Spring phenological responses of marine and freshwater plankton to changing temperature and light conditions

Shifts in the timing and magnitude of the spring plankton bloom in response to climate change have been observed across a wide range of aquatic systems. We used meta-analysis to investigate phenological responses of marine and freshwater plankton communities in mesocosms subjected to experimental manipulations of temperature and light intensity. Systems differed with respect to the dominant mesozooplankton (copepods in seawater and daphnids in freshwater). Higher water temperatures advanced the bloom timing of most functional plankton groups in both marine and freshwater systems. In contrast to timing, responses of bloom magnitudes were more variable among taxa and systems and were influenced by light intensity and trophic interactions. Increased light levels increased the magnitude of the spring peaks of most phytoplankton taxa and of total phytoplankton biomass. Intensified size-selective grazing of copepods in warming scenarios affected phytoplankton size structure and lowered intermediate (20–200?μm)-sized phytoplankton in marine systems. In contrast, plankton peak magnitudes in freshwater systems were unaffected by temperature, but decreased at lower light intensities, suggesting that filter feeding daphnids are sensitive to changes in algal carrying capacity as mediated by light supply. Our analysis confirms the general shift toward earlier blooms at increased temperature in both marine and freshwater systems and supports predictions that effects of climate change on plankton production will vary among sites, depending on resource limitation and species composition.     

Effects of water temperature and mixed layer depth on zooplankton body size

Ecological consequences of global warming include shifts of species ranges toward higher altitudes and latitudes as well as temporal shifts in phenology and life-cycle events. Evidence is accumulating that increasing temperature is also linked to reduced body size of ectotherms. While temperature can act directly on body size, it may also act indirectly by affecting the timing of life-cycle events and the resulting population age and size structure, especially in seasonal environments. Population structure may, in turn, be influenced by temperature-driven changes in resource availability. In a field mesocosm experiment, we investigated how water temperature and mixed surface layer depth (a temperature-dependent determinant of light availability to phytoplankton) affected population dynamics, population age and size structure, and individual size at stage (size at first reproduction) of Daphnia hyalina during and after a phytoplankton spring bloom. Mixed layer depth was inversely related to the magnitudes of the phytoplankton spring bloom and the subsequent Daphnia peak, but had no effect on the body size of Daphnia. Conversely, temperature had no effects on abundance peaks but strongly affected the timing of these events. This resulted in at times positive, at other times negative, transient effects of temperature on mean body size, caused by asynchronous changes in population size structure in cold versus warm treatments. In contrast to mean body size, individual size at stage consistently decreased with increasing temperature. We suggest that size at stage could be used as an unbiased response parameter to temperature that is unaffected by transient, demographically driven changes in population size structure.     

Mechanistic origins of variability in phytoplankton dynamics. Part II: analysis of mesocosm blooms under climate change scenarios

Driving factors of phytoplankton spring blooms have been discussed since long, but rarely analyzed quantitatively. Here, we use a mechanistic size-based ecosystem model to reconstruct observations made during the Kiel mesocosm experiments (2005–2006). The model accurately hindcasts highly variable bloom developments including community shifts in cell size. Under low light, phytoplankton dynamics was mostly controlled by selective mesozooplankton grazing. Selective grazing also explains initial dominance of large diatoms under high light conditions. All blooms were mainly terminated by aggregation and sedimentation. Allometries in nutrient uptake capabilities led to a delayed, post-bloom dominance of small species. In general, biomass and trait dynamics revealed many mutual dependencies, while growth factors decoupled from the respective selective forces. A size shift induced by one factor often changed the growth dependency on other factors. Within climate change scenarios, these indirect effects produced large sensitivities of ecosystem fluxes to the size distribution of winter phytoplankton. These sensitivities exceeded those found for changes in vertical mixing, whereas temperature changes only had minimal impacts.     

