Predicting invasion in grassland ecosystems: is exotic dominance the real embarrassment of richness?

Invasions have increased the size of regional species pools, but are typically assumed to reduce native diversity. However, global-scale tests of this assumption have been elusive because of the focus on exotic species richness, rather than relative abundance. This is problematic because low invader richness can indicate invasion resistance by the native community or, alternatively, dominance by a single exotic species. Here, we used a globally replicated study to quantify relationships between exotic richness and abundance in grass-dominated ecosystems in 13 countries on six continents, ranging from salt marshes to alpine tundra. We tested effects of human land use, native community diversity, herbivore pressure, and nutrient limitation on exotic plant dominance. Despite its widespread use, exotic richness was a poor proxy for exotic dominance at low exotic richness, because sites that contained few exotic species ranged from relatively pristine (low exotic richness and cover) to almost completely exotic-dominated ones (low exotic richness but high exotic cover). Both exotic cover and richness were predicted by native plant diversity (native grass richness) and land use (distance to cultivation). Although climate was important for predicting both exotic cover and richness, climatic factors predicting cover (precipitation variability) differed from those predicting richness (maximum temperature and mean temperature in the wettest quarter). Herbivory and nutrient limitation did not predict exotic richness or cover. Exotic dominance was greatest in areas with low native grass richness at the site- or regional-scale. Although this could reflect native grass displacement, a lack of biotic resistance is a more likely explanation, given that grasses comprise the most aggressive invaders. These findings underscore the need to move beyond richness as a surrogate for the extent of invasion, because this metric confounds monodominance with invasion resistance. Monitoring species' relative abundance will more rapidly advance our understanding of invasions.

Model-based analysis and optimization of bioreactor for hematopoietic stem cell cultivation

One of the problems to be solved in attaining the full potentials of hematopoietic stem cell (HSC) applications is the limited availability of the cells. Growing HSCs in a bioreactor offers an alternative solution to this problem. Besides, it also offers the advantages of eliminating labour intensive process as well as the possible contamination involved in the periodic nutrient replenishments in the traditional T-flask stem cell cultivation. In spite of this, the optimization of HSC cultivation in a bioreactor has been barely explored. This manuscript discusses the development of a mathematical model to describe the dynamics in nutrient distribution and cell concentration of an ex vivo HSC cultivation in a microchannel perfusion bioreactor. The model was further used to optimize the cultivation by proposing three alternative feeding strategies in order to prevent the occurrence of nutrient limitation in the bioreactor. The evaluation of these strategies, the periodic step change increase in the inlet oxygen concentration, the periodic step change increase in the media inflow, and the feedback control of media inflow, shows that these strategies can successfully improve the cell yield of the bioreactor. In general, the developed model is useful for the design and optimization of bioreactor operation.

Grassland productivity limited by multiple nutrients

Terrestrial ecosystem productivity is widely accepted to be nutrient limited1. Although nitrogen (N) is deemed a key determinant of aboveground net primary production (ANPP)2,3, the prevalence of co-limitation by N and phosphorus (P) is increasingly recognized4,​5,​6,​7,​8. However, the extent to which terrestrial productivity is co-limited by nutrients other than N and P has remained unclear. Here, we report results from a standardized factorial nutrient addition experiment, in which we added N, P and potassium (K) combined with a selection of micronutrients (K+μ), alone or in concert, to 42 grassland sites spanning five continents, and monitored ANPP. Nutrient availability limited productivity at 31 of the 42 grassland sites. And pairwise combinations of N, P, and K+μ co-limited ANPP at 29 of the sites. Nitrogen limitation peaked in cool, high latitude sites. Our findings highlight the importance of less studied nutrients, such as K and micronutrients, for grassland productivity, and point to significant variations in the type and degree of nutrient limitation. We suggest that multiple-nutrient constraints must be considered when assessing the ecosystem-scale consequences of nutrient enrichment.

