Climate change and the timing, magnitude, and composition of the phytoplankton spring bloom

In this article, we show by mesocosm experiments that winter and spring warming will lead to substantial changes in the spring bloom of phytoplankton. The timing of the spring bloom shows only little response to warming as such, while light appears to play a more important role in its initiation. The daily light dose needed for the start of the phytoplankton spring bloom in our experiments agrees well with a recently published critical light intensity found in a field survey of the North Atlantic (around 1.3 mol photons m^?2 day^?1). Experimental temperature elevation had a strong effect on phytoplankton peak biomass (decreasing with temperature), mean cell size (decreasing with temperature) and on the share of microplankton diatoms (decreasing with temperature). All these changes will lead to poorer feeding conditions for copepod zooplankton and, thus, to a less efficient energy transfer from primary to fish production under a warmer climate.     

Earlier onset of the spring phytoplankton bloom in lakes of the temperate zone in a warmer climate

The decoupling of trophic interactions is potentially one of the most severe consequences of climate warming. In lakes and oceans the timing of phytoplankton blooms affects competition within the plankton community as well as food–web interactions with zooplankton and fish. Using Upper Lake Constance as an example, we present a model‐based analysis that predicts that in a future warmer climate, the onset of the spring phytoplankton bloom will occur earlier in the year than it does at present. This is a result of the earlier occurrence of the transition from strong to weak vertical mixing in spring, and of the associated earlier onset of stratification. According to our simulations a shift in the timing of phytoplankton growth resulting from a consistently warmer climate will exceed that resulting from a single unusually warm year. The numerical simulations are complemented by a statistical analysis of long‐term data from Upper Lake Constance which demonstrates that oligotrophication has a negligible effect on the timing of phytoplankton growth in spring and that an early onset of the spring phytoplankton bloom is associated with high air temperatures and low wind speeds.     

Warming promotes cold-adapted phytoplankton in temperate lakes and opens a loophole for Oscillatoriales in spring

The effects of the recent warming trend in many northern temperate lakes on the species composition of spring phytoplankton remain poorly understood, especially because a recent change in nutrients has complicated efforts, and previous studies have defined spring according to the calendar. We analysed data from 1979 to 2004 from Lake Müggelsee (Berlin, Germany), using physical and biological parameters to define the spring period. We show that a change in timing of spring plankton events in warm years led to the paradox of lower mean water temperatures during the growth period, favouring cold-adapted diatoms over cyanobacteria, and within the diatoms, some cold-adapted centric forms over pennate forms. Under high P : Si ratios, the increased time between phytoplankton and cladoceran peaks opened a loophole for filamentous cyanobacteria (Oscillatoriales) in warm years to establish dominance after the diatoms, which are silicate limited. Therefore, the warming trend promotes filamentous cyanobacteria, a well-known nuisance in eutrophic lakes, and surprisingly, cold-adapted diatoms.     

Interaction effects of zooplankton and CO2 on phytoplankton communities and the deep chlorophyll maximum

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Climate warming increases spring phenological differences among temperate trees

Climate warming has substantially advanced spring leaf flushing, but winter chilling and photoperiod co‐determine the leaf flushing process in ways that vary among species. As a result, the interspecific differences in spring phenology (IDSP) are expected to change with climate warming, which may, in turn, induce negative or positive ecological consequences. However, the temporal change of IDSP at large spatiotemporal scales remains unclear. In this study, we analyzed long‐term in‐situ observations (1951–2016) of six, coexisting temperate tree species from 305 sites across Central Europe and found that phenological ranking did not change when comparing the rapidly warming period 1984–2016 to the marginally warming period 1951–1983. However, the advance of leaf flushing was significantly larger in early‐flushing species EFS (6.7 ± 0.3 days) than in late‐flushing species LFS (5.9 ± 0.2 days) between the two periods, indicating extended IDSP. This IDSP extension could not be explained by differences in temperature sensitivity between EFS and LFS; however, climatic warming‐induced heat accumulation effects on leaf flushing, which were linked to a greater heat requirement and higher photoperiod sensitivity in LFS, drove the shifts in IDSP. Continued climate warming is expected to further extend IDSP across temperate trees, with associated implications for ecosystem function.     

