytoplankton dominance over large phytoplankton in the clearwater lakes. Possible reasons include the ability of larger phytoplankton to float or swim to reduce sedimentation losses and to acquire nutrients by phagotrophy.     

Occurrence,Dynamics and Production of Picoplankton in Hungarian Shallow Lakes

Phototrophic picoplankton were detected in 10⁵-10⁶ cells/ml concentrations in seven water bodies of differing chemistry and trophic level. Dominant picoplankters were, at all sites, coccoid cyanobacteria of 0.8-1.2 μm dimensions, exhibiting red or yellow autofluorescence. Apart from the effects of water temperature their quantitative dynamics were significantly influenced by the nitrogen supply and the herbivorous zooplankton (Cladocera). Their maximum contribution to the total planktonic primary production was about 50%, both in mesotrophic and hypereutrophic environments. However during the bloom of filamentous nitrogen-fixing blue-greens their role became negligible. In phytoplankton communities the significance of picoplankton is overestimated several times, when based on the cell count, while it is underestimated on the basis of biomass. The most useful characteristics of phytoplankton size groups was the total surface area of their cells.     

Experimental Lake Fertilization in the Kuokkel Area,Northern Sweden. Phytoplankton Biomass and Algal Composition in Natural and Fertilized Subarctic Lakes

Based on 500 quantitative algal analyses from 1971–1980 the seasonal variations of phytoplankton biomass and species diversity have been evaluated in one natural and three fertilized subarctic lakes. Enrichment with P alone did not cause any biomass increase and inhibited dinoflagellates and Dinobryon. Enrichment with N alone stimulated Uroglena and Gymnodinium and the biomass increased. No species were inhibited. Enrichment with both N and P made the biomass increase 50–60 times and the stimulated genera were: Chlorella, Choricystis, Chromulina, Chrysochromulina, Gymnodinium, Monoraphidium, Ochromonas, Spiniferomonas, Rhodomonas and Uroglena. In all fertilized lakes monocultures of chlorococcal green algae finally developed.     

我国不同季节陆地植被NPP对气候变化的响应

阐明不同季节陆地植被净第一性生产力(NPP)对全球变化的响应将有助于理解陆地生态系统和气候系统之间的相互作用以及NPP变化机制.本文使用1982～1999年间的AVHRR/NDVI、气温、降水以及太阳辐射等资料,结合植被分布图和土壤质地图,利用生态过程模型,研究不同季节我国陆地植被NPP的年际变化及其地理分异.结果表明,在1982～1999年的18年间,4个季节的NPP都呈显著增加趋势.其中,春季是NPP增加速率最快的季节,夏季是NPP增加量最大的季节.不同植被类型对全球变化的响应有很大差异.常绿阔叶林、常绿针叶林和落叶针叶林NPP的增加主要由生长季节的提前所致,而落叶阔叶林、针阔混交林、矮林灌丛、温带草原及草甸、稀树草原、高寒植被、荒漠以及人工植被NPP的增加主要来自生长季生长加速的贡献.从区域分布看,在四季中春季NPP增加量最大的地区主要集中在东部季风区域;夏季NPP增加量最大的地区包括西北干旱区域和青藏高原的大部分地区、小兴安岭-长白山区、三江平原、松辽平原、四川盆地、雷州半岛、长江中下游部分地区以及江南山地东部;而秋季植被NPP增加量最大的地区主要有云南高原-西藏东部和呼伦湖的周围等地区.不同植被和地理区域NPP的这些响应方式与区域气候特征及其变化趋势有关.     

Bacterial production measured by leucine and thymidine incorporation rates in French lakes

1. Production of heterotrophic bacterioplankton was estimated monthly by the tritiated thymidine and leucine incorporation methods during the draining and filling of the mesotrophic Lake Pareloup (over a 2.5-years sampling program).
2. Rates of ³H-leucine (leu) and ³H-thymidine (thy _DNA) incorporation generally paralleled each other but the ratio of leu/thy _DNA incorporation rates was higher for the draining period (34.5 mean) than during and after filling (11.5 mean).
3. After draining, the highest ratios were observed during periods of low temperature and low bacterial specific activity, while DNA labeling by ³H-thymidine was reduced. However, bacterial production estimates obtained by ³H-leucine (BPL) and ³H-thymidine (BPT) incorporation methods were generally well correlated and the average BPL/BPT ratio was equal to 0.78.
4. In addition, both methods were applied during a diel cycle in three lakes of different trophic status. An increase of leu/thy _DNA incorporation rates was noted from the oligotrophic to the eutrophic system. In the absence of Cyanobacteria, BPL and BPT values were quite concordant on average.
5. In situations of unbalanced growth, BPL and BPT values can diverge but when considered over a sufficient period of time they were found to be in agreement.     

