On the wind-induced sinking of Sargassum

Sargassum has been photographed on the sea floor at depths of 5000 m. Sargassum was subjected to various pressures to simulate the effect of it being carried down in Langmuir circulation. The time-depth relation for Sargassum to become negatively buoyant was measured; the plant can become negatively buoyant at pressures of only 30 db if left at this pressure for ≈ 5 h. Both the higher critical pressure for the failure of gas vesicles and the longer resistance to subcritical pressure of pelagic Sargassum, compared with the benthonic varieties, suggest an evolutionary adaptation to an open ocean existence. Once negatively buoyant, Sargassum will sink to the sea floor in about 40 h. The measurements are consistent with large scale sinking of Sargassum given the conditions of strong winds and a deep mixed layer both of which exist in winter months in the Sargasso Sea.     

The vertical distribution of phytoplankton in stratified water columns

What determines the vertical distribution of phytoplankton in different aquatic environments remains an open question. To address this question, we develop a model to explore how phytoplankton respond through growth and movement to opposing resource gradients and different mixing conditions. We assume stratification creates a well-mixed surface layer on top of a poorly mixed deep layer and nutrients are supplied from multiple depth-dependent sources. Intraspecific competition leads to a unique strategic equilibrium for phytoplankton, which allows us to classify the distinct vertical distributions that can exist. Biomass can occur as a benthic layer (BL), a deep chlorophyll maximum (DCM), or in the mixed layer (ML), or as a combination of BL+ML or DCM+ML. The ML biomass can be limited by nutrients, light, or both. We predict how the vertical distribution, relative resource limitation, and biomass of phytoplankton will change across environmental gradients. We parameterized our model to represent potentially light and phosphorus limited freshwater lakes, but the model is applicable to a broad range of vertically stratified systems. Increasing nutrient input from the sediments or to the mixed layer increases light limitation, shifts phytoplankton towards the surface, and increases total biomass. Increasing background light attenuation increases light limitation, shifts the phytoplankton towards the surface, and generally decreases total biomass. Increasing mixed layer depth increases, decreases, or has no effect on light limitation and total biomass. Our model is able to replicate the diverse vertical distributions observed in nature and explain what underlying mechanisms drive these distributions.     

Effect of vertical mixing on the vertical distribution of copepods in coastal waters

The vertical distribution of copepod adult stages and naupliihad been studied in a coastal water before and during homogenizationof the water column due to wind. When the water column was stratified,the vertical distribution of the zooplankton was also stratified,as is generally described in the literature. During homogenizationof the water column, different patterns of vertical distributionwere observed. The less active nauplius stage was evenly distributed,even though adult stages always exhibited a stratified distribution.Oithona similis and Microsetclla norvegica exhibited the samedistribution in stratified and unstratified water columns. Temoralongicomis and Pseudocalanus sp. were deeply distributed ina less turbulent area. The causes of these changes are discussed.     

Asynchronous vertical migration and bimodal distribution of motile phytoplankton

Some motile phytoplankton have the capability to exploit deepsources of nutrients in a vertical migration cycle: photosynthesisin the near-surface layer, transit to depth, uptake of the limitingnutrient and transit back to the surface layer. If all foursteps can be completed within 24 h, then migrations can be synchronizedto the day/night cycle to maximize photosynthetic efficiency.Alternatively, if physiological, behavioral or environmentalfactors make it impossible for the cycle to be completed in24 h, then migration may be asynchronous. Many observationsof phytoplankton reveal bimodal vertical distributions of organisms,with maxima near the surface and the nutricline. We demonstratehow bimodal vertical distributions of phytoplankton may be symptomaticof asynchronous vertical migration using a Lagrangian Ensemblenumerical model. We simulate vertical migration of the dinoflagellateAlexandrium fundyense in conditions similar to those in theGulf of Maine, where bimodal distributions of A. fundyense havebeen observed. Migration is regulated by internal nutritionalstate—organisms swim down toward the nitracline when depletedof nitrogen, and return to the surface after nutrient uptake.We test the sensitivity of the results to growth rate, nitrogenuptake rate and swimming speed, and find that organism distributionscan be bimodal or unimodal depending on conditions. Finally,we develop an analytical estimate for population distributionbased on organism characteristics and nutricline depth.     

