Phytoplankton exudates provide full nutrition to a subset of accompanying heterotrophic bacteria via carbon,nitrogen and phosphorus allocation

Marine bacteria rely on phytoplankton exudates as carbon sources (DOCp). Yet, it is unclear to what extent phytoplankton exudates also provide nutrients such as phytoplankton-derived N and P (DONp, DOPp). We address these questions by mesocosm exudate addition experiments with spent media from the ubiquitous pico-cyanobacterium Prochlorococcus to bacterial communities in contrasting ecosystems in the Eastern Mediterranean – a coastal and an open-ocean, oligotrophic station with and without on-top additions of inorganic nutrients. Inorganic nutrient addition did not lower the incorporation of exudate DONp, nor did it reduce alkaline phosphatase activity, suggesting that bacterial communities are able to exclusively cover their nitrogen and phosphorus demands with organic forms provided by phytoplankton exudates. Approximately half of the cells in each ecosystem took up detectable amounts of Prochlorococcus-derived C and N, yet based on 16S rRNA sequencing different bacterial genera were responsible for the observed exudate utilization patterns. In the coastal community, several phylotypes of Aureimarina, Psychrosphaera and Glaciecola responded positively to the addition of phytoplankton exudates, whereas phylotypes of Pseudoalteromonas increased and dominated the open-ocean communities. Together, our results strongly indicate that phytoplankton exudates provide coastal and open-ocean bacterial communities with organic carbon, nitrogen and phosphorus, and that phytoplankton exudate serve a full-fledged meal for the accompanying bacterial community in the nutrient-poor eastern Mediterranean.     

Efficient CO2 fixation by surface Prochlorococcus in the Atlantic Ocean

Nearly half of the Earth''s surface is covered by the ocean populated by the most abundant photosynthetic organisms on the planet—Prochlorococcus cyanobacteria. However, in the oligotrophic open ocean, the majority of their cells in the top half of the photic layer have levels of photosynthetic pigmentation barely detectable by flow cytometry, suggesting low efficiency of CO₂ fixation compared with other phytoplankton living in the same waters. To test the latter assumption, CO₂ fixation rates of flow cytometrically sorted ¹⁴C-labelled phytoplankton cells were directly compared in surface waters of the open Atlantic Ocean (30°S to 30°N). CO₂ fixation rates of Prochlorococcus are at least 1.5–2.0 times higher than CO₂ fixation rates of the smallest plastidic protists and Synechococcus cyanobacteria when normalised to photosynthetic pigmentation assessed using cellular red autofluorescence. Therefore, our data indicate that in oligotrophic oceanic surface waters, pigment minimisation allows Prochlorococcus cells to harvest plentiful sunlight more effectively than other phytoplankton.     

Mineralisation of dissolved organic matter by heterotrophic stream biofilm communities in a large boreal catchment

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Co-culture with Synechococcus facilitates growth of Prochlorococcus under ocean acidification conditions

Anthropogenic CO₂ emissions are projected to lower the pH of the ocean 0.3 units by 2100. Previous studies suggested that Prochlorococcus and Synechococcus, the numerically dominant phytoplankton in the oceans, have different responses to elevated CO₂ that may result in a dramatic shift in their relative abundances in future oceans. Here we showed that the exponential growth rates of these two genera respond to future CO₂ conditions in a manner similar to other cyanobacteria, but Prochlorococcus strains had significantly lower realized growth rates under elevated CO₂ regimes due to poor survival after exposure to fresh culture media. Despite this, a Synechococcus strain was unable to outcompete a Prochlorococcus strain in co-culture at elevated CO₂. Under these conditions, Prochlorococcus' poor response to elevated CO₂ disappeared, and Prochlorococcus' relative fitness showed negative frequency dependence, with both competitors having significant fitness advantages when initially rare. These experiments suggested that the two strains should be able to coexist indefinitely in co-culture despite sharing nearly identical nutritional requirements. We speculate that negative frequency dependence exists due to reductive Black Queen evolution that has resulted in a passively mutualistic relationship analogous to that connecting Prochlorococcus with the ‘helper’ heterotrophic microbes in its environment.     