Mass encystment and sinking of dinoflagellates during a spring bloom

The decline of a spring bloom dominated by dinoflagellates and the mass sedimentation of dinoflagellate cysts was documented in a coastal area of the northern Baltic Sea, SW Finland in 1983. The exceptionally large spring phytoplankton bloom was observed in early May. After depletion of nitrate phytoplankton biomass declined rapidly. The bloom was followed by intense sedimentation of spherical cysts and of organic matter at the end of May. These cysts were presumably hypnozygotes of Peridinium hangoei Schiller. Sedimentation of dinoflagellate cysts was estimated to correspond to ca. 45% of the maximum sedimentation of particulate organic carbon at this time, although most of the dinoflagellate biomass disintegrated already in the water column and was deposited as organic detritus or washed away by advection. It is concluded that the life cycle strategies of the dominant vernal phytoplankton species have a major impact on the sedimentation of the spring bloom.     

Winter atmospheric circulation signature for the timing of the spring bloom of diatoms in the North Sea

Analysing long-term diatom data from the German Bight and observational climate data for the period 1962–2005, we found a close connection of the inter-annual variation of the timing of the spring bloom with the boreal winter atmospheric circulation. We examined the fact that high diatom counts of the spring bloom tended to occur later when the atmospheric circulation was characterized by winter blocking over Scandinavia. The associated pattern in the sea level pressure showed a pressure dipole with two centres located over the Azores and Norway and was tilted compared to the North Atlantic Oscillation. The bloom was earlier when the cyclonic circulation over Scandinavia allowed an increased inflow of Atlantic water into the North Sea which is associated with clearer, more marine water, and warmer conditions. The bloom was later when a more continental atmospheric flow from the east was detected. At Helgoland Roads, it seems that under turbid water conditions (=?low light) zooplankton grazing can affect the timing of the phytoplankton bloom negatively. Warmer water temperatures will facilitate this. Under clear water conditions, light will be the main governing factor with regard to the timing of the spring bloom. These different water conditions are shown here to be mainly related to large-scale weather patterns. We found that the mean diatom bloom could be predicted from the sea level pressure one to three months in advance. Using historical pressure data, we derived a proxy for the timing of the spring bloom over the last centuries, showing an increased number of late (proxy-) blooms during the eighteenth century when the climate was considerably colder than today. We argue that these variations are important for the interpretation of inter-annual to centennial variations of biological processes. This is of particular interest when considering future scenarios, as well to considerations on past and future effects on the primary production and food webs.     

The role of ciliates,heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads,North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P _p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P _p . Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.     

Warming induces shifts in microzooplankton phenology and reduces time-lags between phytoplankton and protozoan production

Indoor mesocosm experiments were conducted to test for potential climate change effects on the spring succession of Baltic Sea plankton. Two different temperature (Δ0?°C and Δ6?°C) and three light scenarios (62, 57 and 49?% of the natural surface light intensity on sunny days), mimicking increasing cloudiness as predicted for warmer winters in the Baltic Sea region, were simulated. By combining experimental and modeling approaches, we were able to test for a potential dietary mismatch between phytoplankton and zooplankton. Two general predator–prey models, one representing the community as a tri-trophic food chain and one as a 5-guild food web were applied to test for the consequences of different temperature sensitivities of heterotrophic components of the plankton. During the experiments, we observed reduced time-lags between the peaks of phytoplankton and protozoan biomass in response to warming. Microzooplankton peak biomass was reached by 2.5 day °C^?1 earlier and occurred almost synchronously with biomass peaks of phytoplankton in the warm mesocosms (Δ6?°C). The peak magnitudes of microzooplankton biomass remained unaffected by temperature, and growth rates of microzooplankton were higher at Δ6?°C (μ_?0?°C?=?0.12 day^?1 and μ_?6?°C?=?0.25 day^?1). Furthermore, warming induced a shift in microzooplankton phenology leading to a faster species turnover and a shorter window of microzooplankton occurrence. Moderate differences in the light levels had no significant effect on the time-lags between autotrophic and heterotrophic biomass and on the timing, biomass maxima and growth rate of microzooplankton biomass. Both models predicted reduced time-lags between the biomass peaks of phytoplankton and its predators (both microzooplankton and copepods) with warming. The reduction of time-lags increased with increasing Q₁₀ values of copepods and protozoans in the tritrophic food chain. Indirect trophic effects modified this pattern in the 5-guild food web. Our study shows that instead of a mismatch, warming might lead to a stronger match between protist grazers and their prey altering in turn the transfer of matter and energy toward higher trophic levels.     

Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

Effect of temperature and light on the growth of micro- nano- and pico-plankton: impact on algal succession

The influence of light and temperature on the phytoplankton succession in a temperate sea area was investigated in laboratory experiments with natural assemblages of micro-, nano-, and picoplankton collected from the northern Baltic Sea during 1989 and 1990. Respiration increased from 0 to 30°C in all groups of phytoplankton, while gross photosynthesis stabilised at 10 to 15°C. Light saturation occurred at 25 to 75 mol quanta m^-2 s^-1, indicating low light adaptation of the algae. Picoplankton showed the strongest temperature response, and at temperatures above 10°C picoplankton obtained a higher biomass specific photosynthesis than that of the other groups. Different light treatments had no effect on the species composition in experiments with natural algal assemblages, while different temperature conditions had a marked effect. With a temperature increase from 0 to 10°C, the algal community changed from a typical spring community, with diatoms and dinoflagellates, to a summer community, dominated by mixotrophic nanoflagellates. The small, or often non-existing, autumn bloom in the sea area studied, can be explained by short day lengths combined with relatively high temperatures, causing high community respiration rates and low gross photosynthesis, resulting in a negative energy balance. The net energy gain depends on a differential temperature effect on gross photosynthesis and endogenous respiration in various plankton groups. This gives the phytoplankton groups diverse competitive advantages during different seasons and thus may be an important factor in controlling algal succession.     

Picophytoplankton responses to changing nutrient and light regimes during a bloom

The spring bloom in seasonally stratified seas is often characterized by a rapid increase in photosynthetic biomass. To clarify how the combined effects of nutrient and light availability influence phytoplankton composition in the oligotrophic Gulf of Aqaba, Red Sea, phytoplankton growth and acclimation responses to various nutrient and light regimes were recorded in three independent bioassays and during a naturally-occurring bloom. We show that picoeukaryotes and Synechococcus maintained a “bloomer” growth strategy, which allowed them to grow quickly when nutrient and light limitation were reversed. During the bloom picoeukaryotes and Synechococcus appeared to have higher P requirements relative to N, and were responsible for the majority of photosynthetic biomass accumulation. Following stratification events, populations limited by light showed rapid photoacclimation (based on analysis of cellular fluorescence levels and photosystem II photosynthetic efficiency) and community composition shifts without substantial changes in photosynthetic biomass. The traditional interpretation of “bloom” dynamics (i.e., as an increase in photosynthetic biomass) may therefore be confined to the upper euphotic zone where light is not limiting, while other acclimation processes are more ecologically relevant at depth. Characterizing acclimation processes and growth strategies is important if we are to clarify mechanisms that underlie productivity in oligotrophic regions, which account for approximately half of the global primary production in the ocean. This information is also important for predicting how phytoplankton may respond to global warming-induced oligotrophic ocean expansion.     

Succession patterns of phytoplankton blooms: directionality and influence of algal cell size

Using four replicate microcosms in the laboratory, we induced a phytoplankton bloom by enclosing a natural community sampled from Masnou Harbor (N.E. Spain) in November 1987, and examined the pattern of algal succession during the bloom. Good replicability of the temporal patterns of the community biomass and the abundance of most species demonstrated that succession was a directional, non-random process. The successional pathway observed (small flagellates » small centric diatoms » small flagellates) resembled that observed by other authors studying phytoplankton blooms. This pattern differed from previous models of algal succession in that dinoflagellates never comprised a substantial fraction of the community biomass, and in that algal cell size did not tend to increase along the successional sequence. Algal cell size, however, was an important determinant of phytoplankton community structure, since it constrained the density, but not the biomass, achievable by the different species. We suggest that there is not a single, general pattern of phytoplankton succession, but that distinction should be made, at least between seasonal and bloom patterns of phytoplankton succession.     

