A coupled 1-D biological/physical model of the northeast subarctic Pacific Ocean with iron limitation

The recent NE subarctic Pacific study of the Canadian JGOFS project was designed primarily to address why phytoplankton biomass and production at Ocean Station Papa (OSP: 50°N, 145°W) are not as high as the nitrate concentrations could potentially support. To examine the possible role of iron (Fe) limitation in concert with microzooplankton grazing and physical supply of nitrate, we have coupled a four-compartment Nitrogen–Phytoplankton–Zooplankton–Detritus planktonic ecosystem model with a 60-layer (each 2 m thick) one-dimensional mixed-layer model (Mellor–Yamada level 2.5), driven by annual forcing characteristic of OSP. Both the physical and ecological models are forced with the same annual heat budget, mean phytoplankton concentration was tuned with the equilibrium solution of the model, and the zooplankton parameter values were chosen to be representative of microzooplankton. Modelled sea surface temperature ranged between 6 (fixed – late winter) and 13–14°C, depending on the distribution and amount of phytoplankton and detritus calculated by the model. Simulations with Fe limitation reducing the maximum specific growth rate of phytoplankton (for Fe-replete conditions) by a factor of ∼3 best reproduced the annual cycle of surface layer nitrate, although the resulting annual f-ratio calculated from the fluxes into and out of the nitrogen compartment was marginally higher than recent estimates of f-ratio based on observations at OSP. The best simulations with Fe limitation agreed with observations of the annual cycle of surface nitrate concentration, the f-ratio, particulate nitrogen concentration in the euphotic layer, the export production, and the remineralization depth scale for sinking detritus, to within ∼50%, probably within the range of observational uncertainty and/or seasonal and interannual variability. Possible modifications include separating the detrital pool into suspended and sinking organic matter, decreasing the rate of remineralization with increasing depth, and examining the supply of nitrate to the surface layer by means of horizontal advection. The observational basis required to formulate these processes is marginal at present.     

Inverse modeling of carbon and nitrogen flows in the pelagic food web of the northeast subarctic Pacific

We use inverse analysis to model carbon and nitrogen flows in the upper ocean food web at Ocean Station Papa (OSP; 50°N, 145°W) for winter, spring, and late summer. The seasonal variability in basic physical, chemical, and biological characteristics is low, and the particulate carbon and nitrogen flux at 200 m is remarkably constant. Despite this apparent uniformity, the food web structure undergoes significant seasonal changes. The diversity of trophic pathways is higher during late summer than during the other two periods. The spring ecosystem is not in steady state and undergoes net phytoplankton growth and macronutrient consumption. The microbial loop is well developed only during late summer. Nevertheless, ammonium regeneration by the food web seems insufficient to meet demand by the primary producers. The difference may be due to recycling of dissolved organic nitrogen (urea+free amino acids), a process not represented in the model. The winter food web is the closest to steady state, with nitrate utilisation approximately in balance with export of particulate nitrogen. The inverse analysis suggests two main seasonally invariant features of the NE Pacific ecosystem. First, the major trophic pathway is always from picophytoplankton (0.2–5 μm) to microzooplankton (heterotrophic dinoflagellates and ciliates) to mesozooplankton. This supports the idea of a strong coupling between the microbial and metazoan food webs. Second, much of the primary production (and bacterial production in late summer) is not grazed and is recycled through the detrital pool. Both these features seem to arise from the requirement to conserve nitrogen as well as carbon in the food web. More complete measurements on the microzooplankton 20–200 μm in size, including the small metazoans like nauplii larvae, are required to improve the models presented here.     

Evidence of change in the winter mixed layer in the Northeast Pacific Ocean

Sea-surface temperatures in the Northeast Pacific Ocean show a warming trend, and salinities show a declining trend, in data collected over the last 60 years. These changes combine to reduce the density of the surface layer over a large area of the Northeast Pacific. The declining surface density changes the energetic requirements for the formation of a surface mixed layer, and observations at Ocean Station Papa indicate that mid-winter mixed layer depths are showing a marked decline. The reduction in the depth of penetration of the winter-time mixed layer should reduce the nutrients entrained into the upper ocean each winter. Observations suggest that near surface nutrient levels are declining at Papa but remain well above levels that might inhibit productivity. However, at present the productivity of large phytoplankton appears to be limited by iron supply which is thought to be mainly from the atmosphere. A shallower mixed layer depth could increase the concentration of iron in this layer. The increase in iron would increase the utilization of nitrate, mainly by diatoms, and new production and the f ratio would increase.     

