A modified one-metre Friedinger sampler

SUMMARY. A 1-m water sampler is described together with results obtained from sampling a vertically stratified phytoplankton population of Ceratium hirundinella . These results were analysed using a one-way analysis of variance to partition the total variance into two components. Population estimates from 1-m sections taken sequentially from the surface waters to 5-m depth and integrated, were not significantly different ( P > 0.05) from those obtained using a 5-m vertical tube sampler: replicate sampling units taken with both samplers showed that Ceratium was contagiouly distributed in the vertical water column.     

Microscale patchiness of plankton within a sharp pycnocline

Microscale distributions of plankton around the pycnocline werestudied over a 24 h period in the southern Kattegat, using agradient sampler which collects 20 samples over a 3 m depthinterval. Moderately elevated concentrations of phyto- and zooplanktonwere observed at the pycnocline. Microscale variance was highestfor adult copepods. Nauplii and copepodites were equally wellrepresented by sampling with ordinary vertical 5 1 water bottlesas with the horizontal 1.5 1 bottles of the gradient sampler.Adult copepods were underestimated by the vertical bottles.No vertical migration of dinoflagellates was observed over the3 m interval covered by the gradient sampler. Microscale correlationsbetween copepods and phytoplankton within the gradient samplerwere weak. Copepodites (mainly Oithona sp.) and the dinoflagellateProrocentrum micans showed the best correlation.     

A multi-purpose modular limnological sampler

This paper describes a new type of multi-purpose limnological sampler, based on several modules which, when fitted together in various ways, and with accessory items, can be used to sample in many different fashions. The main advantages of the sampler are that, in addition to its ability to perform a variety of functions, it is both cheap and light. It is particularly suitable for multi-purpose work on remote waters to which transport of heavy or bulky equipment is difficult or expensive. The complete package described, best operated by two people, currently costs £ 75 and weighs 20 kg. It enables the operators to sample several aspects of water, sediments, phytoplankton, macrophytes, zooplankton, zoobenthos and fish in several ways, both qualitatively and quantitatively, from a boat or from the shore.     

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.     

A segmented pipe sampler for integrated profiling of the upper water column

The segmented integrating pipe sampler (SIPS) samples a continuousvertical profile of physical and biological properties of theupper water column. Composite or detailed samples can be collectedfor analysis of phytoplankton abundance and distribution. Thissampling system requires no power source and can be operatedfrom a small boat or platform in a variety of aquatic habitats.The correlation coefficient calculated for the phytoplanktonabundance sampled by the SIPS and pump was 0.900, while thecoefficient for the SIPS and Niskin bottle comparison was 0.797.     

AN OPTICAL DETECTION SYSTEM FOR THE STUDY OF FINE SCALE VERTICAL DISPLACEMENT OF MICROALGAE IN AN ARTIFICIAL WATER COLUMN

A proper knowledge of the vertical organization of the phytoplankton is of fundamental importance for our understanding of the functions of pelagic ecosystems. Essential in this context is the existence of vertical gradients in environmental parameters. However, little is known about how the fine vertical structures of phytoplankton species are formed and maintained. In situ study of phytoplankton is biased by the fact that submersing instruments can disturb or even destroy the fine vertical gradients in species composition and/or cell numbers. We have designed and constructed an optical instrumentation system by which fine-scale vertical displacements of microalgae can be studied in an artificial water column without influencing fine physical, chemical, and biological structures of the water column. This enables us to find out more about the fine-scale behavioral responses of microalgae to vertical gradients in environmental parameters. We describe the main system, present some test results, and conclude that our optical system is able to reveal fine-scale vertical displacements of microalgae in an artificial water column and that the system can detect differences in cell densities down to 100 cells·mL⁻¹.     

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.     

Seasonal variation in the diel vertical distribution of the migratory alga Cryptomonas marssonii (Cryptophyceae) in a small,highly humic lake

The diel vertical distribution patterns of a migratory alga Cryptomonas marssonii in a small, steeply stratified humic lake were investigated during a summer season (10 diurnal experiments between May and September) using a close-interval Blakar-type sampler. The results indicate that the cells were phototactic; they were typically concentrated at the surface or subsurface during daylight, while in darkness the highest densities were recorded in deeper water, usually near the upper limit of anoxia. During a dense blue-green bloom in August the cells of C. marssonii were also concentrated by day into the same water layer, where oxygen was depleted. However, the cells seemed to avoid totally anoxic water. Because the vertical distribution pattern of C. marssonii had special diurnal and seasonal characteristics, care is needed when designing a sampling programme for a phytoplankton population dominated by this species.     