我国不同季节陆地植被NPP对气候变化的响应

阐明不同季节陆地植被净第一性生产力（NPP）对全球变化的响应将有助于理解陆地生态系统和气候系统之间的相互作用以及NPP变化机制。本文使用1982－1999年间的AVHRR／NDVI、气温、降水以及太阳辐射等资料,结合植被分布图和土壤质地图,利用生态过程模型,研究不同季节我国陆地植被NPP的年际变化及其地理分异。结果表明,在1982－1999年的18年间,4个季节的NPP都呈显著增加趋势。其中,春季是NPP增加速率最快的季节,夏季是NPP增加量最大的季节,不同植被类型对全球变化的响应有很大差异。常绿阔叶林,常绿针叶林和落叶针叶林NPP的增加主要由生长季节的提前所致。而落叶阔叶林、针阔混交林、矮林灌丛,温带草原及草甸,稀树草原、高寒植被,荒漠以及人工植被NPP的增加主要来自生长季生长加速的贡献。从区域分布看,在四季中春季NPP增加量最大的地区主要集中在东部季风区域;夏季NPP增量最大的地区包括西北干旱区域和青藏高原的大部分地区,小兴安岭－长白山区,三江平原,松辽平原,四川盆地,雷州半岛,长江中下游部分地区以及江南山地东部;而秋季植被NPP增加量最大的地区主要有云南高原－西藏东部和呼伦湖的周围等地区。不同植被和地理区域NPP的这些响应方式与区域气候特征及其变化趋势有关。     

Phototrophic Picoplankton in Temperate Lakes: Seasonal Abundance and Importance along a Trophic Gradient

The seasonal development of autotrophic picoplankton was investigated in seven Danish lakes representing a eutrophication gradient. Highest cell abundance between 1.5 to 6 × 10⁵ cells ml⁻¹ were found in mid-summer. Minor peaks were observed in spring. In winter, densities were below 10³ ml⁻¹. The highest relative picoplankton contribution to total autotrophic biomass also occurred in mid-summer. In the eutrophic lakes and one humic lake the average seasonal contribution of picoplankton to total chlorophyll was below 1% increasing to 5-8% in the meso- and oligotrophic clear water lakes. During short periods the proportion of picoplankton did reach 25%. The higher relative importance of picoplankton in less productive lakes was not due to higher actual chlorophyll concentrations, but due to a much more pronounced response by larger algae at higher nutrient loading. Both cyanobacteria and eukaryote organisms were present as picoplankton. Only eukaryotes were found in one eutrophic lake and an acidic, humic lake. In the eutrophic lakes eukaryote picoplankton was dominant; both with respect to cell densities and biovolume, whereas cyanobacteria dominated the two meso-oligotrophic lakes. Autotrophic picoplankton were present in all lake types, however their importance seemed to be less in most eutrophic lakes than in less productive, meso-oligotrophic lakes.     

Effects of Climate Change and Shifts in Forest Composition on Forest Net Primary Production

Forests are dynamic in both structure and species composition, and these dynamics are strongly Influenced by climate.However, the net effects of future tree species composition on net primary production (NPP) are not well understood. The objective of this work was to model the potential range shifts of tree species (DISTRIB Model) and predict their impacts on NPP (PnET-Ⅱ Model) that will be associated with alterations in species composition. We selected four 200 × 200 km areas In Wisconsin, Maine, Arkansas, and the Ohio-West Virginia area, representing focal areas of potential species range shifts. PnET-Ⅱ model simulations were carried out assuming that all forests achieved steady state, of which the species compositions were predicted by DISTRIB model with no migration limitation. The total NPP under the current climate ranged from 552 to 908 g C/m2 per year. The effects of potential species redistributions on NPP were moderate (-12% to 8%) compared with the influence of future climatic changes (-60% to 25%). The direction and magnitude of climate change effects on NPP were largely dependent on the degree of warming and water balance. Thus, the magnitude of future climate change can affect the feedback system between the atmosphere and biosphere.     