Modelling the response of the planktonic food web to iron fertilization and warming in the NE subarctic Pacific

A one-dimensional ecosystem model with two explicit size classes of phytoplankton was developed for the NE subarctic Pacific to investigate variations in the export of organic particles to the ocean interior due to potential changes in the environment. Specifically, the responses of the planktonic ecosystem to permanent removal of iron limitation and to warming (of 2 and 5 °C) were explored. The ecosystem model consists of five components (small and large phytoplankton, microzooplankton, detritus and nitrogen), and includes grazing by mesozooplankton that varies in time according to long-term observations at Ocean Station Papa (OSP). The model addresses the role of iron limitation on phytoplankton growth and includes temperature dependence of physiological rates. The ecosystem model was forced with annual wind and solar heating from OSP. The model best reproduced the low chlorophyll high nitrate conditions of the NE subarctic Pacific when both small and large phytoplankton were limited by iron such that their maximum specific growth rate was reduced by 10 and 70%, respectively. Sensitivity analysis showed that model results depended on the value of the iron limitation parameter of large phytoplankton (L_Fe-L) and the grazing parameters of micro- and mesozooplankton. To explore the effect of iron limitation, simulations were carried out varying the iron limitation parameters while maintaining the nitrogen flux at the base of the model constant and the grazing pressure by mesozooplankton unchanged. In the warming case, simulations were carried out increasing ocean temperatures by 2° and 5 °C applied only to the ecological components, the flux of nitrate at the base of the model was increased to obtain a steady annual cycle, and grazing by mesozooplankton remained constant. When compared with the standard case, model simulations indicated that both permanent removal of iron limitation and warming cause changes in food web structure and the carbon cycle. The response was more dramatic in the iron-replete case where the phytoplankton community structure in spring changed from one dominated by pico- and nanoplankton to one dominated by large phytoplankton, and primary production increased until it consumed all the external nutrient (N) supply to the upper layer. However, reducing iron deficiency actually led to lower annual primary production due to a decrease in the regeneration of nitrogen in the euphotic zone. These changes in food web structure influenced the magnitude, composition and seasonal cycle of sinking particles.     

THE MAXIMUM VALUE OF VERTICAL DISTRIBUTION OF DISSOLVED OXYGEN IN THE SOUTH CHINA SEA

This paper indicates that there is a maximum value of vertical distribution of dissolved oxygen in the South China Sea in summer, and that this maximum value in summer derives mainly from winter. A similar reservation occurs also in temperature under the thermocline. Above and below the depth of the oxygen maximum, the oxygen content is decreased by an increase of temperature, the biological respiration and the decomposition of organic matter respectively. The oxygen maximum usually occurs above 50 meters in depth where the phytoplankton presents in small amounts. The phytoplankton is only an influentical factor for the oxygen maximum, not a decisive one.     

Nutrients in an oligotrophic boundary current: Evidence of a new role for the Leeuwin Current

New observations along the continental shelf of Western Australia provide a novel explanation for the established ∼60 years relationship between Leeuwin Current (LC) strength and greater winter nitrate concentrations at 32°S plus the inter-annual variation in the magnitude of the annual, shelf-scale, phytoplankton bloom. The potential source of dissolved nitrogen to support the annual shelf scale phytoplankton bloom was identified as thin layers of an unprecedented areal extent, nitrate concentration and shallow nature that were observed off the northwest of Australia. We propose that the dissolved inorganic nitrogen (DIN) in these layers enters the LC at depth and then enters the euphotic zone via by three mechanisms: instability that results in a warm core eddy, cooling that deepens the surface mixed layer and shallowing of the thin layer. During the onset of the annual phytoplankton bloom along the west coast of Australia from 22°S to 34°S the poleward flowing LC was clearly evident as a surface intensified ocean boundary current transporting warmer, lower-salinity, greater-silicate waters in a shallow mixed layer rapidly southward. Between 24 and 26°S the core of the LC was present as a 50–100 m deep layer over one or more thin layers, 15–50 m thick, with high nitrate and low dissolved oxygen (DO). These layers were of lower salinity, cooler water with markedly reduced DO, high nitrate concentrations and distinct nitrate:silicate (NO₃:Si(OH)₄) nutrient ratios. As the LC flowed south it cooled and deepened thereby entraining the thin layers of high nitrate water into the euphotic zone. The LC also formed large (greater than 100 km diameter) warm core eddies with a deep surface mixed layer that also entrained nitrate from these thin layers. In some locations as far south as 32°S the LC was still present with the thin layer of high nitrate intact but now within the euphotic zone. Thus, the available evidence suggests the LC arises under conditions that favour rapid and shallow nitrification. This nitrification fuels a shelf-scale bloom on a downwelling favourable coast. Depending upon the rate of nitrification the source of the particular organic matter may be local or delivered from the tropics via horizontal advection in a subsurface layer of the LC.     