Spatial distribution of small phytoplankton composition in the Chukchi Sea

This study was conducted as part of the second Russian American long-term census of the Arctic in 2009 with sampling across the territorial of the Russian Federation in the Chukchi Sea since recent information regarding the standing stocks of pico- and nano-plankton is very limited. Using flow cytometry, small size fractions of the phytoplankton (<20 μm) obtained during the cruise were analyzed for the contributions of pico- and nano-eukaryotic phytoplankton and prokaryotic phytoplankton for the first time in the Chukchi Sea. The salinity and temperature in the southern part were significantly higher than those in the northern part of the Chukchi Sea. The abundance of Prochlorococcus and Synechococcus represented about 30 % of total small phytoplankton cells with their cell abundances reaching 295 and 590 cells ml^?1, respectively, although a significant portion (about 70 %) of the small phytoplankton community (<20 μm) was pico- and nano-eukaryotic plankton in the Chukchi Sea. Among different environmental factors, we found that temperature and NH₄ concentrations were significantly positively correlated with the abundance of total phytoplankton, pico- and nano-eukaryotes, Prochlorococcus, and Synechococcus. Projected higher water temperature and increase in NH₄ concentration condition in the Arctic Ocean as well as the Chukchi Sea could have fostered more small phytoplankton communities especially Prochlorococcus and Synechococcus.     

Soil microbial activity in relation to dissolved organic matter properties under different tree species

Background and aims

The total concentration of dissolved organic carbon (DOC) has often been observed to correlate positively with soil microbial respiration. The aim was to explain the correlation with the properties of dissolved organic matter (DOM).

Methods

A dataset from previously published papers was gathered together and subjected to multivariate analyses. Samples were collected from five tree species experiments in Finland. The degradability of DOM was assessed by measuring bacterial and fungal growth in DOM. The chemical properties of DOM were assessed by XAD resin fractionation and molecular weight. Soil microbial activity was assessed as C and N mineralization and microbial biomass.

Results

Both low and high molecular weight compounds, as well as hydrophilic neutral compounds, seemed to be relatively easily degradable. In contrast to our presupposition, easily degradable DOM seemed to be less abundant in soil where variables describing microbial activity were higher. Birch soil with higher microbial biomass N seemed to contain less easily degradable DOM than spruce soil.

Conclusion

We suggest that DOM collected and characterized at a certain point reflects more the accumulation of refractory compounds following high microbial activity than the easily degradable compounds that microbes would be using when measured.

     

Use of electrospray ionization (ESI) mass spectrometry to investigate complex dissolved organic matter (DOM) and its potential applications in phytoplankton research

Organic nutrients are one of many factors considered to be important in the growth and proliferation of phytoplankton including many species that cause harmful algal blooms (HABs). Several studies have investigated the effects of known organic compounds on phytoplankton growth, however, the role of natural dissolved organic matter (DOM) in phytoplankton nutrition remains understudied at the compound level. This lack of research is due in part to analytical limitations for the characterization of DOM compounds. Electrospray ionization (ESI) mass spectrometry (MS) provides an unprecedented level of chemical information on thousands of organic compounds that comprise the bulk DOM pool. In this paper we provide a brief overview of some of the benefits and caveats of using ESI to investigate DOM in natural freshwater and marine systems and show an example of ESI-MS DOM characterization for a natural bloom of the raphidophyte Chattonella cf. verruculosa.     