Seasonality in Nutrients and Phytoplankton Production in Two Shallow Lakes: Waigani Lake,Papua New Guinea,and Barton Broad,Norfolk, England

Waigani Lake, near Port Moresby, Papua New Guinea and Barton Broad, Norfolk, England are both shallow lakes nutrient-enriched from sewage effluent disposal. In Waigani Lake phytoplankton biomass varied seasonally with lower levels (100-200 mg chlorophyll α m⁻³) during the wet season increasing to over 400 mg chlorophyll α m⁻³ at the end of the dry season. Secchi disc depths varied between 0. 11 and 0. 34 m. Phytoplankton productivity in Waigani Lake was very high throughout the year (range: A_max 4,370-21,000 mg O₂ m⁻³ h⁻¹) but production was lower during the wet season (range: A_max 4,370-12,700 mg O₂ m⁻³ h⁻¹). High surface productivity was recorded from August to December except on sampling days when the weather was overcast. Productivity throughout the year declined rapidly with depth. Algal biomass in Barton Broad varied from 3-10 mg chlorophyll α m⁻³ in winter but increased in spring and was very high in summer (200-500 mg chlorophyll α m⁻³). Secchi disc depth varied from 0.21 m in August 1976 to 1.76 m in December. Phytoplankton production in Barton Broad was low in winter (range: A_max 247-1,250 mg O₂ m⁻³ h⁻¹) but increased markedly in spring and summer with the highest rate (A_max 6,850 mg O₂ m⁻³ h⁻¹) being recorded in August. Surface inhibition was observed during summer except when the weather was overcast. Seasonality in nutrients and phytoplankton in Waigani Lake appear to be related to rainfall. Nutrient concentrations in Barton Broad are more closely related to phytoplankton activity which, in turn, correlates with seasonality in solar radiation.     

Interannual Variability in Terrestrial Net Primary Production: Exploration of Trends and Controls on Regional to Global Scales

Climate and biophysical regulation of terrestrial plant production and interannual responses to anomalous events were investigated using the NASA Ames model version of CASA (Carnegie–Ames–Stanford Approach) in a transient simulation mode. This ecosystem model has been calibrated for simulations driven by satellite vegetation index data from the National Oceanic and Atmospheric Administration (NOAA) Advanced Very High Resolution Radiometer (AVHRR) over the mid-1980s. Relatively large net source fluxes of carbon were estimated from terrestrial vegetation about 6 months to 1 year following El Niño events of 1983 and 1987, whereas the years 1984 and 1988 showed a drop in net primary production (NPP) of 1–2 Pg (10¹⁵ g) C from their respective previous years. Zonal discrimination of model results implies that the northern hemisphere low latitudes could account for almost the entire 2 Pg C decrease in global terrestrial NPP predicted from 1983 to 1984. Model estimates further suggest that from 1985 to 1988, the northern middle-latitude zone (between 30° and 60°N) was the principal region driving progressive increases in NPP, mainly by an expanded growing season moving toward the zonal latitude extremes. Comparative regional analysis of model controls on NPP reveals that although Normalized Difference Vegetation Index “greenness” can alone account for 30%–90% of the variation in NPP interannual anomalies, temperature or radiation loading can have a fairly significant 1-year lag effect on annual NPP at middle- to high-latitude zones, whereas rainfall amount and temperature drying effects may carry over with at least a 2-year lag time to influence NPP in semiarid tropical zones.     

Abundance and Diversity of Sulfur-Oxidizing Bacteria along a Salinity Gradient in Four Qinghai-Tibetan Lakes,China