The influence of fish kairomones on the induction and vertical distribution of sexual individuals of the Daphnia galeata species complex

To assess the potential production of hybrids and backcrosses in a semi-natural environment, we studied the combined effect of fish kairomone, and food level on the production of males and ephippial females in different clones of five Daphnia taxa from the D. galeata species complex. We also studied the diel vertical migration (DVM) of these sexual daphnids under the same varying conditions. This was done to test the hypothesis that males and ephippial females have different migrating strategies, which would increase their mating probability. The study was carried out in two large-scale indoor mesocosms, the so-called `plankton towers' in the Max-Planck Institute in Plön, Germany.Although all of the Daphnia taxa produced ephippial females in the course of the experiment, only D. galeata produced a significant number of males. Fish kairomones had a significant negative influence on the production of ephippial females. We found no DVM in the D. galeata males. They stayed at a depth between 5 and 6 m both day and night, 1 or 2 m above the thermocline. The ephippial females of D. cucullata x hyalina migrated, whereas ephippial females of the other taxa showed no DVM but came significantly closer to the surface in the presence of fish kairomones. We conclude that males and sexual females co-occur in this species complex both in time and space. Therefore, a regular production of hybrids and backcrosses in this species complex seems likely. Fish kairomones do not seem to significantly influence this process.     

The effects of vertical mixing on a phytoplankton community: a modelling approach to the intermediate disturbance hypothesis

1.  Connell's (1978) intermediate disturbance hypothesis (IDH) has been proposed as one explanation of why diversity is often highest at intermediate levels of disturbance. We used a model phytoplankton responses to environmental change (PROTECH) to investigate the validity of this hypothesis.
2. In a simulated phytoplankton assemblage of eight species, we found that the relationship between the increased intensity of a single forcing event and diversity was described by a positively skewed curve.
3. A progressive increase in forcing frequency introduced a sharp decrease in diversity at a threshold frequency. However, the highest diversity values were found at an intermediate frequency of disturbance.
4. We described the shape of this breakpoint response as like a 'cliff' and reconcile it with multiple stable-point theory. It is argued that the IDH should possibly be represented by this 'cliff' relationship, which may be applied to (or encourage the re-examination of) many previous studies.     

Silicate availability, vertical mixing and grazing control of phytoplankton blooms in mesocosms

Lake Sempach, located in the central part of Switzerland, has a surface area of 14 km², a maximum depth of 87 m and a water residence time of 15 years. Restoration measures to correct historic eutrophication, including artificial mixing and oxygenation of the hypolimnion, were implemented in 1984. By means of the combination of external and internal load reductions, total phosphorus concentrations decreased in the period 1984–2000 from 160 to 42 mg P m^–3. Starting from 1997, hypolimnion oxygenation with pure oxygen was replaced by aeration with fine air bubbles. The reaction of the plankton has been investigated as part of a long-term monitoring program. Taxa numbers, evenness and biodiversity of phytoplankton increased significantly during the last 15 years, concomitant with a marked decline of phosphorus concentration in the lake. Seasonal development of phytoplankton seems to be strongly influenced by the artificial mixing during winter and spring and by changes of the trophic state. Dominance of nitrogen fixing cyanobacteria (Aphanizomenon sp.), causing a severe fish kill in 1984, has been correlated with lower N/P-ratio in the epilimnion. Buoyant algae such as Planktothrix rubescens (syn. Oscillatoria rubescens) increased in abundance due to enlargement of the trophogenic layer and extended mixing depth during winter. The interactions between zoo- and phytoplankton seemed to be depressed as a result of restoration measures. Zooplankton composition changed to more carnivorous and less herbivorous species. Oxygenation of the hypolimnion induced bioturbation of sediments, mainly by oligochaetae worms, and stimulated germination of spores and cysts and hatching of resting eggs.     