原绿球藻对环境胁迫的生理和分子响应机制

原绿球藻(Prochlorococcus)作为海洋丰度最高的浮游植物,对海洋生态系统的物质循环和能量流动起着重要的驱动作用。原绿球藻生长和光合作用活性容易受到环境胁迫的影响,进而影响整个海洋生态系统的稳定性。因此,研究原绿球藻应对环境胁迫的响应机制具有重要的生态意义。原绿球藻主要通过分化出不同的生态型来适应不同光照和营养盐的海洋环境,但仍然会很难快速适应各种突如其来的海洋环境变化。本文从原绿球藻应对环境胁迫的角度,探讨了其生理和分子响应机制的最新研究进展,包括光系统I循环电子传递在光照变化时发挥的重要作用,通过RNA快速响应而调控基因表达应对环境胁迫,以及在辅助异养细菌的保护下应对活性氧的胁迫等。本文也展望了原绿球藻对环境胁迫响应的生理和分子机制的未来研究方向,旨在为原绿球藻抗逆机制的深入研究提供参考。     

DOM对米槠次生林不同土层土壤微生物呼吸及其熵值的影响

可溶性有机质(Dissolved organic matter,DOM)作为土壤可溶性有机碳的重要来源,进入土壤之后通过改变土壤微生物数量和活性影响土壤矿化。DOM输入对土壤微生物呼吸和熵值的研究多集中在表层土壤,但对深层土壤微生物呼吸和熵值的影响关注较少。通过室内培养实验(120 d)研究米槠(Castanopsis carlesii)鲜叶DOM添加对表层土壤(0—10 cm)和深层土壤(40—60 cm)微生物呼吸及其土壤代谢熵和微生物熵的影响,为揭示DOM输入对亚热带森林土壤碳过程的影响提供理论依据。结果表明,在培养第1天,添加DOM的表层和深层土壤CO_2瞬时排放速率均显著高于对照(P0.001),分别是对照(不添加DOM)的3.58倍和6.93倍,之后显著下降。就累积排放量而言,无论是DOM添加处理还是对照,表层土壤显著大于深层土壤;在米槠鲜叶DOM添加后,表层土壤累积排放量显著大于对照的表层土壤(P0.001),但DOM添加处理深层土壤累积排放量与对照的深层土壤无明显差异。就微生物生物量碳而言,表层土壤微生物生物量碳含量在培养期间显著大于深层土壤。在整个添加DOM培养期间,表层土壤微生物生物量碳含量显著大于表层对照土壤,深层土壤微生物生物量碳含量显著大于深层对照土壤(第3天除外)。培养结束时(120 d),米槠鲜叶DOM添加处理下,表层土壤和深层土壤有机碳含量与第3天相比分别减少26%和19%。米槠鲜叶DOM添加处理后的深层土壤代谢熵(qCO_2)显著低于对照的深层土壤和DOM添加处理的表层土壤qCO_2(P0.001),说明外源DOM进入深层土壤后提高了土壤微生物对碳的利用效率。米槠鲜叶DOM添加处理后的深层土壤微生物熵是培养第3天的1.58倍,显著大于培养初期(P0.05),而DOM添加处理的表层土壤、对照的表层土壤与深层土壤的微生物熵分别是培养第3天的68%、79%和21%,说明DOM添加提高了深层土壤质量。     

Viral lysis of Micromonas pusilla: impacts on dissolved organic matter production and composition

The viral mediated transformation of phytoplankton organic carbon to dissolved forms (“viral shunt”) has been suggested as a major source of dissolved organic carbon (DOC) in marine systems. Despite the potential implications of viral activity on the global carbon fluxes, studies investigating changes in the DOC composition from viral lysis is still lacking. Micromonas pusilla is an ecologically relevant picoeukaryotic phytoplankter, widely distributed in both coastal and oceanic marine waters. Viruses have been found to play a key role in regulating the population dynamics of this species. In this study we used axenic cultures of exponentially growing M. pusilla to determine the impact of viral lysis on the DOC concentration and composition, as estimated from lysate-derived production of transparent exopolymer particles (TEP) and two fractions of fluorescent dissolved organic matter (DOM): aromatic amino acids (excitation/emission; 280/320 nm; F(280/320)) and marine humic-like fluorescent DOM (320/410 nm; F(320/410)). DOC concentration increased 4.5 times faster and reached 2.6 times higher end concentration in the viral infected compared with the non-infected cultures. The production of F(280/320) and F(320/410) were 4.1 and 2.8 times higher in the infected cultures, and the elevated ratio between F(280/320) and F(320/410) in lysates suggested a higher contribution of labile (protein) components in viral produced DOM than in algal exudates. The TEP production was 1.8 times faster and reached a 1.5 times higher level in the viral infected M. pusilla culture compared with the non- infected cultures. The measured increase in both DOC and TEP concentrations suggests that viral lysis has multiple and opposite implications for the production and export processes in the pelagic ocean: (1) by releasing host biomass as DOC it decreases the organic matter sedimentation and promotes respiration and nutrient retention in the photic zone, whereas (2) the observed enhanced TEP production could stimulate particle aggregation and thus carbon export out of the photic zone.     

