Comparison of two methods to derive the size-structure of natural populations of phytoplankton

Various methods have been proposed to estimate the size structure of phytoplankton in situ , each exhibiting limitations and advantages. Two common approaches are size-fractionated filtration (SFF) and analysis of pigments derived from High Performance Liquid Chromatography (HPLC), and yet these two techniques have rarely been compared. In this paper, size-fractionated chlorophylls for pico- (View the MathML source<2μm), nano- (View the MathML source2–20μm) and micro-phytoplankton (View the MathML source>20μm) were estimated independently from concurrent measurements of HPLC and SFF data collected along Atlantic Meridional Transect cruises. Three methods for estimating size-fractionated chlorophyll from HPLC data were tested. Size-fractionated chlorophylls estimated from HPLC and SFF data were significantly correlated, with HPLC data explaining between 40 and 88% of the variability in the SFF data. However, there were significant biases between the two methods, with HPLC methods overestimating nanoplankton chlorophyll and underestimating picoplankton chlorophyll when compared with SFF. Uncertainty in both HPLC and SFF data makes it difficult to ascertain which is more reliable. Our results highlight the importance of using multiple methods when determining the size-structure of phytoplankton in situ, to reduce uncertainty and facilitate interpretation of data.

An intercomparison of bio-optical techniques for detecting dominant phytoplankton size class from satellite remote sensing

Satellite remote sensing of ocean colour is the only method currently available for synoptically measuring wide-area properties of ocean ecosystems, such as phytoplankton chlorophyll biomass. Recently, a variety of bio-optical and ecological methods have been established that use satellite data to identify and differentiate between either phytoplankton functional types (PFTs) or phytoplankton size classes (PSCs). In this study, several of these techniques were evaluated against in situ observations to determine their ability to detect dominant phytoplankton size classes (micro-, nano- and picoplankton). The techniques are applied to a 10-year ocean-colour data series from the SeaWiFS satellite sensor and compared with in situ data (6504 samples) from a variety of locations in the global ocean. Results show that spectral-response, ecological and abundance-based approaches can all perform with similar accuracy. Detection of microplankton and picoplankton were generally better than detection of nanoplankton. Abundance-based approaches were shown to provide better spatial retrieval of PSCs. Individual model performance varied according to PSC, input satellite data sources and in situ validation data types. Uncertainty in the comparison procedure and data sources was considered. Improved availability of in situ observations would aid ongoing research in this field.

Mechanisms of long-term decline in size of lesser sandeels in the North Sea explored using a growth and phenology model

The lesser sandeel Ammodytes marinus is a key species in the North Sea ecosystem, transferring energy from planktonic producers to top predators. Previous studies have shown a long-term decline in the size of 0-group sandeels in the western North Sea, but they were unable to pinpoint the mechanism (later hatching, slower growth or changes in size-dependent mortality) or cause. To investigate the first 2 possibilities we combined 2 independent time series of sandeel size, namely data from chick-feeding Atlantic puffins Fratercula arctica and from the Continuous Plankton Recorder (CPR), in a novel statistical model implemented using Markov Chain Monte Carlo (MCMC). The model estimated annual mean length on 1 July, as well as hatching date and growth rate for sandeels from 1973 to 2006. Mean length-at-date declined by 22% over this period, corresponding to a 60% decrease in energy content, with a sharper decline since 2002. Up to the mid-1990s, the decline was associated with a trend towards later hatching. Subsequently, hatching became earlier again, and the continued trend towards smaller size appears to have been driven by lower growth rates, particularly in the most recent years, although we could not rule out changes in size-dependent mortality. Our findings point to major changes in key aspects of sandeel life history, which we consider are most likely due to direct and indirect temperature-related changes over a range of biotic factors, including the seasonal distribution of copepods and intra- and inter-specific competition with planktivorous fish. The results have implications both for the many predators of sandeels and for age and size of maturation in this aggregation of North Sea sandeels.

A multicomponent model of phytoplankton size structure

Size-fractionated filtration (SFF) is a direct method for estimating pigment concentration in various size classes. It is also common practice to infer the size structure of phytoplankton communities from diagnostic pigments estimated by high-performance liquid chromatography (HPLC). In this paper, the three-component model of Brewin et al. (2010) was fitted to coincident data from HPLC and from SFF collected along Atlantic Meridional Transect cruises. The model accounted for the variability in each data set, but the fitted model parameters differed for the two data sets. Both HPLC and SFF data supported the conceptual framework of the three-component model, which assumes that the chlorophyll concentration in small cells increases to an asymptotic maximum, beyond which further increase in chlorophyll is achieved by the addition of larger celled phytoplankton. The three-component model was extended to a multicomponent model of size structure using observed relationships between model parameters and assuming that the asymptotic concentration that can be reached by cells increased linearly with increase in the upper bound on the cell size. The multicomponent model was verified using independent SFF data for a variety of size fractions and found to perform well (0.628 ≤ r ≤ 0.989) lending support for the underlying assumptions. An advantage of the multicomponent model over the three-component model is that, for the same number of parameters, it can be applied to any size range in a continuous fashion. The multicomponent model provides a useful tool for studying the distribution of phytoplankton size structure at large scales.