A portable, low-cost, multipurpose, surface-subsurface plankton sampler

This note describes a portable, low-cost, pump-based, multi-purpose,surface–subsurface, phytoplankton and zooplankton sampler.It is self-powered by a small 12 V DC battery, and is portableand ideal for use from a small boat in a river, estuary, orsea. The sampler is equipped with a very sensitive microprocessor-controlledflow sensor for precise determination of the volume of waterprocessed by the sampler. A study was carried out to evaluatethe performance of this sampler in comparison to the conventionaltowed plankton net. The results of this study suggest that thispump-based sampler is as efficient in collecting plankton asusing conventional plankton nets, with little damage to theorganisms.     

Critical conditions for phytoplankton blooms

We motivate and analyse a reaction—advection—diffusion model for the dynamics of a phytoplankton species. The reproductive rate of the phytoplankton is determined by the local light intensity. The light intensity decreases with depth due to absorption by water and phytoplankton. Phytoplankton is transported by turbulent diffusion in a water column of given depth. Furthermore, it might be sinking or buoyant depending on its specific density. Dimensional analysis allows the reduction of the full problem to a problem with four dimensionless parameters that is fully explored. We prove that the critical parameter regime for which a stationary phytoplankton bloom ceases to exist, can be analysed by a reduced linearized equation with particular boundary conditions. This problem is mapped exactly to a Bessel function problem, which is evaluated both numerically and by asymptotic expansions. A final transformation from dimensionless parameters back to laboratory parameters results in a complete set of predictions for the conditions that allow phytoplankton bloom development. Our results show that the conditions for phytoplankton bloom development can be captured by a critical depth, a compensation depth, and zero, one or two critical values of the vertical turbulent diffusion coefficient. These experimentally testable predictions take the form of similarity laws: every plankton—water—light-system characterized by the same dimensionless parameters will show the same dynamics.     

The kinetics of algal photoadaptation in the context of vertical mixing

The responses of phytoplankton to turbulent motions in the surfacemixed layer can be measured to estimate the rate of verticalmixing. If the time scale for the response (photoadaptation)is shorter than that for vertical mixing, phytoplankton willexhibit a vertical gradient associated with adaptation to ambientlight, whereas if mixing occurs with a time scale shorter thanthat of photoadaptation, the surface mixed layer will be uniformwith respect to the photoadaptive parameter. To examine thephysiological bases for a model of vertical mixing and photoadaptation,we grew the marine diatom Thalassiosira pseudonana (clone 3H)at three photon flux densities and subjected the cultures toreciprocal light shifts, measuring physiological and chemicalchanges over the following 10 h. Several parameters, easilymeasured in nature and attributable primarily to phytoplankton,responded to fluctuating light on different time scales. Aftercultures were exposed to relatively bright light, both the initialslope of the photosynthesis-irradiance curve and in vivo fluorescencewere depressed on a time scale of less than an hour. Photosyntheticcapacity was also reduced transiently, but recovered over manyhours to a high level characteristic of an adapted state. First-orderkinetics (the current model of choice for describing photoadaptation)reasonably described the rapid responses of phytoplankton tobright light, but other parameters (i.e. cellular chemical compositionand photosynthetic capacity) changed as a result of unbalancedgrowth and required much longer to adapt from low to high lightas compared to from high to low light. A logistic model of thisadaptation is presented. The model suggests that hysteresisof adaptation during vertical mixing may have important consequences.The vertical distributions of photoadaptive properties in mixedlayers not only reveal the rate of vertical mixing, but showhow phytoplankton integrate environmental fluctuations.     