Analogous nutrient limitations in unicellular diazotrophs and Prochlorococcus in the South Pacific Ocean

Growth limitation of phytoplankton and unicellular nitrogen (N₂) fixers (diazotrophs) were investigated in the oligotrophic Western South Pacific Ocean. Based on change in abundances of nifH or 23S rRNA gene copies during nutrient-enrichment experiments, the factors limiting net growth of the unicellular diazotrophs UCYN-A (Group A), Crocosphaera watsonii, γ-Proteobacterium 24774A11, and the non-diazotrophic picocyanobacterium Prochlorococcus, varied within the region. At the westernmost stations, numbers were enhanced by organic carbon added as simple sugars, a combination of iron and an organic chelator, or iron added with phosphate. At stations nearest the equator, the nutrient-limiting growth was not apparent. Maximum net growth rates for UCYN-A, C. watsonii and γ-24774A11 were 0.19, 0.61 and 0.52 d⁻¹, respectively, which are the first known empirical growth rates reported for the uncultivated UCYN-A and the γ-24774A11. The addition of N enhanced total phytoplankton biomass up to 5-fold, and the non-N₂-fixing Synechococcus was among the groups that responded favorably to N addition. Nitrogen was the major nutrient-limiting phytoplankton biomass in the Western South Pacific Ocean, while availability of organic carbon or iron and organic chelator appear to limit abundances of unicellular diazotrophs. Lack of phytoplankton response to nutrient additions in the Pacific warm pool waters suggests diazotroph growth in this area is controlled by different factors than in the higher latitudes, which may partially explain previously observed variability in community composition in the region.     

Size‐spectrum based differential response of phytoplankton to nutrient and iron‐organic matter combinations in microcosm experiments in a Chilean Patagonian Fjord

The Patagonian fjords have been recognized as a major region of relatively high primary productivity systems during spring–summer bloom periods, where iron‐organic matter forms may be essential complexes involved in key growth processes connected to the carbon and nitrogen cycles. We used two dissolved organic matter (DOM) types, marine polysaccharide and siderophore, as a model to understand how they affect the bioavailability of Fe to phytoplankton and bacteria and to assess their ecological role in fjord systems. A 10‐day microcosm study was performed in the Comau Fjord during summer conditions (March 2012). Pico‐, nano‐, and microphytoplankton abundance, total chlorophyll‐a and bacteria abundance, and bacterial secondary production estimates were analyzed in five treatments: (i) control (no additions), (ii) only nutrients (NUT: PO₄, NO₃, Si), (iii) nutrients + Fe(II), (iv) polysaccharide (natural diatoms extracted: 1–3 beta Glucan), and (v) Hexandentate Desferroxiamine B (DFB, siderophore). Our results showed that while DFB reduced Fe bioavailability for almost all phytoplankton assemblages in the fjord, polysaccharide did not have effects on the iron bioavailability. At Nutrients + Fe and Polysaccharide treatments, chlorophyll‐a concentration abruptly increased from 0.9 to 20 mg m^?3 during the first 4–6 days of the experimental period. Remarkably, at the Nutrients + Fe treatment, the development of the bloom was accompanied by markedly high abundances of Synechococcus, picoeukaryotes, and autotrophic nanoflagellates within the first 4 days of the experiment. Our study indicated that small plankton (phytoplankton <20 μm and bacteria) were the first to respond to dissolved Nutrients + Fe compared to large sized micro‐phytoplankton cells (>20 μm). This could be at least partially attributed to biological utilization of Fe (2 to 3 nM) by <20 μm phytoplankton and bacteria through the interaction with organic ligands released by bacteria that eventually could increase solubility of the Fe dissolved fraction thus having a positive effect on the small‐sized phytoplankton community.     