Seasonal and long-term variations in the body size of planktonic copepods in the northern Baltic Sea

Seasonal and long-term variations in the body size of planktonic copepods were studied in the northern Baltic Sea. The influence of temperature, salinity and phytoplankton concentration to the body size of Eurytemora affinis, Acartia bifilosa and Limnocalanus macrurus (Calanoida) was examined at three sea areas, differing in their hydrographical and trophic conditions (an archipelago area and an enclosed bay on the SW coast of Finland in 1992 and at an open sea station in the Gulf of Bothnia in 1991). There was an inverse relationship between copepod body length and temperature, while there was a direct effect of phytoplankton concentration. According to multiple regression analysis, the relative importance of these factors varied according to species, developmental stage and study area. In the archipelago area, copepod body size was mainly determined by temperature, while in the open Gulf of Bothnia, phytoplankton concentration was usually more important. Interannual variation in the summer body length of E. affinis and A. bifilosa was examined using samples collected over 18 years (1967 to 1984) in the archipelago area. In this analysis, the inverse relationship between water temperature and copepod body size disappeared; in E. affinis the average female length correlated positively with summer temperature. We suggest that the length and timing of the study period greatly influence the investigator's view of the factors regulating copepod body size.     

Pelagic-benthic coupling in a subtidal system of the North-Western Mediterranean

Suspended particulate matter, zooplankton, and macrobenthos dynamics were investigated in a shallow area of the Ligurian Sea (north-west Mediterranean) characterized by wide temporal variability over an annual cycle. As indicated by multivariate analyses, the seasonal dynamics can be summarized as follows: (1) a late winter-early spring phytoplankton bloom followed by high zooplankton and macrobenthos densities during the spring months; (2) low-quality particulate suspended matter in summer, and an increase in the importance of zooplankton taxa with a wide range of feeding strategies, a decrease in macrofaunal abundance, and an increase in deposit-feeders and predators; and (3) a second phytoplankton bloom in autumn, followed by an increase in copepod density and a low macrofaunal abundance. In conclusion, pelagic and benthic communities in the coastal area of the Ligurian Sea mainly seem to be controlled bottom-up. Our results suggest that the quality of the particulate organic matter may play an important role in determining the temporal changes of both plankton and benthic assemblages, while the direct influence of other environmental features (such as sediment grain size) is relevant only for some macrobenthic taxa (e.g. crustaceans).     

Climate change effects on phytoplankton depend on cell size and food web structure

We investigated the effects of warming on a natural phytoplankton community from the Baltic Sea, based on six mesocosm experiments conducted 2005–2009. We focused on differences in the dynamics of three phytoplankton size groups which are grazed to a variable extent by different zooplankton groups. While small-sized algae were mostly grazer-controlled, light and nutrient availability largely determined the growth of medium- and large-sized algae. Thus, the latter groups dominated at increased light levels. Warming increased mesozooplankton grazing on medium-sized algae, reducing their biomass. The biomass of small-sized algae was not affected by temperature, probably due to an interplay between indirect effects spreading through the food web. Thus, under the higher temperature and lower light levels anticipated for the next decades in the southern Baltic Sea, a higher share of smaller phytoplankton is expected. We conclude that considering the size structure of the phytoplankton community strongly improves the reliability of projections of climate change effects.     

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern?

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the population's reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female^-1 d^-1, equivalent to 5.7% body C d^-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch^-1) vary with the age of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female^-1 clutch^-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.     

The influence of temperature on the development of Baltic Sea sprat (Sprattus sprattus) eggs and yolk sac larvae

In spring 2004 and 2005 we performed two sets of experiments with Baltic sprat (Sprattus sprattus balticus Schneider) eggs and larvae from the Bornholm Basin simulating ten different temperature scenarios. The goal of the present study was to analyse and parameterise temperature effects on the duration of developmental stages, on the timing of important ontogenetic transitions, growth during the yolk sac phase as well as on the survival success of eggs and early larval stages. Egg development and hatching showed exponential temperature dependence. No hatching was observed above 14.7°C and hatching success was significantly reduced below 3.4°C. Time to eye pigmentation, as a proxy for mouth gape opening, decreased with increasing temperatures from 17 days post hatch at 3.4°C to 7 days at 13°C whereas the larval yolk sac phase was shortened from 20 to 10 days at 3.8 and 10°C respectively. Maximum survival duration of non-fed larvae was 25 days at 6.8°C. Comparing the experimental results of Baltic sprat with existing information on sprat from the English Channel and North Sea differences were detected in egg development rate, thermal adaptation and in yolk sac depletion rate (YSDR). Sprat eggs from the English Channel showed significantly faster development and the potential to develop at temperatures higher than 14.7°C. North Sea sprat larvae were found to have a lower YSDR compared to larvae from the Baltic Sea. In light of the predictions for global warming, Baltic sprat stocks could experience improved conditions for egg development and survival.     