瓯江口春季营养盐、浮游植物和浮游动物的分布

研究2007年4月瓯江口海域(27°38′～28°02′N、120°50′～121°14′E)浮游生物空间分布特征,分析这一分布特征与营养盐和其他水文要素之间的联系.结果表明,悬浮物浓度、DIP和DIN分布特征均是近河口最高,由近河口向外数量逐渐减少.在灵昆岛南侧和东南侧近口门水域,是DIP和DIN的高值中心,但该水域高浓度悬沙使水体透光率较低,不利于浮游植物生长.在口门外侧海域,悬沙浓度已经明显降低,因而是浮游植物高密度区域.浮游植物和浮游动物丰度分布趋势基本相同,由内海向外海数量逐渐递减;其中,大型浮游动物丰度和浮游植物丰度,小型浮游动物丰度和浮游植物丰度之间显著地呈线性正相关关系.这一特征的形成,主要由瓯江口营养盐和悬浮物分布特征决定,浮游动物与浮游植物空间分布的一致性,很好地反映出浮游动物对浮游植物有效的下行控制,从而使水域生态系统在此季节保持稳定.     

Phytoplankton diversity loss along a gradient of future warming and brownification in freshwater mesocosms

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Spring bloom succession,grazing impact and herbivore selectivity of ciliate communities in response to winter warming

This study aimed at simulating different degrees of winter warming and at assessing its potential effects on ciliate succession and grazing-related patterns. By using indoor mesocosms filled with unfiltered water from Kiel Bight, natural light and four different temperature regimes, phytoplankton spring blooms were induced and the thermal responses of ciliates were quantified. Two distinct ciliate assemblages, a pre-spring and a spring bloom assemblage, could be detected, while their formation was strongly temperature-dependent. Both assemblages were dominated by Strobilidiids; the pre-spring bloom phase was dominated by the small Strobilidiids Lohmaniella oviformis, and the spring bloom was mainly dominated by large Strobilidiids of the genus Strobilidium. The numerical response of ciliates to increasing food concentrations showed a strong acceleration by temperature. Grazing rates of ciliates and copepods were low during the pre-spring bloom period and high during the bloom ranging from 0.06 (Δ0°C) to 0.23 day⁻¹ (Δ4°C) for ciliates and 0.09 (Δ0°C) to 1.62 day⁻¹ (Δ4°C) for copepods. During the spring bloom ciliates and copepods showed a strong dietary overlap characterized by a wide food spectrum consisting mainly of Chrysochromulina sp., diatom chains and large, single-celled diatoms. Priority programme of the German Research Foundation—contribution 4.     

Water temperature and stratification depth independently shift cardinal events during plankton spring succession

In deep temperate lakes, the beginning of the growing season is triggered by thermal stratification, which alleviates light limitation of planktonic producers in the surface layer and prevents heat loss to deeper strata. The sequence of subsequent phenological events (phytoplankton spring bloom, grazer peak, clearwater phase) results in part from coupled phytoplankton–grazer interactions. Disentangling the separate, direct effects of correlated climatic drivers (stratification‐dependent underwater light climate vs. water temperature) from their indirect effects mediated through trophic feedbacks is impossible using observational field data, which challenges our understanding of global warming effects on seasonal plankton dynamics. We therefore manipulated water temperature and stratification depth independently in experimental field mesocosms containing ambient microplankton and inocula of the resident grazer Daphnia hyalina. Higher light availability in shallower surface layers accelerated primary production, warming accelerated consumption and growth of Daphnia, and both factors speeded up successional dynamics driven by trophic feedbacks. Specifically, phytoplankton peaked and decreased earlier and Daphnia populations increased and peaked earlier at both shallower stratification and higher temperature. The timing of ciliate dynamics was unrelated to both factors. Volumetric peak densities of phytoplankton, ciliates and Daphnia in the surface layer were also unaffected by temperature but declined with stratification depth in parallel with light availability. The latter relationship vanished, however, when population sizes were integrated over the entire water column. Overall our results suggest that, integrated over the entire water column of a deep lake, surface warming and shallower stratification independently speed up spring successional events, whereas the magnitudes of phytoplankton and zooplankton spring peaks are less sensitive to these factors. Therefore, accelerated dynamics under warming need not lead to a trophic mismatch (given similar grazer inocula at the time of stratification). We emphasize that entire water column dynamics must be studied to estimate global warming effects on lake ecosystems.     

Predicted warming and browning affect timing and magnitude of plankton phenological events in lakes: a mesocosm study

1. Aquatic ecosystems in Northern Europe are expected to face increases in temperature and water colour (TB) in future. While effects of these factors have been studied separately, it is unknown whether and how a combination of them might affect phenological events and trophic interactions. 2. In a mesocosm study, we combined both factors to create conditions expected to arise during the coming century. We focused on quantifying effects on timing and magnitude of plankton spring phenological events and identifying possible mismatches between resources (phytoplankton) and consumers (zooplankton). 3. We found that the increases in TB had important effects on timing and abundance of different plankton groups. While increased temperature led to an earlier peak in phytoplankton and zooplankton and a change in the relative timing of different zooplankton groups, increased water colour reduced chlorophyll‐a concentrations. 4. Increased TB together benefitted cladocerans and calanoid copepods and led to stronger top‐down control of algae by zooplankton. There was no sign of a mismatch between primary producers and grazers as reported from other studies. 5. Our results point towards an earlier onset of plankton spring growth in shallow lakes in future with a stronger top‐down control of phytoplankton by zooplankton grazers.     