ANTARCTIC AQUATIC ECOSYSTEMS AS HABITATS FOR PHYTOPLANKTON

1. The Southern Ocean is a large-scale, relatively homogeneous upwelling ecosystem whose phytoplankton apparently grows suboptimally over much of its area. By contrast there is a wide variety of freshwater habitats in the Antarctic and in some of these phytoplankton growth efficiency is very high. The two habitats share similar temperature and irradiance regimes, but differ markedly in availability of inorganic nutrients, in grazing pressure and in the time- and space-scales on which various physical processes act. 2. Concentrations of inorganic nutrients in the marine ecosystem have been represented as being in excess of phytoplankton requirements, but the ionic composition of some nutrient pools may not conform to phytoplankton preferences. 3. Nutrient-limitation determines phytoplankton production in Antarctic lakes and gives rise to gross differences between lakes. 4. Irradiance in the water column varies greatly over the year in both marine and freshwater ecosystems. Most algae are shade-adapted, with the ability to utilize low irradiance but with sub-optimal response to high irradiance. However, local phytoplankton maxima may attain very high carbon fixation and growth rates. 5. Consistently low temperatures characterize both systems. Their effects on photo-synthetic carbon uptake mirror shade-adaptation. Division rates of marine phytoplankton may however be very much higher than predicted for ambient temperatures. 6. Vertical mixing is important in both ecosystems and influences the environment experienced by phytoplankton cells. This appears to have little effect on the average performance of phytoplankton in the strongly mixed surface water column of the Southern Ocean, where the mixed depth may exceed 100 m. This can be related partly to the shade-adapted photosynthetic response. Euphotic depths range from 20 to 100 m. 7. Strong vertical mixing under ice-free conditions in lakes may maximize photosynthetic efficiency, whilst distinct vertical stratification in permanently ice-covered lakes gives rise to segregation of nutrient uptake and regeneration. 8. Physical removal of phytoplankton biomass by grazing is locally important in the Southern Ocean, in contrast to the estimated mean mesoscale impact of grazing. Vertical sedimentation losses appear important in the context of mixing depth and generation time, and may be modified by vertical circulation of water. 9. Loss of phytoplankton biomass from lakes during the ice-free period is dominated by physical removal via the lake outflow. Grazing is generally unimportant, except where larvae of otherwise nektobenthic zooplankton hatch in synchrony with a phytoplankton maximum. Sedimentation is important under ice-cover.     

Species Richness and Invasion Vectors: Sampling Techniques and Biases

During a European Union Concerted Action study on species introductions, an intercalibration workshop on ship ballast water sampling techniques considered various phytoplankton and zooplankton sampling methods. For the first time, all the techniques presently in use worldwide were compared using a plankton tower as a model ballast tank spiked with the brine shrimp and oyster larvae while phytoplankton samples were taken simultaneously in the field (Helgoland Harbour, Germany). Three cone-shaped and 11 non-cone shaped plankton nets of different sizes and designs were employed. Net lengths varied from 50 to 300 cm, diameters 9.7–50 cm, and mesh sizes 10–100 μm. Three pumps, a Ruttner sampler, and a bucket previously used in ballast water sampling studies were also compared. This first assessment indicates that for sampling ballast water a wide range of techniques may be needed. Each method showed different results in efficiency and it is unlikely that any of the methods will sample all taxa. Although several methods proved to be valid elements of a hypothetical `tool box' of effective ship sampling techniques. The Ruttner water sampler and the pump P30 provide suitable means for the quantitative phytoplankton sampling, whereas other pumps prevailed during the qualitative trial. Pump P15 and cone-shaped nets were the best methods used for quantitative zooplankton sampling. It is recommended that a further exercise involving a wider range of taxa be examined in a larger series of mesocosms in conjunction with promising treatment measures for managing ballast water. This revised version was published online in July 2006 with corrections to the Cover Date.     

<Emphasis Type="Italic">In situ</Emphasis> Quantification of Phytoplankton in Reservoirs Using a Submersible Spectrofluorometer

A submersible in situ spectrofluorometer, which permits the differentiation of four algal groups (green algae, diatoms, cryptophytes and cyanobacteria), was used for phytoplankton monitoring in five reservoirs with varying levels of eutrophication and composition of their phytoplankton communities. Data obtained in situ were compared to standard laboratory methods for phytoplankton quantification; concentration of chlorophyll a and microscopy analysis. A high correlation (r = 0.95, n = 96) between chlorophyll a levels using different methods was found in all types of phytoplankton community. Taxonomic analyses and cell counts were closely related to the ratio of algal classes measured by the in situ spectrofluorometer. The submersible device used in the study measures in a continuous mode, which is advantageous in comparison with discrete sampling. This method appears to be a good tool for water quality management and can be used in the detection of natural horizontal and vertical variability in phytoplankton communities or for the early detection of cyanobacterial blooms. The device used in this study is recommended as a screening tool that enables more effective sampling that can be focused on the localities and depths where changes in phytoplankton composition occur.     