The impact of climate on the geographical distribution of phytoplankton species in boreal lakes

Here, we use a novel space-by-time approach to study large-scale changes in phytoplankton species distribution in Swedish boreal lakes in response to climate variability. Using phytoplankton samples from 27 lakes, evenly distributed across Sweden, all relatively unimpacted by anthropogenic disturbance and sampled annually between 1996 and 2010, we found significant shifts in the geographical distribution of 18 species. We also found significant changes in the prevalence of 45 species (33 became more common and 12 less common) over the study period. Using species distribution models and phytoplankton samples from 60 lakes sampled at least twice between 1992 and 2010, we evaluated the importance of climate variability and other environmental variables on species distribution. We found that temperature (e.g., extreme events and the duration of the growing season) was the most important predictor for species detections. Many cyanobacteria, chlorophytes, and, to a lesser extent, diatoms and zygnematophytes, showed congruent and positive responses to temperature. In contrast, precipitation explained little variation and was important only for a few taxa (e.g., Staurodesmus spp., Trachelomonas volvocina). At the community level, our results suggest a change in community composition at temperatures over 20 °C and growing seasons longer than 40 days. We conclude that climate is an important driver of the distributional patterns of individual phytoplankton species and may drive changes in community composition in minimally disturbed boreal lakes.     

The influence of fluctuating light intensities on species composition and diversity of natural phytoplankton communities

The influence of fluctuating light intensities on phytoplankton composition and diversity was investigated for 49 days under semi-continuous culture conditions with sufficient nutrient supply, using phytoplankton assemblages from Lake Biwa, Japan. Light conditions were either periodically changed from high intensity (100 µmol photons m^-2 s^-1) to low intensity (20 µmol photons m^-2 s^-1) at intervals of 1, 3, 6 and 12 days, or fixed to constant intensities (permanent high and low light levels). All treatments additionally experienced a day:night cycle of 16:8 h. Phytoplankton abundance increased and reached a saturation level on day 19 of the treatment with permanent high light, but increased continuously until the end of the experiment (day 49) in the treatment with permanent low light intensity. In treatments with periodically changing light intensities, the phytoplankton abundance reached saturation levels between these dates. Under phytoplankton abundance saturation, chlorophytes predominated in the treatment with permanent high light, while either cyanophytes or diatoms were abundant under permanent low light intensity. Treatments with changing light supply had chlorophyte- and cyanobacteria-dominated replicates as well as replicates with balanced proportions of both. Furthermore, species diversity, measured by the Shannon index, was low in cultures under permanent light intensity, while slow fluctuating light at the scale of 3 -12 days resulted in an increased diversity index. These results indicate that species composition and diversity of the phytoplankton were affected by the periodically changing light regime in the order of days, and suggest that temporal changes in weather conditions are a major impediment to competitive exclusion of phytoplankton species in nature.     

The influence of predation on horizontal distribution of zooplankton species

SUMMARY. 1. The horizontal distributions of Daphnia longispina and Bosmina tongispina in Lake Kvernavatn (Norway) were investigated twice in 1982. In late spring, when populations were small, the two species inhabited the same areas, and they were evenly distributed from the littoral to the pelagic. At high population densities, during midsummer, the species were spatially segregated, D. longispina being pelagic and B. longispina littoral in distribution.
2. The distribution and feeding of three-spined sticklebacks (Gasteros-teus aculeatus) were also studied. The sticklebacks were apparently forced into littoral areas by larger piscivorous predators in the pelagic and they were consequently restricted to foraging primarily on B. longispina, which formed dense swarms during daytime in summer.
3. We suggest that predation and competition influence the spatial distribution of zooplankton species. The feeding efficiency of fish foraging on high-density zooplankton populations can be reduced by spatial segregation of zooplankton species. Where high local densities occur, due to swarm formation, predation is changed from size-selective feeding to consumption of spatially isolated individuals.     