东海春季水华期浮游植物生长与微型浮游动物摄食

2005年4～6月在东海有害水华频发区14个站位采样,通过现场稀释法实验对春季东海水域浮游植物比生长率和微型浮游动物比摄食率进行了研究.结果表明东海有害水华频发区浮游植物群落以甲藻为优势.浮游植物比生长率在水华爆发前相对较低,平均为1.18 d~(-1);进入水华期后比生长率明显升高,但在水华站位随现存量增加而降低;非水华区比生长率近岸高、远岸低.微型浮游动物主要以急游虫和桡足类幼体为主,而种类上以砂壳纤毛虫居多.微型浮游动物比摄食率在水华爆发前波动较大,介于0.53～1.73 d~(-1),平均为0.90 d~(-1);在水华区比摄食率较为稳定,浮游植物比生长率的降低导致群落净生长率持续下降;在非水华区,比摄食率整体较高,近岸低而远岸高.微型浮游动物的摄食对浮游植物群落的生长有一定的控制作用,但在水华爆发后这种控制作用将减弱.     

Physical processes and their role on the spatial and temporal variability of the spring phytoplankton bloom in the central Yellow Sea

The spatial and temporal variability of the spring phytoplankton bloom (SPB) in the central Yellow Sea is studied, using SeaWiFS surface chlorophyll remote-sensing data, AVHRR sea surface temperatures (SST), QuikSCAT sea surface wind speed (SSW) from 1998 to 2009 and the cruise survey data in 2007 and 2009. The influences of the hydrological conditions on the SPB are significant. (1) The rising SST and low SSW in spring play an important role in the development of the SPB. The SPB in the central Yellow Sea occurs primarily in April (from April 3 to April 24) and at this period the mean SST is generally greater than 10 °C, and 24 h averaged SSW is less than 5.4 m/s. The 99% of the SPB occurs when the SST is 9–14 °C and SSW is 0–7.9 m/s. (2) Specifically, the development of the SPB is from April 4 to April 7 and from April 4 to April 22 in 2007 and 2009 respectively. The longer duration of the SPB in 2009 than that in 2007 is related to the high SST in 2009, which is approximately 2 °C greater than that in 2007, and the weak SSW in 2009, which is much lower than that in 2007, which further indicate that these two factors are critical to the duration of the SPB. (3) The horizontal distribution of surface chlorophyll a in 2007 is found to be greater and located more northward than that in 2009. Comparing the spatial coverage of the SPB, the path of the Yellow Sea warm current, which is warmer and salty, and the location of maximum SST reveals that the spatial coverage of the SPB locates in the warmer temperature (>9 °C) and higher salinity (>33) waters in March and this warm and salty water was much more evident in 2007 than that in 2009 because a stronger warm current of the Yellow Sea in winter. (4) The vertical depths of the maximum chlorophyll layer (MCL) appear at three levels: surface, 10 m and 30 m in 2007, whereas the MCL occurs only at a sub-surface level in 2009 which is related to vertical stability of the water column, i.e., the water is vertically mixed in 2007 while vertically stratified in 2009.     

Physical processes and their role on the spatial and temporal variability of the spring phytoplankton bloom in the central Yellow Sea

The spatial and temporal variability of the spring phytoplankton bloom (SPB) in the central Yellow Sea is studied, using SeaWiFS surface chlorophyll remote-sensing data, AVHRR sea surface temperatures (SST), QuikSCAT sea surface wind speed (SSW) from 1998 to 2009 and the cruise survey data in 2007 and 2009. The influences of the hydrological conditions on the SPB are significant. (1) The rising SST and low SSW in spring play an important role in the development of the SPB. The SPB in the central Yellow Sea occurs primarily in April (from April 3 to April 24) and at this period the mean SST is generally greater than 10 °C, and 24 h averaged SSW is less than 5.4 m/s. The 99% of the SPB occurs when the SST is 9–14 °C and SSW is 0–7.9 m/s. (2) Specifically, the development of the SPB is from April 4 to April 7 and from April 4 to April 22 in 2007 and 2009 respectively. The longer duration of the SPB in 2009 than that in 2007 is related to the high SST in 2009, which is approximately 2 °C greater than that in 2007, and the weak SSW in 2009, which is much lower than that in 2007, which further indicate that these two factors are critical to the duration of the SPB. (3) The horizontal distribution of surface chlorophyll a in 2007 is found to be greater and located more northward than that in 2009. Comparing the spatial coverage of the SPB, the path of the Yellow Sea warm current, which is warmer and salty, and the location of maximum SST reveals that the spatial coverage of the SPB locates in the warmer temperature (>9 °C) and higher salinity (>33) waters in March and this warm and salty water was much more evident in 2007 than that in 2009 because a stronger warm current of the Yellow Sea in winter. (4) The vertical depths of the maximum chlorophyll layer (MCL) appear at three levels: surface, 10 m and 30 m in 2007, whereas the MCL occurs only at a sub-surface level in 2009 which is related to vertical stability of the water column, i.e., the water is vertically mixed in 2007 while vertically stratified in 2009.     