Sedimentary and water column processes in the Oyster Grounds: a potentially hypoxic region of the North Sea

The purpose of this research was to investigate the potential causes of low oxygen levels in the bottom water of the Oyster Grounds region of the shallow southern North Sea, an area which provides suitable conditions for low oxygen levels to develop. At the end of the summer stratified period, relevant biogeochemical processes were investigated using a combination of sedimentary and water column rate measurements. Phytoplankton nitrate and ammonium uptake was measured throughout the water column using (15)N labelled isotopes and showed ammonium uptake dominated in the upper and bottom mixed layer with a maximum 294.4 micromol N m(-3)h(-1). In the deep chlorophyll maximum at the thermocline, primary production was dominated by nitrate uptake, with an average of 35.0 micromol N m(-3)h(-1), relative to ammonium uptake, with an average of 24.6 micromol N m(-3)h(-1). This high relative nitrate uptake will in part result in exportable new production to the isolated bottom mixed layer and sediments, as opposed to regenerated ammonium driven uptake. This biomass export was indicated by significant benthic oxygen consumption rates in the stratified region (782-1275 micromol O(2)m(-2)h(-1)micromol N m(-3)h(-1)) long after the end of the spring bloom. The sediments were also an active net source of nitrate, ammonium, phosphate and silicate into the bottom mixed layer of 4.4, 8.4, 2.3 and 68.8 micromol m(-2)h(-1), respectively. The export of new production within the thermocline to the bottom mixed layer and the consequent sediment oxygen consumption in the isolated bottom mixed layer in the Oyster Grounds are expected to have contributed to the low bottom water oxygen concentrations of 2.07 mg l(-1) (64.7 micromol l(-1)) measured. The long stratified period associated with this low oxygen is predicted to occur more regularly in the future and continued monitoring of this ecologically important region is therefore essential if the causes of these potentially damaging low oxygen levels are to be fully understood.     

A general ecosystem model for applications to primary productivity and carbon cycle studies in the global oceans

We have developed a general 1-D multi-component ecosystem model that incorporates a skillful upper ocean mixed layer model based on second moment closure of turbulence. The model is intended for eventual incorporation into coupled 3-D physical–biogeochemical ocean models with potential applications to modeling and studying primary productivity and carbon cycling in the global oceans as well as to promote the use of chlorophyll concentrations, in concert with satellite-sensed ocean color, as a diagnostic tool to delineate circulation features in numerical circulation models. The model is nitrogen-based and the design is deliberately general enough and modular to enable many of the existing ecosystem model formulations to be simulated and hence model-to-model comparisons rendered feasible. In its more general form (GEM10), the model solves for nitrate, ammonium, dissolved nitrogen, bacteria and two size categories of phytoplankton, zooplankton and detritus, in addition to solving for dissolved inorganic carbon and total alkalinity to enable estimation of the carbon dioxide flux at the air–sea interface. Dissolved oxygen is another prognostic variable enabling air–sea exchange of oxygen to be calculated. For potential applications to HNLC regions where productivity is constrained by the availability of a trace constituent such as iron, the model carries the trace constituent as an additional prognostic variable. Here we present 1-D model simulations for the Black Sea, Station PAPA and the BATS site. The Black Sea simulations assimilate seasonal monthly SST, SSS and surface chlorophyll, and the seasonal modulations compare favorably with earlier work. Station PAPA simulations for 1975–1977 with GEM5 assimilating observed SST and a plausible seasonal modulation of surface chlorophyll concentration also compare favorably with earlier work and with the limited observations on nitrate and pCO₂ available. Finally, GEM5 simulations at BATS for 1985–1997 are consistent with the available time series. The simulations suggest that while it is generally desirable to employ a comprehensive ecosystem model with a large number of components when accurate depiction of the entire ecosystem is desirable, as is the prevailing practice, a simpler formulation such as GEM5 (N²PZD model) combined with assimilation of remotely sensed SST and chlorophyll concentrations may suffice for incorporation into 3-D prediction models of primary productivity, upper ocean optical clarity and carbon cycling.     