Vertical Distribution of Soil Denitrifying Communities in a Wet Sclerophyll Forest under Long-Term Repeated Burning

Soil biogeochemical cycles are largely mediated by microorganisms, while fire significantly modifies biogeochemical cycles mainly via altering microbial community and substrate availability. Majority of studies on fire effects have focused on the surface soil; therefore, our understanding of the vertical distribution of microbial communities and the impacts of fire on nitrogen (N) dynamics in the soil profile is limited. Here, we examined the changes of soil denitrification capacity (DNC) and denitrifying communities with depth under different burning regimes, and their interaction with environmental gradients along the soil profile. Results showed that soil depth had a more pronounced impact than the burning treatment on the bacterial community size. The abundance of 16S rRNA and denitrification genes (narG, nirK, and nirS) declined exponentially with soil depth. Surprisingly, the nosZ-harboring denitrifiers were enriched in the deeper soil layers, which was likely to indicate that the nosZ-harboring denitrifiers could better adapt to the stress conditions (i.e., oxygen deficiency, nutrient limitation, etc.) than other denitrifiers. Soil nutrients, including dissolved organic carbon (DOC), total soluble N (TSN), ammonium (NH4 +), and nitrate (NO3 −), declined significantly with soil depth, which probably contributed to the vertical distribution of denitrifying communities. Soil DNC decreased significantly with soil depth, which was negligible in the depths below 20 cm. These findings have provided new insights into niche separation of the N-cycling functional guilds along the soil profile, under a varied fire disturbance regime.

Fluorescence properties of Baltic Sea phytoplankton

To obtain data on phytoplankton dynamics with improved spatial and temporal resolution, and at reduced cost, traditional phytoplankton monitoring methods have been supplemented with optical approaches. In this thesis, I have explored various fluorescence-based techniques for detection of phytoplankton abundance, taxonomy and physiology in the Baltic Sea. In algal cultures used in this thesis, the availability of nitrogen and light conditions caused changes in pigmentation, and consequently in light absorption and fluorescence properties of cells. In the Baltic Sea, physical environmental factors (e.g. mixing depth, irradiance and temperature) and related seasonal succession in the phytoplankton community explained a large part of the seasonal variability in the magnitude and shape of Chlorophyll a (Chla)-specific absorption. The variability in Chla-specific fluorescence was related to the abundance of cyanobacteria, the size structure of the phytoplankton community, and absorption characteristics of phytoplankton. Cyanobacteria show very low Chla-specific fluorescence. In the presence of eukaryotic species, Chla fluorescence describes poorly cyanobacteria. During cyanobacterial bloom in the Baltic Sea, phycocyanin fluorescence explained large part of the variability in Chla concentrations. Thus, both Chla and phycocyanin fluorescence were required to predict Chla concentration. Phycobilins are major light harvesting pigments for cyanobacteria. In the open Baltic Sea, small picoplanktonic cyanobacteria were the main source of phycoerythrin fluorescence and absorption signal. Large filamentous cyanobacteria, forming harmful blooms, were the main source of the phycocyanin fluorescence signal and typically their biomass and phycocyanin fluorescence were linearly related. Using phycocyanin fluorescence, dynamics of cyanobacterial blooms can be detected at high spatial and seasonal resolution not possible with other methods. Various taxonomic phytoplankton pigment groups can be separated by spectral fluorescence. I compared multivariate calibration methods for the retrieval of phytoplankton biomass in different taxonomic groups. Partial least squares regression method gave the closest predictions for all taxonomic groups, and the accuracy was adequate for phytoplankton bloom detection. Variable fluorescence has been proposed as a tool to study the physiological state of phytoplankton. My results from the Baltic Sea emphasize that variable fluorescence alone cannot be used to detect nutrient limitation of phytoplankton. However, when combined with experiments with active nutrient manipulation, and other nutrient limitation indices, variable fluorescence provided valuable information on the physiological responses of the phytoplankton community. This thesis found a severe limitation of a commercial fast repetition rate fluorometer, which couldn t detect the variable fluorescence of phycoerythrin-lacking cyanobacteria. For these species, the Photosystem II absorption of blue light is very low, and fluorometer excitation light did not saturate Photosystem II during a measurement. This thesis encourages the use of various in vivo fluorescence methods for the detection of bulk phytoplankton biomass, biomass of cyanobacteria, chemotaxonomy of phytoplankton community, and phytoplankton physiology. Fluorescence methods can support traditional phytoplankton monitoring by providing continuous measurements of phytoplankton, and thereby strengthen the understanding of the links between biological, chemical and physical processes in aquatic ecosystems.

