Nutritional diagnosis of phytoplankton in Lake Baikal

To diagnose the nutritional status of phytoplankton in Lake Baikal, surveys for the determination of concentrations of particulate carbon (PC), nitrogen (PN) and phosphorus (PP) and their ratios were conducted at six stations in March, June, August and October 1999. The concentrations of PC and PN were lower than, and those of PP were similar to, those in another mesotrophic lake except at the station near the mouth of the largest input river, Selenga River, of Lake Baikal. The PC : PN : PP ratio was 102 : 13 : 1, considerably close to the Redfield ratio. The ratio was constant against spatiotemporal changes. These indicate that phytoplankton in Lake Baikal were exposed to no deficiency in nitrogen nor phosphorus. From a viewpoint of the nutritional status of phytoplankton, Lake Baikal might be viewed as an ocean rather than as a lake.     

Beyond macronutrients: element variability and multielement stoichiometry in freshwater invertebrates

We contrasted concentrations of macronutrients (C, N and P), essential (As, Cu, Zn and Se) and non-essential metals (Pb, Hg and Cd) in invertebrates across five lakes and June to October in one lake. We predicted that somatic concentrations of tightly regulated elements would be less variable than weakly and unregulated elements. Within each taxon, variation was lowest in macronutrients, intermediate in essential micronutrients, and highest in non-essential metals, which corresponded in rank to homeostatic regulation strength for the same elements calculated from the literature. Hence, homeostatic regulation may strongly influence variation in element concentrations of biota in situ . Of the individual elements, only taxonomic differences in C and N were consistent across lakes and over a season. Nevertheless, canonical discriminant analyses successfully discriminated among taxa based on taxonomic multielement composition. Thus, relative taxonomic differences in multielement composition appear more informative than absolute stoichiometric formulae when considering the role of inherently variable trace elements in ecological investigations.     

Phytoplankton stoichiometry

Because phytoplankton live at the interface between the abiotic and the biotic compartments of ecosystems, they play an important role in coupling multiple nutrient cycles. The quantitative details of how these multiple nutrient cycles intersect is determined by phytoplankton stoichiometry. Here we review some classic work and recent advances on the determinants of phytoplankton stoichiometry and their role in determining ecosystem stoichiometry. First, we use a model of growth with flexible stoichiometry to reexamine Rhee and Goldman’s classic chemostat data. We also discuss a recent data compilation by Hall and colleagues that illustrates some limits to phytoplankton flexibility, and a model of physiological adaptation that can account for these results. Second, we use a model of resource allocation to determine the how the optimal nitrogen-to-phosphorus stoichiometry depends on the ecological conditions under which species grow and compete. Third, we discuss Redfield’s mechanism for the homeostasis of the oceans’ nitrogen-to-phosphorus stoichiometry and show its robustness to additional factors such as iron-limitation and temporal fluctuations. Finally, we suggest areas for future research.     

浮游动物化学计量学稳态性特征研究进展

稳态性是有机体的基本属性,也是生态化学计量学理论成立的前提和基础。一般来讲,浮游植物的元素组成变化较大,而浮游动物具有明显的稳态性特征。浮游动物稳态性特征的研究不仅有助于了解水生生态系统的能量流动和物质循环,同时也对研究营养元素如何调节生物生长、繁殖和代谢起到促进作用。在综述生态化学计量学研究的基础上,主要介绍了稳态性的概念和浮游动物稳态性特征的基本框架及变化规律,以期为促进国内相关研究工作的开展提供参考。     

THE ELEMENTAL COMPOSITION OF SOME MARINE PHYTOPLANKTON1

We analyzed the cellular content of C, N, P, S, K, Mg, Ca, Sr, Fe, Mn, Zn, Cu, Co, Cd, and Mo in 15 marine eukaryotic phytoplankton species in culture representing the major marine phyla. All the organisms were grown under identical culture conditions, in a medium designed to allow rapid growth while minimizing precipitation of iron hydroxide. The cellular concentrations of all metals, phosphorus, and sulfur were determined by high‐resolution inductively coupled plasma mass spectrometry (HR‐ICPMS) and those of carbon and nitrogen by a carbon hydrogen nitrogen analyzer. Accuracy of the HR‐ICPMS method was validated by comparison with data obtained with ⁵⁵Fe radioactive tracer and by a planktonic reference material. The cellular quotas (normalized to P) of trace metals and major cations in the biomass varied by a factor of about 20 among species (except for Cd, which varied over two orders of magnitude) compared with factors of 5 to 10 for major nutrients. Green algae had generally higher C, N, Fe, Zn, and Cu quotas and lower S, K, Ca, Sr, Mn, Co, and Cd quotas than coccolithophores and diatoms. Co and Cd quotas were also lower in diatoms than in coccolithophores. Although trace element quotas are influenced by a variety of growth conditions, a comparison of our results with published data suggests that the measured compositions reflect chiefly the intrinsic (i.e. genetically encoded) trace element physiology of the individual species. Published field data on the composition of the planktonic biomass fall within the range of laboratory values and are generally close to the approximate extended Redfield formula given by the average stoichiometry of our model species (excluding the hard parts): While clearly this elemental stoichiometry varies between species and, potentially, in response to changes in the chemistry of seawater, it provides a basis for examining how phytoplankton influence the relative distributions of the ensemble of major and trace elements in the ocean.     