Vertical distribution and diel migration of flagellated phytoplankton in a small humic lake

The vertical distributions and migrations are described of the most abundant flagellated phytoplankton species from the summer community of a small forest lake in southern Finland. The lake showed a steep and stable thermal stratification with a shallow oxygenated epilimnion. Horizontal variation of phytoplankton distribution within the lake was tested on two scales and found to be statistically significant only in the case of Mallomonas reginae. The vertical distribution of flagellated phytoplankton was assessed by reference to the distribution of a non-motile, neutrally buoyant species Ankyra judayi. Statistically significant, active vertical positioning was demonstrated for all the flagellates examined with the exception of Spiniferomonas bourrellyi. Diel vertical migrations were apparent for all species showing active positioning and the pattern of an evening descent and a morning ascent was ubiquitous. The extent and timing of diel migrations varied between species. The most extensive migrations were by Cryptomonas marssonii which crossed a temperature gradient of 14 °C and penetrated far into the anoxic hypolimnion. Several categories of competitive advantage can be gained by species undertaking such diel vertical migrations.     

Depth integrating samplers for use in limnocorrals

Constructional details of a zooplankton and a phytoplankton sampler are given. Both samplers are designed for use in shallow limnocorrals (5 m deep enclosures) and both give samples integrated with respect to depth and offer a considerable saving in time over other sampling methods.     

Stepping back to improve sprint performance: a kinetic analysis of the first step forwards

Using a step backward to initiate forward movement can increase force and power at push-off and improve sprint performance over short distances. However, it is not clear whether the benefit provided by this paradoxical step influences the mechanics of the first step forwards. Twenty-seven men of an athletic background performed maximal effort 5-m sprints from a standing start and employed a step forwards (parallel and split stance) or backwards (false) to initiate movement. Each sprint was started with an audio cue that also activated the timing gates. Three trials of each starting style were performed and movement (0 m), 2.5-, and 5-m times were recorded. An in-ground force plate placed at the 0-m mark measured the kinetic and temporal characteristics of the first step. Sprint times to 2.5 and 5 m were slower (p < 0.05) when a parallel start was used. No differences were seen in the normalized peak forces (vertical and horizontal) or the vertical impulse between starts, but the vertical mean force was 11 and 12% higher for the false and split starts, respectively. Surprisingly, the parallel start's impulse was significantly greater than that of the false (24%) and split (22%) styles, a consequence of the additional time spent in contact with the ground. The ground contact time, time to peak force, and time from peak force to toe-off (vertical and horizontal) were significantly longer for the parallel start. These temporal variables were also better correlated with sprint performance than any kinetic measure (0.42 ≤ r ≤ 0.75). The false start appears to be advantageous over short distances by improving push-off and the temporal characteristics of the first step.     

Thermal stability and phytoplankton distribution

Thermal stability is the potential of water columns to mix, and has long been known to fundamentally influence the vertical and temporal distribution of phytoplankton. Essentially this is because it indirectly controls the amount of light available to phytoplankton.Under stable conditions of strong temperature gradients algal species (or assemblages of associated species) distribute vertically because they have sufficient time to exploit the attenuated light field at their preferred depths. This encourages a species diversity which, in the Southern Hemisphere, is especially exemplified by the extremely stable conditions under the permanent ice of Antarctic lakes.In other lakes stability commonly encourages growth of blue-green algae by permitting their positive buoyancy to place them in optimal light conditions, and by inhibiting the resuspension of competing non-buoyant species. Analogous patterns occur with motile species (Dinophyceae, Cryptophyceae, etc.), and with non-motile forms whose physiological adaptations allow growth to large sub-surface peaks at preferred depths. These sub-surface maxima can be upwelled to the water surface, in a manner controlled by thermal stability and vertical shear, and horizontally transported to give large variations in horizontal distribution.At all latitudes diel stability cycles in surface waters can affect physiological properties important for growth, and in some circumstances can dominate the phytoplankton dynamics and distribution.Such short-term stability events merge with longer-term (e.g. annual) events with no conceptual distinction. A modern way to integrate this continuity is by scaling using spectral analysis of cyclicity. This allows biological variables (algal biomass, numbers, production) to be stochastically related to indices of stability (e.g. Brunt-Väisälä frequency).     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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A new sampler for collecting invertebrates in submerged vegetation

A new sampler is described that can rapidly enclose a specific volume of water at a given depth, including submerged macrophytes and associated invertebrates. The sampler is a tong-shaped instrument consisting of two metal rods connected by a flexible joint. At the end of each arm is a metal cylinder (diameter 10 cm); one end of the cylinder has a sharpened edge and the other is equipped with a net. Powerful metal springs force the cylinders together at high speed, and macrophytes are enclosed in the nets. The sampler is suitable for studying vertical and horizontal distribution of invertebrates, even fast-moving taxa, within submerged vegetation. The device causes little disturbance of the vegetation, hence it is also suitable for repeated sampling within mesocosms.