Effects of temperature on the threshold of phosphorus for algal blooms

Algal blooms, worsening marine ecosystems and causing great economic loss, have been paid much attention to for a long time. Such environmental factors as light penetration, water temperature, and nutrient concentration are crucial in blooms processes. Among them, only nutrients can be controlled. Therefore, the threshold of nutrients for algal blooms is of great concern. To begin with, a dynamic eutrophication model has been constructed to simulate the algal growth and phosphorus cycling. The model encapsulates the essential biological processes of algal growth and decay, and phosphorus regeneration due to algal decay. The nutrient limitation is based upon commonly used Monod's kinetics. The effects of temperature and phosphorus limitation are particularly addressed. Then, we have endeavored to elucidate the threshold of phosphorus at different temperature for algal blooms. Based on the numerical simulation, the isoquant contours of change rate of alga as shown in the figure are obtained, which obviously demonstrate the threshold of nutrient at an arbitrary reasonable temperature. The larger the change rate is, the more rapidly the alga grows. If the phosphorus concentration at a given temperature remains larger than the threshold the algal biomass may increase monotonically, leading to the algal blooming. With the rising of temperature, the threshold is apparently reduced, which may explain why likely red tide disasters occur in a fine summer day. So, high temperature and sufficient phosphorus supply are the major factors which result in algal growth and blowout of red tide.

Variação temporal de uma comunidade fitoplanctônica do reservatório de APM-Manso através de modelagem ecológica tridimensional.

A ecologia de reservatórios, que são ecossistemas complexos, dinâmicos e artificiais, vem assumindo destaque no Brasil. O objetivo deste trabalho foi avaliar a viabilidade da aplicação, no reservatório de APM-Manso, de um modelo ecológico tridimensional em estudos sobre a dinâmica fitoplanctônica, simulando a variação temporal do fitoplâncton para cenários distintos de carga de nutrientes. O modelo CAEDYM foi acoplado ao ELCOM e simulação foi realizada em duas etapas: uma hidrodinâmica e outra ecológica. Escolheu-se para as simulações o período de cinco meses, a partir de 1 de setembro de 2005. Foram construídos dois cenários de simulação, o primeiro contendo os valores reais de carga de nutrientes dos principais rios contribuintes medidos em campo, e o segundo com redução na carga nutricional destes rios, simulando um possível processo de substituição de áreas florestadas por áreas de pastagem na bacia do rio Manso. A comunidade fitoplanctônica simulada apresentou rápidas respostas à disponibilidade nutricional do ambiente, e os resultados obtidos corroboraram com diversas teorias sobre as estratégias adaptativas e sobre as dinâmicas algais. Dentre as classes simuladas, Bacillariophyceae e Cryptophyceae se mostraram mais sensíveis às reduções de carga, enquanto Chrolophyceae e Cyanophyceae, apesar de terem suas biomassas reduzidas, sofreram menos com o impacto, sugerindo estarem mais adaptadas à limitação de nutrientes. Os picos chuvosos influenciaram positivamente as taxas de crescimento das Bacillariophyceae apenas no Cenário 1, uma vez que a limitação por nutrientes foi mais decisiva para esta classe no Cenário 2. Observou-se em ambas as simulações uma tendência de substituição na dominância de Cyanophyceae por Chlorophyceae.