Sulfur-oxidizing bacteria (SOB) play important roles in the sulfur cycle and are widespread in a number of environments, but their occurrence and relationship to geochemical conditions in (hyper)saline lakes are still poorly understood. In this study, the abundance and diversity of SOB populations were investigated in four Qinghai-Tibetan lakes (Erhai Lake, Gahai Lake 1, Gahai Lake 2 and Xiaochaidan Lake) by using quantitative polymerase chain reaction (qPCR) and soxB gene- (encoding sulfate thiohydrolase) based phylogenectic analyses. qPCR analyses showed that in the studied lakes, the total bacterial 16S rRNA and soxB gene abundances in the sediments were distinctly higher than in the overlying waters. The 16S rRNA gene abundance in the waters ranged 5.27 × 10⁶–6.09 × 10⁸ copies per mL and 7.39 × 10¹⁰–2.9 × 10¹¹ copies per gram sediment. The soxB gene abundance in the waters ranged from 1.88 × 10⁴ to 5.21 × 10⁵ per mL and 4.73 × 10⁶–2.65 × 10⁷ copies per gram sediment. The soxB gene in the waters of the two hypersaline lakes (Gahai Lake 2 and Xiaochaidan Lake) was more abundant (2.97 × 10⁵ and 5.21 × 10⁵ copies per mL) than that in the two low-salinity lakes (1.88 × 10⁴ and 3.36 × 10⁴ copies per mL). Phylogenetic analysis showed that Alpha- and Betaproteobacteria were dominant SOB in the investigated lakes, and the composition of proteobacterial subgroups varied with salinity: in freshwater Erhai Lake and low-salinity Gahai Lake 1, the SOB populations were dominated by the Betaproteobacteria, whereas in hypersaline Lake Gahai 2 and Xiaochaidan Lake, the SOB populations were dominated by Alphaproteobacteria. Overall, salinity played a key role in controlling the diversity and distribution of SOB populations in the investigated Qinghai-Tibetan lakes.     

The significance of extracellular production and winter photosynthesis to estimates of primary production in a woodland stream community

Both winter photosynthesis and the release of extracellular DOC are commonly ignored in stream production studies. We examined these contributions in a second-order stream under a completely closed deciduous canopy. We estimate that in Sandy Run approximately 26% of the annual autochthonous particulate carbon is produced between December and March. Measured winter rates of photosynthesis were not significantly different than summertime rates. Contrary to implicit assumptions often made about stream primary productivity, winter production was as important as summer production. Highest rates of carbon assimilation, however, were measured in the spring and fall, and were significantly correlated with standing crops of stream algae as measured by chlorophyll concentration. The recovery of released DOC from stream algae indicated that this contribution was equivalent to 5% of the particulate contribution. Rates of DOC production were significantly correlated with rates of particulate production. We estimate that had winter photosynthesis and extracellular DOC production been ignored in Sandy Run, annual productivity would have been underestimated by about a third. This revised version was published online in July 2006 with corrections to the Cover Date.     

Decline in the Primary Productivity of Northwestern European Forests as a Consequence of Climate Aridization

Average annual age-dependent changes of carbon accumulation in the stemwood of major forest species (pine, spruce, and birch) of the taiga zone of the northwestern European Russia (Karelia) were analyzed. The changes in carbon accumulation were assessed by comparing carbon reserves in tree stands of various ages. Net primary productivity of photosynthesis (NPP) and the proportionality coefficient between respiratory decarboxylation and carbon reserves in wood were calculated. NPP clearly decreased with increasing climate aridization (aridity index). However, the time of the attainment of climax state by a stand did not depend on the latitudinal climate gradient. Hence, only the size of heterotrophic part of phytomass determines annual carbon losses in northern-taiga stands. It is concluded that the climate dependency of the long-term carbon storage in the phytomass of boreal forests is mainly determined by the climate effect on photosynthetic carbon sequestration.__________Translated from Fiziologiya Rastenii, Vol. 52, No. 4, 2005, pp. 513–517.Original Russian Text Copyright © 2005 by Voronin, Konovalov, Bolondinskii, Kaipiainen, Mao.     

华南南亚热带植被第一性生产量的影响因素及预测模型

讨论植被类型、降雨、湿度、土壤条件对华南南亚热带植被第一性生产量的影响。比较各种预测第一性生产量的气候模型,为更好地预测和提高华南南亚热带植被第一性生产量提供依据。     