An Ecosystem Model Including Nitrogen Isotopes: Perspectives on a Study of the Marine Nitrogen Cycle

We have developed an ecosystem model including two nitrogen isotopes (¹⁴N and ¹⁵N), and validated this model using an actual data set. A study of nitrogen isotopic ratios (δ¹⁵N) using a marine ecosystem model is thought to be most helpful in quantitatively understanding the marine nitrogen cycle. Moreover, the model study may indicate a new potential of δ¹⁵N as a tracer. This model has six compartments: phytoplankton, zooplankton, particulate organic nitrogen, dissolved organic nitrogen, nitrate and ammonium in a two-box model, and has biological processes with/without isotopic fractionation. We have applied this model to the Sea of Okhotsk and successfully reproduced the δ¹⁵N of nitrate measured in seawater and the seasonal variations in δ¹⁵N of sinking particles obtained from sediment trap experiments. Simulated δ¹⁵N of phytoplankton are determined by δ ¹⁵N of nitrate and ammonium, and the nitrogen f-ratio, defined as the ratio of nitrate assimilation by phytoplankton to total nitrogenous nutrient assimilation. Detailed considerations of biological processes in the spring and autumn blooms have demonstrated that there is a significant difference between simulated δ¹⁵N values of phytoplankton, which assimilates only nitrate, and only ammonium, respectively. We suggest that observations of δ ¹⁵N values of phytoplankton, nitrate and ammonium in the spring and autumn blooms may indicate the ratios of nutrient selectivity by phytoplankton. In winter, most of the simulated biogeochemical fluxes decrease rapidly, but nitrification flux decreases much more slowly than the other biogeochemical fluxes. Therefore, simulated δ¹⁵N values and concentrations of ammonium reflect almost only nitrification. We suggest that the nitrification rate can be parameterized with observations of δ¹⁵N of ammonium in winter and a sensitive study varying the parameter of nitrification rate.     

Adsorption—desorption control of phosphate in anoxic sediment of a coastal sea,Funka Bay,Japan

Sediment core samples were taken once a month from July 1980 to September 1981 at a station in Funka Bay (92-m depth) for the determination of phosphate, silicate and alkalinity in interstitial water. A remarkable seasonal variation was found for interstitial phosphate, that is, distinct maxima appeared in spring (March—April), just after a phytoplankton bloom which brought a large amount of settling particles to the bottom, and in summer (July—August) when the water was stratified and the dissolved oxygen content of the bottom water decreased due to the decomposition of organic matter. The high interstitial phosphate concentration was always accompanied by a sharp increase in alkalinity, indicating sulfate reduction. This large seasonal variation in interstitial phosphate cannot be explained by in situ decomposition of organic matter and/or the diffusive loss of interstitial phosphate. A more likely explanation is adsorption and desorption of interstitial phosphate coincident with the depth of the active sulfate reduction layer.     

黄海溶解氧垂直分布最大值的成因

本文对黄海溶解氧垂直分布中最大值的成因进行了探讨．认为：浮游植物春花期光合作用产生的大量氧，是溶解氧垂直分布最大值得以在春季（5月）形成的基础；而良好的温、密跃层的存在是氧最大值得以形成的必要条件，夏季叶绿素最大值层中较强的光合作用，是氧最大值在夏季得以维持甚至增强的主要原因．提出：黄海春、夏季溶解氧垂直分布中的最大值并非主要由冬季保持而来．     

马里亚纳海沟“挑战者深渊”营养盐的垂直分布特征

2015年12月在马里亚纳海沟"挑战者深渊"进行了定点样品采集,对温度、盐度、溶解氧、pH等环境参数进行了分析,讨论了营养盐的垂直分布特征、各形态营养盐结构特征及影响因素。研究发现,溶解氧在表层具有最大值,在1000 m左右出现极小值,而在8700 m深度具有较高溶解氧值(5.79 mg·L^-1),这可能与富氧水团的存在有关。硝酸盐表层含量较低,在1000和5367 m处出现双峰值。在表层水体中,溶解有机氮、磷是溶解总氮、溶解总磷的主要存在形式,表层以深,溶解无机氮、磷逐渐占据主导地位。磷酸盐表层含量最低,在1000 m处达到最大值,之后随着深度的增加浓度逐渐降低;硅酸盐在表层含量较低,在约4000 m处有最大值161.65μmol·L^-1,在4000 m以深,硅酸盐仍维持较高浓度。结果表明马里亚纳海沟"挑战者深渊"的溶解氧、pH及营养盐的垂直分布特征与大洋环流、海沟形态以及生物活动密切相关。     