Increased toxicity of Karenia brevis during phosphate limited growth: ecological and evolutionary implications

Karenia brevis is the dominant toxic red tide algal species in the Gulf of Mexico. It produces potent neurotoxins (brevetoxins [PbTxs]), which negatively impact human and animal health, local economies, and ecosystem function. Field measurements have shown that cellular brevetoxin contents vary from 1–68 pg/cell but the source of this variability is uncertain. Increases in cellular toxicity caused by nutrient-limitation and inter-strain differences have been observed in many algal species. This study examined the effect of P-limitation of growth rate on cellular toxin concentrations in five Karenia brevis strains from different geographic locations. Phosphorous was selected because of evidence for regional P-limitation of algal growth in the Gulf of Mexico. Depending on the isolate, P-limited cells had 2.3- to 7.3-fold higher PbTx per cell than P-replete cells. The percent of cellular carbon associated with brevetoxins (%C-PbTx) was ~ 0.7 to 2.1% in P-replete cells, but increased to 1.6–5% under P-limitation. Because PbTxs are potent anti-grazing compounds, this increased investment in PbTxs should enhance cellular survival during periods of nutrient-limited growth. The %C-PbTx was inversely related to the specific growth rate in both the nutrient-replete and P-limited cultures of all strains. This inverse relationship is consistent with an evolutionary tradeoff between carbon investment in PbTxs and other grazing defenses, and C investment in growth and reproduction. In aquatic environments where nutrient supply and grazing pressure often vary on different temporal and spatial scales, this tradeoff would be selectively advantageous as it would result in increased net population growth rates. The variation in PbTx/cell values observed in this study can account for the range of values observed in the field, including the highest values, which are not observed under N-limitation. These results suggest P-limitation is an important factor regulating cellular toxicity and adverse impacts during at least some K. brevis blooms.

Forecasting Daily Chlorophyll a Concentration during the Spring Phytoplankton Bloom Period in Xiangxi Bay of the Three-Gorges Reservoir by Means of a Recurrent Artificial Neural Network

A recurrent artificial neural network was used for 0-and 7-days-ahead forecasting of daily spring phytoplankton bloom dynamics in Xiangxi Bay of Three-Gorges Reservoir with meteorological, hydrological, and limnological parameters as input variables. Daily data from the depth of 0.5 m was used to train the model, and data from the depth of 2.0 m was used to validate the calibrated model. The trained model achieved reasonable accuracy in predicting the daily dynamics of chlorophyll a both in 0-and 7-days-ahead forecasting. In 0-day-ahead forecasting, the R-2 values of observed and predicted data were 0.85 for training and 0.89 for validating. In 7-days-ahead forecasting, the R-2 values of training and validating were 0.68 and 0.66, respectively. Sensitivity analysis indicated that most ecological relationships between chlorophyll a and input environmental variables in 0-and 7-days-ahead models were reasonable. In the 0-day model, Secchi depth, water temperature, and dissolved silicate were the most important factors influencing the daily dynamics of chlorophyll a. And in 7-days-ahead predicting model, chlorophyll a was sensitive to most environmental variables except water level, DO, and NH3N.

Effects of macroalgae Ulva pertusa (Chlorophyta) and Gracilaria lemaneiformis (Rhodophyta) on growth of four species of bloom-forming dinoflagellates

The effects of fresh thalli and culture medium filtrates from two species of marine macroalgae, Ulva pertusa Kjellm (Chlorophyta) and Gracilaria lemaneiformis (Bory) Dawson (Rhodophyta), on growth of marine microalgae were investigated in co-culture under controlled laboratory conditions. A selection of microalgal species were used, all, being identified as bloom-forming dinoflagellates: Prorocentrum donghaiense Lu sp., Alexandrium tamarense (Lebour) Balech, Amphidinium carterae Hulburt and Scrippsiella trochoide (Stein) Loeblich III. Results showed that the fresh thalli of either U. pertusa or G. lemaneiformis significantly inhibited the microalgal growth, or caused mortality at the end of the experiment. However, the overall effects of the macroalgal culture filtrates on the growth of the dinoflagellates were species-specific (inhibitory, stimulatory or none) for different microalgal species. Results indicated an allelopathic effect of macroalga on the co-cultured dinoflagellate. We then took P. donghaiense as an example to further assess this hypothesis. The present study was carried out under controlled conditions, thereby excluded the fluctuation in light and temperature. Nutrient assays showed that nitrate and phosphate were almost exhausted in G. lemaneiformis co-culture. but remained at enough high levels in U pertusa co-culture, which were well above the nutrient limitation for the microalgal growth, when all cells of P. donghaiense were killed in the co-culture. Daily f/2 medium enrichment greatly alleviated the growth inhibition on P. donghaiense in G. lemaneiformis co-culture, but could not eliminate it. Other environmental factors, such as carbonate limitation, bacterial presence and the change of pH were also not necessary for the results. We thus concluded that allelopathy was the most possible reason leading to the negative effect of U. pertusa on P. donghaiense, and the combined roles of allelopathy and nutrient competition were essential for the effect of G. lemaneiformis on P. donghaiense. (c) 2006 Elsevier B.V. All rights reserved.