Seasonal variation of mixing depth and its influence on phytoplankton dynamics in the Zeya reservoir, China

In reservoirs or lakes, mixing depth affects growth and loss rates of phytoplankton populations. Based on 1-year data from the Zeya reservoir, China, we scaled the mixing depth throughout a whole year by utilizing cluster analysis, and then investigated its influence on phytoplankton dynamics and other physical and chemical parameters. Over the whole year, all physical and chemical parameters except TN and temperature had significant correlations with mixing depth, indicating that mixing depth is one of the important driving factors influencing water environment. According to mixing depth, a year can be divided into three different periods, including the thermally stratified period, isothermally mixed period, and transition period between them. When considering the former two different periods separately, mixing depth had no correlation with the phytoplankton biovolume. However, over the whole year a significant correlation was observed, which indicated that the influence of mixing depth on phytoplankton growth in the Zeya reservoir still followed Diehl’s theory. Furthermore, according to the steady-state assumption, a unimodal curve (mixing depth—phytoplankton biovolume) with a significant peak appearing at a mixing depth of 2 m was observed, closely agreeing with Diehl’ prediction.     

How internal waves influence the vertical distribution of zooplankton

1. We present data with a high spatio‐temporal resolution from a 72‐h field survey in Bautzen Reservoir (Saxony, Germany). The aims of this survey were to observe hydrophysical processes during a period of unstable stratification in spring and investigate the effect of wind‐induced internal waves on the vertical distribution of zooplankton. 2. Wind velocities up to 10 m s⁻¹ caused a strong downwelling event of warm water at the sampling site and led to the generation of internal waves with an amplitude of 4 m. 3. The zooplankton community, which was dominated by Daphnia galeata, inhabited epilimnetic waters. Downwelling enlarged the thickness of the epilimnetic layer and, hence, led to high zooplankton abundances down to relatively deep water strata indicating lateral transport of zooplankton. As a consequence, area‐specific zooplankton abundances increased considerably (max. fourfold) during downwelling. 4. We conclude that classical limnological field sampling, such as for monitoring purposes, can lead to severely biased estimates of zooplankton abundance due to the interfering effects of hydrophysical processes like internal waves. 5. Backscattering strengths measured by a simultaneously deployed Acoustic Doppler Current Profiler (600 kHz) were found to be correlated with estimated zooplankton abundances based on plankton samples.     

The influence of copepod and krill grazing on the species composition of phytoplankton communities from the Scotia Weddell sea

The influence of copepods (mainly Oithona sim-ilis) and krill (Euphausia superba) grazing on the species composition of plankton communities in ship board con tainers was investigated during the spring and post spring period in the Scotia Weddell Sea in the Antarctic ocean. Numbers of grazers were experimentally manipulated in containers with natural phytoplankton assemblages. With ratural levels of copepods but no krill a high (700–950 g C·l¹, ca 30 g chl a·.l¹) phytoplankton biomass developed. In these cultures large diatoms, e.g. Corethron criophilum and chains of Thalassiosira sp., made up 80% of total phytoplankton cell carbon at the end of the experiment. In cultures with elevated numbers of copepods (5X or 10X the natural level) phytoplankton biomass was somewhat reduced (ca 23 g chl a · l¹) compared to cultures with natural copepod abundance, but still high. Phytoplankton species composition was on the other hand greatly influenced. Instead of large diatoms these cultures were dominated by Phaeocystis pouchetii (70%) together with small Nitszchia sp. and Chaetoceros neogracile (20%). In containers with krill (both juveniles and adults), but without elevated numbers of copepods, phytoplankton biomass rapidly approached zero. With 10X the in situ level of copepods, krill first preyed on these before Corethron criophilum and Thalassiosira sp. were grazed. When krill were removed a plankton community dominated by flagellates (60–90%), e.g. Pyramimonas sp. and a Cryptophycean species, grazed by an unidentified droplet-shaped heterothrophic flagellate, developed. These flagellates were the same as those which dominated the plankton community in the Weddell Sea after the spring bloom. A similar succession was observed in situ when a krill swarm grazed down a phytoplankton bloom in a few hours. Our experiments show that copepods cannot control phytoplankton biomass in shipboard cultures even at artificially elevated numbers. Krill at concentrations similar to those in natural swarms have a great impact on both phytoplankton biomass and species composition in shipboard cultures. Both copepods and krill may have an impact on phytoplankton species composition and biomass in situ since the rates of phytoplankton cell division were probably artificially increased in shipboard cultures compared to natural conditions, where lower growth rates make phytoplankton more vulnerable to grazing. A similarity between phytoplankton successions in containers and in situ, especially with respect to krill grazing, supports the conclusion that grazing may structure phytoplankton communities in the Scotia-Weddell Sea.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