Elemental stoichiometry of the key calcifying marine phytoplankton Emiliania huxleyi under ocean climate change: A meta-analysis

The elemental composition of marine microorganisms (their C:N:P ratio, or stoichiometry) is central to understanding the biotic and biogeochemical processes underlying key marine ecosystem functions. Phytoplankton C:N:P is species specific and flexible to changing environmental conditions. However, bulk or fixed phytoplankton stoichiometry is usually assumed in biogeochemical and ecological models because more realistic, environmentally responsive C:N:P ratios have yet to be defined for key functional groups. Here, a comprehensive meta-analysis of experimental laboratory data reveals the variable C:N:P stoichiometry of Emiliania huxleyi, a globally significant calcifying phytoplankton species. Mean C:N:P of E. huxleyi is 124C:16N:1P under control conditions (i.e. growth not limited by one or more environmental stressors) and shows a range of responses to changes in nutrient and light availability, temperature and pCO₂. Macronutrient limitation caused strong shifts in stoichiometry, increasing N:P and C:P under P deficiency (by 305% and 493% respectively) and doubling C:N under N deficiency. Responses to light, temperature and pCO₂ were mixed but typically shifted cellular elemental content and C:N:P stoichiometry by ca. 30% or less. Besides these independent effects, the interactive effects of multiple environmental changes on E. huxleyi stoichiometry under future ocean conditions could be additive, synergistic or antagonistic. To synthesise our meta-analysis results, we explored how the cellular elemental content and C:N:P stoichiometry of E. huxleyi may respond to two hypothetical future ocean scenarios (increased temperature, irradiance and pCO₂ combined with either N deficiency or P deficiency) if an additive effect is assumed. Both future scenarios indicate decreased calcification (which is predominantly sensitive to elevated pCO₂), increased C:N, and up to fourfold shifts in C:P and N:P. Our results strongly suggest that climate change will significantly alter the role of E. huxleyi (and potentially other calcifying phytoplankton species) in marine biogeochemical processes.     

Multiple environmental controls on phytoplankton growth strategies determine adaptive responses of the N : P ratio

The controls on the ‘Redfield’ N : P stoichiometry of marine phytoplankton and hence the N : P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource‐allocation trade‐offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per‐ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N : P ratios and N : P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.     

Monitoring cellular C:N ratio in phytoplankton by means of FTIR‐spectroscopy

Statistical growth rate modelling can be applied in a variety of ecological and biotechnological applications. Such models are frequently based on Monod or Droop equations and, especially for the latter, require reliable determination of model input parameters such as C:N quotas. Besides growth rate modelling, a C:N quota quantification can be useful for monitoring and interpretation of physiological acclimation to abiotic and biotic disturbances (e.g., nutrient limitations). However, as high throughput C:N quota determination is difficult to perform, alternatives need to be established. Fourier‐transformed infrared (FTIR) spectroscopy is used to analyze a variety of biochemical, chemical, and physiological parameters in phytoplankton. Hence, a quantification of the C:N quota should also be feasible. Therefore, using FTIR spectroscopy, six phytoplankton species from among different phylogenetic groups have been analyzed to determine the effect of nutrient limitation on C:N quota patterns. The typical species‐specific response to increasing nitrogen limitation was an increase in the C:N quota. Irrespective of this species specificity, we were able to develop a reliable multi‐species C:N quota prediction model based on FTIR spectroscopy using the partial least square regression (PLSR) algorithm. Our data demonstrate that the PLSR approach is more robust in C:N quota quantification (R² = 0.93) than linear correlation of C:N quota versus growth rate (R² ranges from 0.74 to 0.86) or biochemical information based on FTIR spectra (R² ranges from 0.82 to 0.89). This accurate prediction of C:N values may support high throughput measurements in a broad range of future approaches.     