Continuous Plankton Records - Seasonal-Variations In The Distribution And Abundance Of Plankton In The North-Atlantic Ocean And The North-Sea

Charts are presented of the seasonal variations in the distribution of four phytoplankton and five zooplankton taxa in the North Atlantic and the North Sea. The main factors determining the seasonal variations appear to be the distribution of the main overwintering stocks, the current system and, in some instances, temperature control of the rate of population increase. Information is presented about the variation with latitude (over the range from 34° N to 65 ° N) of the seasonal regime of the plankton. On the assumption that there is a relationship between nutrient supply and vertical temperature stratification the main features of this variability can be interpreted. In the south (to about 43° N) nutrient limitation plus grazing appear to be dominant, resulting in a bimodal seasonal cycle of phytoplankton. North of about 60° N the system appears to be limited by the size of the phytoplankton stocks being grazed primarily by Calanus Finmarchicus and Euphausiacea. In an extensive zone, from about 44° N to 60° N, it would appear that the spring bloom of phytoplankton is under-exploited by grazing while in summer the zooplankton graze the daily production of the phytoplankton, the stocks of which are probably maintained by in situ nutrient regeneration. The implications, for at least this mid-latitude zone, that rates and fluxes of processes, as opposed to density dependent interactions between stocks, play a major role in the dynamics of the seasonal cycle is consistent with previously reported observations suggesting that physical environmental factors play a major role in determining year-to-year fluctuations in the abundance of the plankton.

Identification of senescence and death in Emiliania huxleyi and Thalassiosira pseudonana: Cell staining chlorophyll alterations and dimethylsulfoniopropionate (DMSP) metabolism

We measured membrane permeability, hydrolytic enzyme, and caspase-like activities using fluorescent cell stains to document changes caused by nutrient exhaustion in the coccolithophore Emiliania huxleyi and the diatom Thalassiosira pseudonana, during batch-culture nutrient limitation. We related these changes to cell death, pigment alteration, and concentrations of dimethylsulfide (DMS) and dimethylsulfoniopropionate (DMSP) to assess the transformation of these compounds as cell physiological condition changes. E. huxleyi persisted for 1 month in stationary phase; in contrast, T. pseudonana cells rapidly declined within 10 d of nutrient depletion. T. pseudonana progressively lost membrane integrity and the ability to metabolize 5-chloromethylfluorescein diacetate (CMFDA; hydrolytic activity), whereas E. huxleyi developed two distinct CMFDA populations and retained membrane integrity (SYTOX Green). Caspase-like activity appeared higher in E. huxleyi than in T. pseudonana during the post-growth phase, despite a lack of apparent mortality and cell lysis. Photosynthetic pigment degradation and transformation occurred in both species after growth; chlorophyll a (Chl a) degradation was characterized by an increase in the ratio of methoxy Chl a : Chl a in T. pseudonana but not in E. huxleyi, and the increase in this ratio preceded loss of membrane integrity. Total DMSP declined in T. pseudonana during cell death and DMS increased. In contrast, and in the absence of cell death, total DMSP and DMS increased in E. huxleyi. Our data show a novel chlorophyll alteration product associated with T. pseudonana death, suggesting a promising approach to discriminate nonviable cells in nature.