The influence of bacteria on the arsenic species produced by laboratory cultures of the marine phytoplankton Dunaliella tertiolecta

To assess whether bacteria influence the biogeochemical cycling of arsenic by laboratory cultures of the marine phytoplankton Dunaliella tertiolecta, the arsenic species produced by D. tertiolecta were compared in “operationally sterile” and bacteria spiked cultures. It was observed that glycerol (Gly-) arsenoriboside (41–78 %), phosphate (PO₄?) arsenoriboside (7–38 %) and arsenate (As(V)) (15–21 %) were the major water-soluble arsenic species in all D. tertiolecta cultures irrespective of whether cultures were operationally sterile or contained added bacteria. PO₄-riboside (46–74 %) and Gly-riboside (24–36 %) were also the major arsenic species in hydrolysed lipid extracts of D. tertiolecta irrespective of whether cultures were operationally sterile or contained bacteria. In addition to similarities in the arsenic species produced, total arsenic concentrations and culture growth did not differ relative to whether cultures were operationally sterile or not. Similar bacterial strains were identified in all D. tertiolecta cultures irrespective of whether bacteria were added or not. Consequently, it is evident that the presence of “foreign” or “added” bacteria in D. tertiolecta has minimal influence on the metabolism and cycling of arsenic by phytoplankton. Thus, the use of laboratory phytoplankton cultures containing bacteria may be appropriate means to investigate arsenic biogeochemical cycling unlike previously believed.     

Regional, vertical and seasonal distribution of phytoplankton and photosynthetic pigments in Lake Baikal

A 3-year phytoplankton study was carried out in Lake Baikal(Siberia) as part of the CONTINENT project and in conjunctionwith a 60-year long monitoring programme by the Irkutsk StateUniversity. A combination of microscopy and high performanceliquid chromatography (HPLC) pigment analysis was used. Allover the lake, the dominant functional group (by biovolume)was the vernal diatom blooms, due to the dominance of endemicCyclotella species. Chlorophyll a (Chl a) was significantlyhighest at the Selenga and Barguzin inflows (2.39 ± 0.34and 2.49 ± 0.18 nmol L^–1, mean ± 95% CI,respectively) and higher in the South than in the North (1.43± 0.26 and 0.96 ± 0.13 nmol L^–1). This variationof Chl a reflected changes in the phytoplankton composition.Diatoms and Chrysophyceae were the major contributors to thetotal Chl a except in the South (Chlorophyceae) and SelengaDelta (cyanobacterial picoplankton). There were also indicationsof species composition changes due to enhanced P-loading fromthe Selenga River. However, canonical analyses indicated thattemperature and stratification were the major driving forcesfor regional distribution patterns and seasonal succession.It seems likely that further global warming will cause a shiftin the species and group composition towards small cells atthe expense of the large endemic diatom flora.     