Absorption-based algorithm of primary production for total and size-fractionated phytoplankton in coastal waters

Most satellite models of production have been designed and calibrated for use in the open ocean. Coastal waters are optically more complex, and the use of chlorophyll a (chl a) as a first-order predictor of primary production may lead to substantial errors due to significant quantities of coloured dissolved organic matter (CDOM) and total suspended material (TSM) within the first optical depth. We demonstrate the use of phytoplankton absorption as a proxy to estimate primary production in the coastal waters of the North Sea and Western English Channel for both total, micro- and nano+pico-phytoplankton production. The method is implemented to extrapolate the absorption coefficient of phytoplankton and production at the sea surface to depth to give integrated fields of total and micro- and nano+pico-phytoplankton primary production using the peak in absorption coefficient at red wavelengths. The model is accurate to 8% in the Western English Channel and 22% in this region and the North Sea. By comparison, the accuracy of similar chl a based production models was >250%. The applicability of the method to autonomous optical sensors and remotely sensed aircraft data in both coastal and estuarine environments is discussed.

Meiobenthos and macrobenthos are discrete entities and not artefacts of sampling a size continuum: Comment on Bett (2013)

Species size distributions for metazoan benthic invertebrates conform to the highly conservative bimodal pattern, regardless of the sieve mesh sizes or numbers of sieves used in their extraction. This pattern is not an artefact of sampling a size continuum as suggested by computer simulations using just 2 fixed mesh sizes in Bett (2013; Mar Ecol Prog Ser 487:1-6). Meiobenthos and macrobenthos are coherent entities, each with a distinct suite of functional attributes, and should not be regarded as a single unit for ecological modelling purposes.

Effect of phytoplankton size classes on bio-optical properties of phytoplankton in the Western Iberian coast: Application of models

Chlorophyll-a satellite products are routinely used in oceanography, providing a synoptic and global view of phytoplankton abundance. However, these products lack information on the community structure of the phytoplankton, which is crucial for ecological modelling and ecosystem studies. To assess the usefulness of existing methods to differentiate phytoplankton functional types (PFT) or phytoplankton size classes from satellite data, in-situ phytoplankton samples collected in the Western Iberian coast, on the North-East Atlantic, were analysed for pigments and absorption spectra. Water samples were collected in five different locations, four of which were located near the shore and another in an open-ocean, seamount region. Three different modelling approaches for deriving phytoplankton size classes were applied to the in situ data. Approaches tested provide phytoplankton size class information based on the input of pigments data (Brewin et al., 2010), absorption spectra data (Ciotti et al., 2002) or both (Uitz et al., 2008). Following Uitz et al. (2008), results revealed high variability in microphytoplankton chlorophyll-specific absorption coefficients, ranging from 0.01 to 0.09 m2 (mg chl)− 1 between 400 and 500 nm. This spectral analysis suggested, in one of the regions, the existence of small cells (< 20 μm) in the fraction of phytoplankton presumed to be microphytoplankton (based on diagnostic pigments). Ciotti et al. (2002) approach yielded the highest differences between modelled and measured absorption spectra for the locations where samples had high variability in community structure and cell size. The Brewin et al. (2010) pigment-based model was adjusted and a set of model coefficients are presented and recommended for future studies in offshore water of the Western Iberian coast.

Mechanisms shaping size structure and functional diversity of phytoplankton communities in the ocean

The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.

Temporal changes in total and size-fractioned chlorophyll-a in surface waters of three provinces in the Atlantic Ocean (September to November) between 2003 and 2010

Phytoplankton total chlorophyll concentration (TCHLa) and phytoplankton size structure are two important ecological indicators in biological oceanography. Using high performance liquid chromatography (HPLC) pigment data, collected from surface waters along the Atlantic Meridional Transect (AMT), we examine temporal changes in TCHLa and phytoplankton size class (PSC: micro-, nano- and pico-phytoplankton) between 2003 and 2010 (September to November cruises only), in three ecological provinces of the Atlantic Ocean. The HPLC data indicate no significant change in TCHLa in northern and equatorial provinces, and an increase in the southern province. These changes were not significantly different to changes in TCHLa derived using satellite ocean-colour data over the same study period. Despite no change in AMT TCHLa in northern and equatorial provinces, significant differences in PSC were observed, related to changes in key diagnostic pigments (fucoxanthin, peridinin, 19′-hexanoyloxyfucoxanthin and zeaxanthin), with an increase in small cells (nano- and pico-phytoplankton) and a decrease in larger cells (micro-phytoplankton). When fitting a three-component model of phytoplankton size structure — designed to quantify the relationship between PSC and TCHLa to each AMT cruise, model parameters varied over the study period. Changes in the relationship between PSC and TCHLa have wide implications in ecology and marine biogeochemistry, and provide key information for the development and use of empirical ocean-colour algorithms. Results illustrate the importance of maintaining a time-series of in-situ observations in remote regions of the ocean, such as that acquired in the AMT programme.

Influence of light in the mixed-layer on the parameters of a three-component model of phytoplankton size class

Phytoplankton size structure is an important indicator of the state of the pelagic ecosystem. Stimulated by the paucity of in situ observations on size structure, and by the sampling advantages of autonomous remote platforms, new efforts are being made to infer the size-structure of the phytoplankton from oceanographic variables that may be measured at high temporal and spatial resolution, such as total chlorophyll concentration. Large-scale analysis of in situ data has revealed coherent relationships between size-fractionated chlorophyll and total chlorophyll that can be quantified using the three-component model of Brewin et al. (2010). However, there are variations surrounding these general relationships. In this paper, we first revise the three-component model using a global dataset of surface phytoplankton pigment measurements. Then, using estimates of the average irradiance in the mixed-layer, we investigate the influence of ambient light on the parameters of the three-component model. We observe significant relationships between model parameters and the average irradiance in the mixed-layer, consistent with ecological knowledge. These relationships are incorporated explicitly into the three-component model to illustrate variations in the relationship between size-structure and total chlorophyll, ensuing from variations in light availability. The new model may be used as a tool to investigate modifications in size-structure in the context of a changing climate.