Primary Production of Phytoplankton in a Strongly Stratified Temperate Lake

Lake Verevi (12.6 ha, maximum depth 11.0 m, mean depth 3.6 m) is a strongly eutrophic and stratified lake. Planktothrix agardhii is the most characteristic phytoplankton species in summer and autumn, while photosynthesizing sulphur bacteria can occur massively in the metalimnion. Primary production (PP) and chlorophyll a concentration (Chl a) were seasonally studied in 1991, 1993, 2000, and 2001. Vertical distribution of PP was rather complex, having usually two peaks, one at or near the surface (0–1 m), and another deeper (at 3–7 m) in the metalimnion. The values of dark fixation of CO₂ in the metalimnion were in most cases higher than those in the upper water layer. Considering the average daily PP 896 mg C m⁻² and yearly PP 162 mg C m⁻², Secchi depth 2.34 m, and epilimnetic concentrations of chlorophyll a (19.6 mg m⁻³), total nitrogen and total phosphorus (TP, 52 mg m⁻³) in 2000, L. Verevi is a eutrophic lake of a ‘good’ status. Considering the total amounts of nutrients stored in the hypolimnion, the average potential concentrations in the whole water column could achieve 1885 mg m⁻³ of TN and 170 mg m⁻³ of TP reflecting hypertrophic conditions and a ‚bad’ status. Improvement of the epilimnetic water quality from the 1990s to the 2000s may have resulted from incomplete spring mixing and might not reflect the real improvement. A decreased nutrient concentration in the epilimnion has supported the establishment of a ‘clear epilimnion state’ allowing light to penetrate into the nutrient-rich metalimnion and sustaining a high production of cyanobacteria and phototrophic sulphur bacteria.     

Diagnosing and assessing uncertainties of terrestrial ecosystem models in a multimodel ensemble experiment: 1. Primary production

We conducted an ensemble modeling exercise using the Terrestrial Observation and Prediction System (TOPS) to evaluate sources of uncertainty in carbon flux estimates resulting from structural differences among ecosystem models. The experiment ran public‐domain versions of biome‐bgc, lpj, casa , and tops‐bgc over North America at 8 km resolution and for the period of 1982–2006. We developed the Hierarchical Framework for Diagnosing Ecosystem Models (HFDEM) to separate the simulated biogeochemistry into a cascade of three functional tiers and sequentially examine their characteristics in climate (temperature–precipitation) and other spaces. Analysis of the simulated annual gross primary production (GPP) in the climate domain indicates a general agreement among the models, all showing optimal GPP in regions where the relationship between annual average temperature (T, °C) and annual total precipitation (P, mm) is defined by P=50T+500. However, differences in simulated GPP are identified in magnitudes and distribution patterns. For forests, the GPP gradient along P=50T+500 ranges from ～50 g C yr^?1 m^?2 °C^?1 (casa ) to ～125 g C yr^?1 m^?2 °C^?1 (biome‐bgc ) in cold/temperate regions; for nonforests, the diversity among GPP distributions is even larger. Positive linear relationships are found between annual GPP and annual mean leaf area index (LAI) in all models. For biome‐bgc and lpj , such relationships lead to a positive feedback from LAI growth to GPP enhancement. Different approaches to constrain this feedback lead to different sensitivity of the models to disturbances such as fire, which contribute significantly to the diversity in GPP stated above. The ratios between independently simulated NPP and GPP are close to 50% on average; however, their distribution patterns vary significantly between models, reflecting the difficulties in estimating autotrophic respiration across various climate regimes. Although these results are drawn from our experiments with the tested model versions, the developed methodology has potential for other model exercises.     

土地利用约束下中国东部南北样带生产力和植被分布对全球变化的响应

对现有的区域植被动态模拟模型进行了改进,使之包含了土地利用分布格局对植被和生态系统相关过程的影响。改进后的模型被用地研究中国东部南北样带(NSTEC)植被和净第一性生产力对未来气候变化的响应。模拟结果显示土地利用格局对未来气候条件下植被分布的变迁和生产力形成过程有非常显著的影响。与没有土地利用约束的情形相比较,土地利用作为限制条件缓减了植被类型之间的竞争,从而减少了模拟的样带区域内常绿阔叶林,但增加了模拟灌木和草地的分布。土地利用约束使得模拟得到的当前条件下的净第一性生产力更为接近实际情况,且未来气候条件下的生产力改变量更为可信。对未来CO2倍增条件下7个大气环流模型预测的气候情景的模拟结果表明：落叶阔叶林将显著增加,但针叶林、灌木和草原的分布将下降。未来气候条件下NSTEC样带的净第一性生产力总量将增加。预测样带北部的净第一性生产力的变化范围大于样带南部。温度变化比降水变化对样带的生产力具有更强的控制。