Marine nitrogen: Phosphorus stoichiometry and the global N:P cycle

Nitrogen supply is often assumed to limitmarine primary production. A global analysis of totalnitrogen (N) to phosphorus (P) molar ratios shows thattotal N:P is low (<16:1) in some estuarine andcoastal ecosystems, but up to 100:1 in open oceans.This implies that elements other than N may limitmarine production, except in human impacted, estuarineor coastal ecosystems. This pattern may reconcileconflicting enrichment studies, because N additionfrequently increases phytoplankton growth where totalN:P is expected to be low, but P, Fe, or Si augmentphytoplankton growth in waters where total N:P ishigh. Comparison of total N:P stoichiometry betweenmarine and freshwaters yields a model of the form ofthe aquatic N:P cycle.     

Multielement compositions of marine phytoplankton samples from coastal areas of japan by instrumental neutron activation analysis

Phytoplankton samples were collected during spring bloom of diatoms from three coastal areas of Japan using a NORPAC P-25 net (25-Μm opening) with a NGG52 prenet (335-Μm opening), and 25 major and trace elements have been analyzed by INAA. Concentration ranges of analyzed phytoplankton samples are much wider than the concentration ranges compiled by Bowen (1979) except for As, and data of marine phytoplankton samples for Br, Sb, Hf, Sc, La, Ce, Sm, and Eu were not included in the compilation. The 25 analyzed elements have been categorized into three groups: elements showing positive correlation with Br, positive correlation with Al, and no positive correlation with Br or Al. The marine phytoplankton samples have been plotted on a Masuzawa-Koyama-Terazaki (MKT) plot and it proved that the MKT plot is applicable to marine phytoplankton samples.     

Evolutionary temperature compensation of carbon fixation in marine phytoplankton

The efficiency of carbon sequestration by the biological pump could decline in the coming decades because respiration tends to increase more with temperature than photosynthesis. Despite these differences in the short‐term temperature sensitivities of photosynthesis and respiration, it remains unknown whether the long‐term impacts of global warming on metabolic rates of phytoplankton can be modulated by evolutionary adaptation. We found that respiration was consistently more temperature dependent than photosynthesis across 18 diverse marine phytoplankton, resulting in universal declines in the rate of carbon fixation with short‐term increases in temperature. Long‐term experimental evolution under high temperature reversed the short‐term stimulation of metabolic rates, resulting in increased rates of carbon fixation. Our findings suggest that thermal adaptation may therefore have an ameliorating impact on the efficiency of phytoplankton as primary mediators of the biological carbon pump.     

THE MESOZOIC RADIATION OF EUKARYOTIC ALGAE: THE PORTABLE PLASTID HYPOTHESIS1

Although all chloroplasts appear to have been derived from a common ancestor, a major schism occurred early in the evolution of eukaryotic algae that gave rise to red and green photoautotrophic lineages. In Paleozoic and earlier times, the fossil record suggests that oceanic eukaryotic phytoplankton were dominated by the green (chl b‐containing) algal line. However, following the end‐Permian extinction, a diverse group of eukaryotic phytoplankton evolved from secondary symbiotic associations in the red (chl c‐containing) line and subsequently rose to ecological prominence. In the contemporary oceans, red eukaryotic phytoplankton taxa continue to dominate marine pelagic food webs, whereas the green line is relegated to comparatively minor ecological and biogeochemical roles. To help elucidate why the oceans are not dominated by green taxa, we analyzed and compared whole plastid genomes in both the red and green lineages. Our results suggest that whereas all algal plastids retain a core set of genes, red plastids retain a complementary set of genes that potentially confer more capacity to autonomously express proteins regulating oxygenic photosynthetic and energy transduction pathways. We hypothesize that specific gene losses in the primary endosymbiotic green plastid reduced its portability for subsequent symbiotic associations. This corollary of the plastid “enslavement” hypothesis may have limited subsequent evolutionary advances in the green lineage while simultaneously providing a competitive advantage to the red lineage.     