Residence time and exposure time of sinking phytoplankton in the euphotic layer

The residence time of a sinking particle in the euphotic layer is usually defined as the time taken by this particle to reach for the first time the bottom of the euphotic layer. According to this definition, the concept of residence time does not take into account the fact that many cells leaving the euphotic layer at some time can re-enter the euphotic layer at a later time. Therefore, the exposure time in the surface layer, i.e. the total time spent by the particles in the euphotic layer irrespective of their possible excursions outside the surface layer, is a more relevant concept to diagnose the effect of diffusion on the survival of phytoplankton cells sinking through the water column.While increasing the diffusion coefficient can induce both a decrease or an increase of the residence time, the exposure time in the euphotic layer increases monotonically with the diffusion coefficient, at least when the settling velocity does not increase with depth. Turbulence is therefore shown to increase the total time spent by phytoplankton cells in the euphotic layer.The generalization of the concept of exposure time to take into account the variations of the light intensity with depth or the functional response of phytoplankton cells to irradiance leads to the definition of the concepts of light exposure and effective light exposure. The former provides a measure of the total light energy received by the cells during their cycling through the water column while the latter diagnose the potential growth rate.The exposure time, the light exposure and the effective light exposure can all be computed as the solution of a differential problem that generalizes the adjoint approach introduced by Delhez et al. (2004) for the residence time. A general analytical solution of the 1D steady-state version of this equation is derived from which the properties of the different diagnostic tools can be obtained.     

北京地区蝗虫种类研究及其垂直分布调查

经过2003~2004年对北京市平原区和山区的调查研究,初步发现北京市蝗虫有49种,隶属8科33属。由于植被、生态因素与环境条件和海拔高度的不同,蝗虫种类的垂直分布存在较明显的差异。对农林业为害较重的主要优势种有10种。     

The influence of different disturbance frequencies on the species richness,diversity and equitability of phytoplankton in shallow lakes

The relationships between the species richness, diversity and equitability of phytoplankton is discussed in the context of Connell's (1978, Science 199: 1304–1310) Intermediate Disturbance Hypothesis (IDH). The records of 759 vertical phytoplankton samples, which were obtained from four shallow central European lakes (Balaton, Neusiedlersee, and two small artificial ponds) at daily to weekly intervals were analysed.

1. The Shannon-Weaver function was used to measure diversity of the recorded species compositionof the phytoplankton. It is shown on fictitious data that compositional diversity is sensitive to the numberof coequilibrating species provided that the suspected interrelationship between diversity and ‘complexity’is amenable to the application of this method.
2. The disturbance scale that was developed on the basis of the field records fits well to Reynolds'(1988, Verh. int. Ver. Limnol. 23: 683–691) derivation: < 3 days qualifies as high frequency, approximatel3–8 days as intermediate frequency and > 8–9 days as low frequency of disturbance for phytoplankton.
3. Arithmetical means of the compositional diversity of phytoplankton under different frequencies ofdisturbance support the hypothesis that maximal diversity appears at intermediate frequencies.
4. There are different reasons for decrease in diversity at higher and lower frequencies. Inequitabilitydiminishes diversity at low disturbance; while species number decreases at high frequencies.
5. The case of Neusiedlersee calls attention to the fact that it is difficult, if at all possible, to differentiatebetween the indices under continuous stress and high frequency of disturbance in lakes in temperateregions. Similar species number-equitability pattern are induced by both and it is also presumablethat high frequency disturbance can itself effect a serious stress.
6. The striking effects that regular major periodic events (e.g. significant changes in the grazing pressureat the onset of the clear-water phase, autumnal cooling) in the plankton have on its species diversity areevident. Thus, the relative importance of intermediate frequency disturbances has its own seasonality:it is increasingly important in periods (partly in the spring, but mostly in the summer-autumn equilibriumphases), in which competition among phytoplankton species is increasing. This observation suggestsa way by which the stochasticity-based IDH can be incorporated into rather more deterministicexplanations (e.g. PEG-model; Sommer et al., 1986. Archiv für Hydrobiologie 106: 433–471) of planktonsuccession.
7. The most controversial issue and, therefore, the main difficulty, with IDH is that it not onlymaintains species richness in an ecosystem but it also supposes its presence. The lack of either earlyor late successional species in a given community can inactivate the mechanism. From the point of viewof the diversity-species richness relationship, the persistence of disturbance at given frequencies is ofgreater importance than the temporal alterations themselves in the evolutionary ecology of the phytoplankton.
8. For characteristically unperturbed phytoplankton communities (no case was studied here), equilibriumconcepts (niche diversification, etc.) should be more strongly applicable to their diversity andspecies richness.