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 influence of vertical mixing on the photoinhibition of variable chlorophyll a fluorescence and its inclusion in a model of phytoplankton photosynthesis

The maximum effective quantum yield of photosystem II was estimatedfrom measurements of variable chlorophyll a fluorescence [(F'_m- F'_o)/F'_m = F'_v/F'_m] in samples of phytoplankton collectedfrom various depths in Chaffey Reservoir, Australia. Duringstratified conditions, F'_v/F'_m showed depth-dependent decreasesas irradiance increased during the morning, and increases asirradiancereduced in the afternoon. Wind-induced mixing disrupted thediel pattern, but even under well-mixed conditions a verticalgradient in F'_v/F'_m remained. Differences in F'_v/F'_m valuesbetween samples incubated at fixed depths and unconstrainedlake samples enabled identification of the phytoplankton mixingdepth. Recovery of F'_v/F'_m was modelled as a function of timeand the degree of F'_v/F'_m inhibition, while damage was considereda function of photon dose. A combined, numerical model was fittedto diel sequences of F'_v/F'_m to estimate rate constants fordamage and repair. Recovery rate constants ranged from r = 0.7x 10^-4 to 9.1 x 10^-4 s^-1, while damage rate constants rangedfrom k = 0.03 to 0.22 m² mol photon^-1. A fluorescence-basedmodel of photosynthesis was used to investigate the effectsof wind speed, euphotic depth and mixed layer depth on photoinhibition.At different mixing rates, depth-integrated photosynthesis wasenhanced by up to 16% under the conditions tested, while increasesof 9% occurred between phytoplankton with different measureddamage and repair characteristics.     

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.     

垂向湍流、温、光对浅水湖泊下沉藻类垂向分布的影响

垂向湍流、温、光等是影响藻类生长和垂向分布的关键因素。基于大型浅水湖泊（太湖）的背景物理场（辐射、水深、水温和消光等）,利用藻类生长-扩散模型的敏感性试验,探讨物理过程对浅水湖泊中下沉藻的垂向分布的影响机制。水温是次表层叶绿素峰值（Subsurface Chlorophyll Maxinmum：SCMax）或谷值（Subsurface Chlorophyll Minimum：SCMin）形成的限制性条件;当水温大于阈值时,随深度的增大,过强光能抑制藻类的生长转变为弱光限制藻类的生长是SCMax形成的关键;垂向湍流的裹挟作用是弱化SCMax强度的关键过程。该研究有利于厘清物理过程对藻类群落演替的作用机制,强化对水生态系统的修复效果。     

长江口锋面附近咸淡水混合对浮游植物生长影响的现场培养

通过2009年6月调查航次,获得了营养盐等参数断面分布,表明咸淡水混合是控制营养盐分布的主要因素。为了解不同盐度梯度下浮游植物生长与营养盐吸收的关系,采集两个站位水样分别代表长江冲淡水(C1站)和外海水(I10站),按C1站水样比例,分100%、75%、50%、25%、0%不同比例混合进行现场模拟咸淡水混合培养,有以下认识:(1)平行结果表明培养过程中活体荧光极大值在100%混合组,且淡水比例越低,指数生长期0—48 h内生长速率越低,100%、75%、50%、25%组分别为1.18/d、1.12/d、1.14/d、0.77/d;(2)低于26盐度的水体中PO34-在48 h内可被迅速消耗而产生限制作用,是控制浮游植物生长的潜在限制因子;(3)除0%组外,各混合组DIN/P(DIN:溶解无机氮,Dissolved Inorganic Nitrogen,DIN=NO3-+NO-2+NH4+)比值在浮游植物指数生长期有升高趋势,100%组DIN/P比值增加了一倍。各组培养48 h后DIN/Si比值逐渐降低至原来的0.7左右,初始DIN/Si小于一定时间内硅藻吸收的(ΔDIN/ΔSi)比,是造成各组DIN/Si比值减小的原因。以上结果表明咸淡水混合过程中形成的营养盐梯度可造成浮游植物生长程度和速率差异,且可因局部浮游植物旺发而改变海水营养结构。     

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 vertical distributions of chlorophyll and phytoplankton species in the North Pacific central environment

The phytoplankton species in the North Pacific central environmentare known to be distributed into two vertically distinct assemblagesduring most of the year. Key species are defined for each assemblage.The vertical distributions of these key species indicate thatthe increase in abundance of deep species closely parallelsthe increase in chlorophyll a at the top of the chlorophyllmaximum layer. The chlorophyll maximum is comprised of speciescharacteristic of the deep assemblage, with only insignificantnumbers of shallow species.     

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.     

东海浮游植物数量分布与优势种

根据1997～2000年东海23°30′～33°N、118°30′～128°E海域4个季节海洋调查资料,运用定量、定性方法,探讨了东海浮游植物总丰度的平面分布、季节分布及变化的动力学机制,同时还探讨了东海浮游植物优势种的生态学特征和对温盐环境适应特征。结果表明:东海浮游植物总丰度有明显的季节分布,秋季总丰度达到4季最高峰,均值为211.91×104cell.m-3,夏季次之(50.40×104cell.m-3),冬季11.34×104cell.m-3,春季(2.01×104cel.lm-3)最低。春夏季东海近海(Ⅰ和Ⅲ)高于外海(Ⅱ和Ⅳ),秋冬季东海北部外海(Ⅱ)高于近海(Ⅰ),南部近海(Ⅲ)高于外海(Ⅳ)。4季共出现优势种(Y≥0.02)11种,冬季的主要优势种为洛氏角毛藻和细弱海链藻,春季的主要优势种为洛氏角毛藻和夜光藻,夏季以拟弯角毛藻和细长翼根管藻为主要优势种,秋季优势种仅为聚生角毛藻。浮游植物的聚集强度指数为较大正值,聚集作用明显。从全年的逐步回归分析结果看,温度是影响浮游植物总丰度季节分布的主要因子,盐度是次要因子。浮游植物的11个优势种之间生存环境适应比较,洛氏角毛藻和中华盒形藻生长温度幅度较大,可以在12～28℃水温增长,因而春、夏和冬季都成为优势种。中肋骨条藻适合生长的水温范围都较小,仅为22～28℃。聚生角毛藻更小,仅21～25℃。洛氏角毛藻和中华盒形藻生长温盐度和高分布区温盐度范围比聚生角毛藻和中肋骨条藻都广,而秋季数量却低于聚生角毛藻。     

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