Nitrogen limitation in the tropical waters of the Bay of Bengal

A 3 year study (1986–1989) was carried out in the Bay of Bengal off Madras in order to understand the influence of physical and chemical variables on the occurrence, abundance and productivity of its phytoplankton. Biochemical oxygen demand and nutrient concentrations were highest near the mouth of river Cooum. Pigments and net primary production in nearshore waters varied between 4.1 and 1113 C mg m^–3 h^–1, while in offshore waters the maximum was only 201 mg C m^–3 h^–1. Multiple regression analysis with net photosynthesis as the dependent variable and other variates as the independent variables revealed that nutrients did not account for much variation in net photosynthesis in nearshore stations but contributed significantly to variation in offshore station. Analyses of seawater collected during two cruises in coastal waters at 18–22° N latitude revealed that nitrogen was low in comparison to phosphorus and could be limiting primary production in the surface waters of the Bay.     

Carbon allocation under light and nitrogen resource gradients in two model marine phytoplankton1

Marine phytoplankton have conserved elemental stoichiometry, but there can be significant deviations from this Redfield ratio. Moreover, phytoplankton allocate reduced carbon (C) to different biochemical pools based on nutritional status and light availability, adding complexity to this relationship. This allocation influences physiology, ecology, and biogeochemistry. Here, we present results on the physiological and biochemical properties of two evolutionarily distinct model marine phytoplankton, a diatom (cf. Staurosira sp. Ehrenberg) and a chlorophyte (Chlorella sp. M. Beijerinck) grown under light and nitrogen resource gradients to characterize how carbon is allocated under different energy and substrate conditions. We found that nitrogen (N)‐replete growth rate increased monotonically with light until it reached a threshold intensity (~200 μmol photons · m^?2 · s^?1). For Chlorella sp., the nitrogen quota (pg · μm^?3) was greatest below this threshold, beyond which it was reduced by the effect of N‐stress, while for Staurosira sp. there was no trend. Both species maintained constant maximum quantum yield of photosynthesis (mol C · mol photons^?1) over the range of light and N‐gradients studied (although each species used different photophysiological strategies). In both species, C:chl a (g · g^?1) increased as a function of light and N‐stress, while C:N (mol · mol^?1) and relative neutral lipid:C (rel. lipid · g^?1) were most strongly influenced by N‐stress above the threshold light intensity. These results demonstrated that the interaction of substrate (N‐availability) and energy gradients influenced C‐allocation, and that general patterns of biochemical responses may be conserved among phytoplankton; they provided a framework for predicting phytoplankton biochemical composition in ecological, biogeochemical, or biotechnological applications.     

Rapid evolution of metabolic traits explains thermal adaptation in phytoplankton

Understanding the mechanisms that determine how phytoplankton adapt to warming will substantially improve the realism of models describing ecological and biogeochemical effects of climate change. Here, we quantify the evolution of elevated thermal tolerance in the phytoplankton, Chlorella vulgaris. Initially, population growth was limited at higher temperatures because respiration was more sensitive to temperature than photosynthesis meaning less carbon was available for growth. Tolerance to high temperature evolved after ≈ 100 generations via greater down‐regulation of respiration relative to photosynthesis. By down‐regulating respiration, phytoplankton overcame the metabolic constraint imposed by the greater temperature sensitivity of respiration and more efficiently allocated fixed carbon to growth. Rapid evolution of carbon‐use efficiency provides a potentially general mechanism for thermal adaptation in phytoplankton and implies that evolutionary responses in phytoplankton will modify biogeochemical cycles and hence food web structure and function under warming. Models of climate futures that ignore adaptation would usefully be revisited.     

中亚热带不同森林更新方式生态酶化学计量特征

了解土壤生态化学计量特征对预测不同生态系统养分变化、功能以及植物生产力具有重要意义。森林更新是维持中亚热带森林生态系统可持续发展的重要手段。选取福建省三明市陈大林业采育场3种不同森林更新方式进行研究,包括米槠次生林（SF）、米槠人工促进天然更新林（AR）和杉木人工林（CF）,测定其土壤理化性质及土壤3种酶,计算酶化学计量。结果显示：1）AR的土壤总氮、全磷、铵态氮含量以及含水量最高（P < 0.05）,土壤有效磷的含量则是CF最高（P < 0.05）;2）生态酶化学计量结果表明AR的土壤微生物处于氮限制状态,CF的土壤微生物处于磷限制状态;3）冗余分析表明,土壤含水量和铵态氮是驱动不同森林更新方式土壤生态酶化学计量变异的重要环境因子。研究表明,人促更新方式更有利于土壤有效氮的积累,而人工林则有利于土壤有效磷积累,这可能与造林树种有关。土壤生态酶化学计量更易受到土壤含水量以及有效性养分的驱动,而与土壤化学计量未呈现良好的耦合性。