Wintertime rates of net primary production and nitrate and ammonium uptake in the southern Benguela upwelling system

The elevated levels of primary productivity associated with eastern boundary currents are driven by nutrient- rich waters upwelled from depth, such that these regions are typically characterised by high rates of nitrate-fuelled phytoplankton growth. Production studies from the southern Benguela upwelling system (SBUS) tend to be biased towards the summer upwelling season, yet winter data are required to compute annual budgets and understand seasonal variability. Net primary production (NPP) and nitrate and ammonium uptake were measured concurrently at six stations in the SBUS in early winter. While euphotic zone NPP was highest at the stations nearest to the coast and declined with distance from the shore, a greater proportion was potentially exportable from open-ocean surface waters, as indicated by the higher specific nitrate uptake rates and f-ratios (ratio of nitrate uptake to total nitrogen consumption) at the stations located off the continental shelf. Near the coast, phytoplankton growth was predominantly supported by ammonium despite the high ambient nitrate concentrations. Along with ammonium concentrations as high as 3.6 µmol l^–1, this strongly suggests that nitrate uptake in the inshore SBUS, and by extension carbon drawdown, is inhibited by ammonium, at least in winter, although this has also been hypothesised for the summer.     

An Ecological-Physical Coupled Model with Ontogenetic Vertical Migration of Zooplankton in the Northwestern Pacific

A vertical one-dimensional ecosystem model with vertical migration of zooplankton was constructed and applied to Station A-7 off Sanriku district of Japan in the northwest Pacific Ocean. The model consists of an eight-compartment ecosystem model coupled with a physical model of the oceanic mixed layer. The transition of phytoplankton species responsible for the spring bloom is well reproduced by this model with vertical migration of zooplankton but is not simulated by the model without vertical migration. This new model also simulates an observed inter-annual variability of the spring bloom, with the timing and intensity of the simulated bloom in a given year depending upon the strength of mixing during the preceding winter. This revised version was published online in August 2006 with corrections to the Cover Date.     

Summer hypoxia in the northern Gulf of Mexico and its prediction from 1978 to 1995

An 18-year monitoring record (1978-1995) of dissolved oxygen within a region having hypoxia (dissolved oxygen less than 2 mgl(-1)) in the bottom layer was examined to describe seasonal and annual trends. The monitoring location was near or within a well-described summer hypoxic zone whose size has been up to 20,000 km(2). The monitoring data were used to hindcast the size of the hypoxic zone for before consistent shelfwide surveys started, and to predict it for 1989, when a complete shelfwide survey was not made. The concentration of total Kjeldahl nitrogen (TKN) in surface waters and concentration of bottom water oxygen were directly related, as anticipated if organic loading from surface to bottom was from in situ processes. The TKN data were used to develop a predictive relationship that suggested there was no substantial hypoxia before the 1970s, which was before nitrate flux from the Mississippi River to the Gulf of Mexico began to rise. The peak frequency in monthly hypoxic events is two to three months after both the spring maximum in discharge and nitrate loading of the Mississippi River. These results support the conclusion that persistent, large-sized summer hypoxia is a recently-developed phenomenon that began in the 1970s or early 1980s.     

黄河口区及其附近海域溶解氧的分布变化

本文讨论了1984年5月、8月、11月调查的黄河口区溶解氧的分布。黄河口区溶解氧的分布主要受水温的影响,在春季还受硅藻的影响,而夏季受微型浮游植物的影响。 夏季,黄河口门底层水的表观耗氧量高达2.04ml╱L。反映了黄河悬浮物所携带的有机物氧化的结果。     

Spatial and seasonal evolution of dissolved oxygen and nutrients in the Southern Levantine Basin (Eastern Mediterranean Sea): chemical characterization of the water masses and inferences on the N : P ratios