有机碳质量对黄河三角洲芦苇凋落物分解及其温度敏感性的影响

凋落物分解速率及其温度敏感性Q_(10)能够影响凋落物对土壤的碳归还及其对全球变暖的响应。然而,凋落物有机碳质量对凋落物分解及其温度敏感性的影响研究仍不充分。以黄河三角洲芦苇(Phragmites australi)为例,通过凋落物袋法、室内模拟实验及固态~(13)C核磁共振技术,研究有机碳质量对凋落物分解及其温度敏感性的影响,探讨预测凋落物分解及其温度敏感性的指标。结果表明:(1)随着凋落物分解,易分解碳组分(烷氧碳、双烷氧碳)相对含量逐渐降低,而难分解碳组分(芳香碳)相对含量显著增加,疏水碳/亲水碳、芳香碳/烷氧碳比值逐渐增大,凋落物有机碳更加稳定,凋落物呼吸速率及失重率呈下降趋势。(2)凋落物失重主要受烷基碳、烷氧碳相对含量及C/N的影响,凋落物CO_2累积释放量主要受烷氧碳及双烷氧碳相对含量的影响。羰基碳相对含量可以用来解释Q_(10)的变异。因此,相对于生态化学计量比,烷基碳、烷氧碳、双烷氧碳、羰基碳相对含量是预测凋落物分解及其温度敏感性的敏感性指标。     

Elevated pCO2 alters marine heterotrophic bacterial community composition and metabolic potential in response to a pulse of phytoplankton organic matter

Factors that affect the respiration of organic carbon by marine bacteria can alter the extent to which the oceans act as a sink of atmospheric carbon dioxide. We designed seawater dilution experiments to assess the effect of pCO₂ enrichment on heterotrophic bacterial community composition and metabolic potential in response to a pulse of phytoplankton-derived organic carbon. Experiments included treatments of elevated (1000 p.p.m.) and low (250 p.p.m.) pCO₂ amended with 10 μmol L⁻¹ dissolved organic carbon from Emiliana huxleyi lysates, and were conducted using surface-seawater collected from the South Pacific Subtropical Gyre. To assess differences in community composition and metabolic potential, shotgun metagenomic libraries were sequenced from low and elevated pCO₂ treatments collected at the start of the experiment and following exponential growth. Our results indicate bacterial communities changed markedly in response to the organic matter pulse over time and were significantly affected by pCO₂ enrichment. Elevated pCO₂ also had disproportionate effects on the abundance of sequences related to proton pumps, carbohydrate metabolism, modifications of the phospholipid bilayer, resistance to toxic compounds and conjugative transfer. These results contribute to a growing understanding of the effects of elevated pCO₂ on bacteria-mediated carbon cycling during phytoplankton bloom conditions in the marine environment.     

有机磷培养下水体浮游植物竞争与群落结构演替

磷是浮游植物生长的必要元素,也是调控水体富营养化的重要因子之一。在无机磷缺乏的水域,能够有效利用有机磷的藻类有可能成为浮游植物群落的优势种。为了研究有机磷对浮游植物群落结构演替的影响,分别从3个水库采集表层水样,西陂水库(硅藻70.3%,甲藻19.1%,绿藻9.3%)、山美水库(硅藻31.3%,甲藻59.2%,绿藻3.5%)、三十六脚湖(硅藻43.1%,甲藻50.5%,绿藻5.0%),选用有机磷单磷酸腺苷(Adenosine monophosphate,AMP)作为磷源进行实验室模拟实验,并与无机磷(NaH_2PO_4·2H_2O)作为磷源实验组做同步对比,探讨有机磷源条件下浮游植物竞争及群落结构演替规律。结果显示:AMP培养20 d后,西陂水库、山美水库和三十六脚湖的浮游植物生物量分别增加了6.5倍,2.1倍和1.9倍,浮游植物群落结构都演替为以甲藻门和绿藻门为优势藻,所占比例分别为81.8%和16.7%、55.2%和33.5%、73.2%和24.3%,其中西陂水库甲藻门拟多甲藻对有机磷AMP的利用能力强,其所占比例达到81.8%。表明有机磷AMP能有效地促进甲藻门拟多甲藻属的增殖,进而改变水体浮游植物群落结构。     