A mechanistic explanation of the Sargasso Sea DMS summer paradox

In the Sargasso Sea, maximum dimethylsulfide (DMS) accumulation occurs in summer, concomitant with the minimum of chlorophyll and 2 months later than its precursor, dimethylsulfoniopropionate (DMSP). This phenomenon is often referred to as the DMS "summer paradox". It has been previously suggested that the main agent triggering this pattern is increasing irradiance leading to light stress-induced DMS release from phytoplankton cells. We have developed a new model describing DMS(P) dynamics in the water column and used it to investigate how and to what extent processes other than light induced DMS exudation from phytoplankton, may contribute to the DMS summer paradox. To do this, we have conceptually divided the DMS "summer paradox" into two components: (1) the temporal decoupling between chlorophyll and DMSP and (2) the temporal decoupling between DMSP and DMS. Our results suggest that it is possible to explain the above cited patterns by means of two different dynamics, respectively: (1) a succession of phytoplankton types in the surface water and (2) the bacterially mediated DMSP(d) to DMS conversion, seasonally varying as a function of nutrient limitation. This work differs from previous modelling studies in that the presented model suggests that phytoplankton light-stress induced processes may only partially explain the summer paradox, not being able to explain the decoupling between DMSP and DMS, which is possibly the more challenging aspect of this phenomenon. Our study, therefore, provides an "alternative" explanation to the summer paradox further underlining the major role that bacteria potentially play in DMS production and fate.

On the roles of cell size and trophic strategy in North Atlantic diatom and dinoflagellate communities

We have examined the inter- and intra-group seasonal succession of 113 diatom and dinoflagellate taxa, as surveyed by the Continuous Plankton Recorder (CPR) in the North Atlantic, by grouping taxa according to two key functional traits: cell size (mg C cell21) and trophic strategy (photoautotrophy, mixotrophy, or heterotrophy). Mixotrophic dinoflagellates follow photoautotrophic diatoms but precede their obligate heterotrophic counterparts in the succession because of the relative advantages afforded by photosynthesizing when light and nutrients are available in spring. The mean cell size of the sampled diatoms is smallest in the summer, likely because of the higher specific nutrient affinity of smaller relative to larger cells. Contrastingly, we hypothesize that mixotrophy diminishes the size selection based on nutrient limitation and accounts for the lack of a seasonal size shift among surveyed dinoflagellates. Relatively small, heterotrophic dinoflagellates (mg C cell21 , 1023) peak after other, larger dinoflagellates, in part because of the increased abundance of their small prey during nutrientdeplete summer months. The largest surveyed diatoms (mg C cell21 . 1022) bloom later than others, and we hypothesize that this may be because of their relatively slow maximum potential growth rates and high internal nutrient storage, as well as to the slower predation of these larger cells. The new trait database and analysis presented here helps translate the taxonomic information of the CPR survey into metrics that can be directly compared with trait-based models.

Flexible C : N ratio enhances metabolism of large phytoplankton when resource supply is intermittent

Phytoplankton cell size influences particle sinking rate, food web interactions and biogeographical distributions. We present a model in which the uptake, storage and assimilation of nitrogen and carbon are explicitly resolved in different-sized phytoplankton cells. In the model, metabolism and cellular C :N ratio are influenced by the accumulation of carbon polymers such as carbohydrate and lipid, which is greatest when cells are nutrient starved, or exposed to high light. Allometric relations and empirical data sets are used to constrain the range of possible C : N, and indicate that larger cells can accumulate significantly more carbon storage compounds than smaller cells. When forced with extended periods of darkness combined with brief exposure to saturating irradiance, the model predicts organisms large enough to accumulate significant carbon reserves may on average synthesize protein and other functional apparatus up to five times faster than smaller organisms. The advantage of storage in terms of average daily protein synthesis rate is greatest when modeled organisms were previously nutrient starved, and carbon storage reservoirs saturated. Small organisms may therefore be at a disadvantage in terms of average daily growth rate in environments that involve prolonged periods of darkness and intermittent nutrient limitation. We suggest this mechanism is a significant constraint on phytoplankton C :N variability and cell size distribution in different oceanic regimes.