A bioassay study of effects of zooplankton,iron and NTA on the phytoplankton productivity of a marl lake

Additions of iron and NTA had minor stimulatory effects on the phytoplankton productivity of samples of water from two calcareous Michigan lakes in autumn and winter contrary to effects at other seasons. Added Daphnia pulex significantly reduced phytoplankton productivity by grazing at this time of year, in all tests at all levels of addition from 4–32 animals/l, greatly in excess of any possible beneficial effects of increased nutrient availability.     

The effect of small zooplankton on the microbial loop and edible algae during a cyanobacterial bloom

SUMMARY 1. We studied the effect of the small crustacean zooplankton on heterotrophic micro-organisms and edible phytoplankton in a eutrophic lake during a cyanobacterial bloom.
2. Small (15 L) enclosures were filled with natural or screened (100 μm) lake water and incubated for 5 days in the lake. Screening removed crustacean zooplankton but the initial density of rotifers and phytoplankton remained the same in control and removal treatments. Changes in the abundance and biomass of bacteria, autotrophic picoplankton (APP), heterotrophic nanoflagellates (HNF) and ciliates were measured daily.
3. The crustacean zooplankton, dominated by the small cladoceran Chydorus sphaericus , did not affect cyanobacteria, the main phytoplankton group during the experiment.
4. The removal of the crustacean zooplankton induced a higher abundance of ciliates and reduced that of the HNF, indicating the importance of ciliates in controlling HNF in this system.     

Nutrients associated with terrestrial dissolved organic matter drive changes in zooplankton:phytoplankton biomass ratios in an alpine lake

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Life‐history traits buffer against heat wave effects on predator–prey dynamics in zooplankton

In addition to an increase in mean temperature, extreme climatic events, such as heat waves, are predicted to increase in frequency and intensity with climate change, which are likely to affect organism interactions, seasonal succession, and resting stage recruitment patterns in terrestrial as well as in aquatic ecosystems. For example, freshwater zooplankton with different life‐history strategies, such as sexual or parthenogenetic reproduction, may respond differently to increased mean temperatures and rapid temperature fluctuations. Therefore, we conducted a long‐term (18 months) mesocosm experiment where we evaluated the effects of increased mean temperature (4°C) and an identical energy input but delivered through temperature fluctuations, i.e., as heat waves. We show that different rotifer prey species have specific temperature requirements and use limited and species‐specific temperature windows for recruiting from the sediment. On the contrary, co‐occurring predatory cyclopoid copepods recruit from adult or subadult resting stages and are therefore able to respond to short‐term temperature fluctuations. Hence, these different life‐history strategies affect the interactions between cyclopoid copepods and rotifers by reducing the risk of a temporal mismatch in predator–prey dynamics in a climate change scenario. Thus, we conclude that predatory cyclopoid copepods with long generation time are likely to benefit from heat waves since they rapidly “wake up” even at short temperature elevations and thereby suppress fast reproducing prey populations, such as rotifers. In a broader perspective, our findings suggest that differences in life‐history traits will affect predator–prey interactions, and thereby alter community dynamics, in a future climate change scenario.     

Metabolic traits predict the effects of warming on phytoplankton competition

Understanding how changes in temperature affect interspecific competition is critical for predicting changes in ecological communities with global warming. Here, we develop a theoretical model that links interspecific differences in the temperature dependence of resource acquisition and growth to the outcome of pairwise competition in phytoplankton. We parameterised our model with these metabolic traits derived from six species of freshwater phytoplankton and tested its ability to predict the outcome of competition in all pairwise combinations of the species in a factorial experiment, manipulating temperature and nutrient availability. The model correctly predicted the outcome of competition in 72% of the pairwise experiments, with competitive advantage determined by difference in thermal sensitivity of growth rates of the two species. These results demonstrate that metabolic traits play a key role in determining how changes in temperature influence interspecific competition and lay the foundation for mechanistically predicting the effects of warming in complex, multi‐species communities.