Role of cold resistance and burial for winter survival and spring initiation of an Ulva spp. (Chlorophyta) bloom in a eutrophic lagoon (Veerse Meer lagoon, The Netherlands)

In the eutrophic Veerse Meer lagoon (The Netherlands) large amounts of free-floating thalli from Ulva spp. are present from May to October. In winter however, no algae seem to occur in the lagoon. Sexual reproduction appears to be negligible, as spore formation and germling growth are observed only sporadically. Results of a field survey showed that in winter, viable Ulva biomass is present buried in the sediment of the shallow parts of the lagoon. Freezing experiments demonstrated that the algae are able to survive temperatures of −5 °C for 2 weeks when kept in darkness. In spring, the buried Ulva thalli are liberated out of the sediment to initiate a bloom. A field experiment indicates that bioturbation by the lugworm Arenicola marina does not stimulate the release of the thalli. Burial and winter survival can explain the rapid increase in Ulva biomass in spring and suggest that the initial spring biomass is one of the major factors determining the maximal biomass in summer. Received: 28 October 1997 / Accepted: 20 January 1998     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

黄海春季表层叶绿素和初级生产力及其粒径结构研究

根据2006年4月对黄海浮游植物分级叶绿素及初级生产力的调查,研究了黄海叶绿素及初级生产力的水平分布及粒级结构特征,并分析了其主要影响因素。黄海海域调查站位表层叶绿素a质量浓度变化范围为0.20～4.94μg·L-1,平均值为0.96μg·L-1。叶绿素最大值出现在临近长江口的站位。叶绿素分级结果表明黄海春季以粒径〉5μm的浮游植物占优势。黄海表层初级生产力的变化范围为2.03～15.64mg·m-3·h-1,平均值为6.08mg·m-3·h-1。其中南黄海海域初级生产力平均为6.58mg·m-3·h-1,北黄海海域初级生产力平均为4.92mg·m-3·h-1。高值区分布在南黄海中部。受水体透明度的影响,低值区出现在临近长江口的站位。断面站位分析表明浮游植物初级生产力由北向南逐步升高,温度随纬度的变化是南北海域初级生产力水平差异的主要原因。由于粒径较小（〈5μm）的浮游植物单位叶绿素具有较高的碳固定能力,调查期间整个海区初级生产力以粒径〈5μm的浮游植物贡献为主。     

Microphytoplankton in the Strait of Otranto (eastern Mediterranean)

The Strait of Otranto is the connection between the Adriatic and Ionian Seas. Low nutrient concentrations, high transparency, and low phytoplankton cell density and biomass reflect the oligotrophic character of the area. Enrichment of the euphotic layer with nutrients is mainly due to discharge of Albanian and Greek rivers, as well as mixing and upwelling in winter/early spring. Following phytoplankton bloom in April, a progressive decrease of phytoplankton cell density is due to the consumption of nutrients throughout the proceeding summer and autumn. Nitrogen was a strong limiting factor for phytoplankton growth in summer. Deep biomass maxima were detected in the 50 to 100 m layer and corresponded mostly to cells smaller than 20 m. The eastern part of the strait is mostly influenced by the northerly inflowing current from the Ionian Sea, and the western part by the southerly outflowing current from the Adriatic Sea. This typical circulation could be disturbed by inertial oscillations in the current field, generated by the strong oscillating winds and cyclonic eddies. The type of circulation determined the distribution of thermohaline characteristics, abundance, biomass, as well as taxonomic composition of phytoplankton, across the strait. Ecological characteristics of the water masses on two sides of the strait were significantly different during the formation of a longitudinal thermohaline front in May 1990.