The spatial and seasonal variability of nutrients and dissolved oxygen concentrations as well as the chemical characterization of the different water masses of the Southern Levantine Basin were determined in detail. In summer, the upper 150 m of the water body was stratified and the cross basin distribution of dissolved oxygen and nutrients was fairly constant. Surficial waters were saturated with dissolved oxygen, and a shallow oxygen maximum (oversaturated) was present at about 80 m depth. Oversaturation was attributed mainly to the physical process of rapid capping and trapping of oxygen in the Atlantic water (AW) mass, with only 28% of the excess oxygen originating from biological production. Nutrient concentrations were very low and showed an increase in the intermediate levels, coupled with a decrease in oxygen. The winter cross-section distribution showed an upper mixed layer of 100 m, with dissolved oxygen and nutrient concentrations fairly constant across the basin. The concentration of nitrate was higher than in summer, while phosphate was slightly lower and silicic acid similar. In winter, the influence of the physical features (gyres) could be detected up to the surface, and in summer they were detected by the chemical properties in the 150–600 m layer. In the transition layer between the Levantine intermediate water (LIW) and the deep water (DW) (400–700 m) there was a gradual decrease in dissolved oxygen and an increase in nutrient concentrations eastwards. The DW showed no seasonal variation, only spatial variability: dissolved oxygen decreased and silicic acid increased eastwards. No differences were found in nitrate and phosphate concentrations between the DW in the western and eastern provinces, indicating the oxidation of organic matter poor in N and P.N : P ratios in the upper water masses were seasonally dependent. The largest variation was found in the Levantine surface water (LSW), from an average of 52 in winter to 5 in summer. It is hypothesized that the gradual decrease from winter to summer values was due mainly to preferential atmospheric input of N in winter and P in summer, together with biological consumption and differential regeneration of N and P. In the DW, the N : P ratios were constant throughout the year (25.2±2.7, n=567), and higher than Redfield's ratio. It was speculated that the high N : P ratio in the DW was a result of oxidation of particulate organic matter deficient in P.The winter wet atmospheric input of N provided 12% of new N to the LSW. Average new production for the Southern Levantine Basin was estimated from the new N as 4.75 g C m⁻² yr⁻¹. The dry atmospheric contribution of P was estimated to significantly increase the P pool in the LSW. Dry deposition is not evenly distributed and occurs in episodic and localized events, which may have a large effect on productivity in the short periods when deposition occurs.There have been recently reported changes in the deep thermohaline circulation of the Eastern Mediterranean, with main contribution of the Aegean Sea as a source of DW. The data presented here can serve as a reference for assessing future changes in the chemical composition of the water masses in the Southern Levantine.     

珠江河口氮和磷循环及溶解氧的数值模拟 Ⅰ.模式建立

为了研究珠江河口营养物质循环和溶解氧,建立了一个生态型水质模式.在该水质模式中营养物质以溶解无机态、碎屑有机物质、底栖物质、浮游植物和浮游动物等5种形态出现,而每种形态均分氮和磷两种形式.水质模式采用三维形式,并与斜压水动力和泥沙模式联立运行.     

Modeling of spring bloom in the western subarctic Pacific (off Japan) with observed vertical density structure

Effects of vertical stability on spring blooms of phytoplankton were investigated for the western subarctic Pacific ocean using a one-dimensional (depth) ecosystem model. In the model, vertical stability was expressed by diffusion constants calculated from observed density distribution. Dynamics of phytoplankton in blooms was calculated by the model using the vertical diffusion. Then, the calculated results were compared with the Coastal Zone Color Scanner (CZCS) data. The comparison shows that the shallow surface mixed layer causes early start days of spring blooms at inshore (northern) stations. In addition, spring blooms continue long at inshore (northern) stations since a water column has weak stability. This is because weak stability of a water column causes large nutrient supply from a deep layer and large diffusive transport of phytoplankton biomass from the subsurface maximum.     

Modelling planktonic ecosystems: parameterizing complexity

This paper explores several simplified representations of complexity or ecological ‘texture’ in models of the marine planktonic ecosystem. It is relatively straightforward to formulate more complex models to include explicitly different functional groups of phytoplankton, zooplankton and bacteria, and to include regulation by multiple nutrients such a nitrate, ammonium, silica, and iron. However, the number of parameters that must be specified from observations increases approximately as the square of the number of compartments and quickly surpasses our ability to constrain them properly from observations. Moreover, ecosystem models often become unstable for small changes in parameter values, and increasing complexity may not lead to increased stability. Here we consider alternative formulations for ecosystem models that try to represent complex interactions, such as the microbial loop, in simpler, less explicit ways. Results are presented demonstrating diagnostic methods to represent (1) multiple size classes of phytoplankton according to total biomass of a single phytoplankton compartment, and (2) partitioning a single compartment for nitrogen into nitrate and ammonium according to the origin of incoming fluxes, nitrification by bacteria, and a relative preference algorithm for uptake by phytoplankton. Ambient concentrations of ammonium are simulated with this model and evaluated against observations from Ocean Station P in the subarctic Northeast Pacific Ocean.