杉木和米槠凋落叶DOM对土壤碳矿化的影响

DOM(Dissolved organic matter)是土壤微生物呼吸的重要底物,凋落物淋溶的DOM对土壤碳矿化具有重要影响。选择中亚热带地区具有代表性的杉木(Cunninghamia lanceolata)和米槠(Castanopsis carlesii)凋落叶作为研究对象,通过两个月的短期室内培养,把不同凋落叶浸提出的DOM添加到培养瓶中,定期测定土壤碳矿化速率,计算土壤碳累积矿化量,探讨两种等浓度等量DOM添加对土壤碳矿化的影响,并分析DOM化学性质在土壤碳矿化过程中的重要性。结果表明:米槠凋落叶浸提得到的DOC(Dissolved organic carbon)和DON(Dissolved organic nitrogen)浓度均显著高于杉木凋落叶的(P0.05),而杉木凋落叶浸提得到的DOM的UV吸收值(SUVA_(254))和HIX(Humification index)均显著低于米槠凋落叶的(P0.01)。添加等浓度等量杉木和米槠凋落叶DOM到土壤中均显著增加了土壤碳矿化速率,在第1天内分别比对照高198%和168%,3d后下降到61.8%和44.1%,14d后基本处于平稳状态,表明外源有机物添加对土壤碳矿化的前期影响较大。培养过程中,添加杉木和米槠凋落叶DOM的土壤碳矿化累积量均能采用双因素指数模型进行拟合(r~2=0.99),但添加两者凋落叶DOM后土壤碳矿化累积量没有显著差异。     

Environmental Stress Determines the Quality of Bacterial Lysate and Its Utilization Efficiency in a Simple Microbial Loop

Heterotrophic bacteria provide the critical link in the microbial loop by converting dissolved organic matter (DOM) into particulate form. In this study, DOM was prepared from recently isolated estuarine bacterial strain Vibrio sp. (DSM14379) grown at different salinities [0.2%, 0.5%, 3%, 5%, or 10% (w/v)], washed, concentrated, and lysed by autoclaving. The corresponding lysate-containing media were designated LM_0.2, LM_0.5, LM₃, LM₅, and LM₁₀. Vibrio sp. cells grown at different salinities had similar C/N/P ratios, but different C/S ratios, different trace element composition, and different 2D gel electrophoresis protein profiles. Pseudoalteromonas sp. (DSM06238) isolated from a similar environment was able to grow on all lysates, and its biomass production was dependent on lysate type. The highest growth rate and biomass production of Pseudoalteromonas sp. at saturation lysate concentrations were observed in LM₃. The biomass production at saturation lysate concentrations was about 3-fold higher as compared to LM_0.2 and LM₁₀. The initial respiration rate, intracellular adenosine triphosphate (ATP) levels, and ³H-Leu and ³H-TdR incorporation rates were lowest in LM₃. On the other hand, in LM_0.2 or LM₁₀ lysates the situation was reversed, the growth rates and biomass production were lowest, whereas ³H-Leu and ³H-TdR incorporation, respiration rates, as well as ATP levels, were highest. These results imply uncoupling of catabolism from growth in either high- or low-salinity lysates. The results also suggest that differences in organic carbon quality generated during Vibrio sp. growth at different NaCl concentrations were propagated through the simple microbial loop, which may have important ecological implications for higher trophic levels that depend on microbial grazing.